Amaranto

Saubhik
Das
Amaranthus:
A Promising
Crop of Future
Amaranthus: A Promising Crop of Future
Saubhik Das
Amaranthus:
A Promising Crop of
Future
Saubhik Das
Department of Botany
Taki Government College
West Bengal, India
ISBN 978-981-10-1468-0 ISBN 978-981-10-1469-7 (eBook)

DOI 10.1007/978-981-10-1469-7
Library of Congress Control Number: 2016945253
© Springer Science+Business Media Singapore 2016

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Preface
Pseudocereals are promising crops for coming decades keeping in view the
global food security. Amaranths are a leading group of plants among the
pseudocereals that have the great potentiality to prevent malnutrition espe-
cially in the low-income food-deficient countries. Though its cultivation and
use as food grains have great antiquity, but later its cultivation was ignored by
a larger section of people of the world. It lagged behind the conventional
cereal crops in spite of being nutritionally very much competitive to those. In
the last few decades, the nutritive potentialities and other unique features of
amaranths have been rediscovered in different corners of the world. Research
done so far on amaranths has surfaced its unique and unparallel nutritive
value, wide adaptability, herbicide resistance property and vast germplasm
variability. Its germplasm variability has opened up a new avenue to evolve
much improved varieties or cultivars. Little efforts have been devoted to
improve its genetic background by applying biotechnological methods. But
that yielded significant achievements. Keeping in view its tremendous poten-
tiality to become a super crop of coming decades, much more attention is to
be given to it especially when the conventional crops are overburdened with
a task to feed the huge world population. To increase the food production at a
global level and at a sustainable basis, the importance of amaranths cannot be
ignored.
In this book, attempts have been made to accumulate updated information,
significant research achievements and knowledge about amaranths also to
revive interest about amaranths and to explore them comprehensively.
The author expresses his sincere gratitude to Dr. David Brenner (curator of
amaranth, North Central Regional Plant Introduction Station, Iowa State
University, Ames, Iowa), Dr. Duilio Iamonico (Department of PDTA, Section
of Environment and Landscape, University of Rome Sapienza, Italy) and
Dr. J. C. Rana (head, Division of Germplasm Evaluation, ICAR-NBPGR,
New Delhi, India), having pioneering contribution in amaranth research, for
their help and cooperation. The author is also thankful to Dr. G. Nessom
(Flora of North America Association); Dr. K. N. Gandhi (Harvard University
Herbaria); the director of the Central National Herbarium, Howrah, Shibpur,
v
vi Preface
West Bengal, India; and the head of the Department of Vegetable Crop,
TNAU, Coimbatore for their necessary Cooperation.
The effort devoted in this book will be successful if the book gets recogni-
tion and appreciation. The author welcomes relevant comments and sugges-
tions for future improvement of this book.
Taki, West Bengal, India Saubhik Das

Contents
1 Introduction .................................................................................. 1
1.1 General ................................................................................ 1
1.2 Agricultural Development and Global Food Security ........ 1
1.3 Underutilised Crop.............................................................. 3
2 Pseudocereals: An Efficient Food Supplement .......................... 5
2.1 General ................................................................................ 5
2.2 Floristic Composition of India ............................................ 5
2.3 Pseudocereals: An Alternative Food Resource ................... 6
2.4 Nutritive Value of Pseudocereals ........................................ 7
3 Amaranths: The Crop of Great Prospect .................................. 13
3.1 General ................................................................................ 13
3.2 Grain Amaranths: A Nutritive Supplement
to Major Cereals ................................................................. 14
3.3 Different Species of Grain Amaranths ................................ 15
3.3.1 Amaranthus hypochondriacus L. ......................... 18
3.3.2 Amaranthus cruentus L. ....................................... 20
3.3.3 Amaranthus caudatus L. ...................................... 21
3.4 Potentiality of Grain Amaranth as Food ............................. 23
3.5 Vegetable Amaranths .......................................................... 25
3.6 Species of Vegetable Amaranths ......................................... 27
3.6.1 Amaranthus tricolor L. ......................................... 27
3.6.2 Amaranthus blitum L. ........................................... 29
3.6.3 Amaranthus dubius L. .......................................... 30
3.6.4 Amaranthus cruentus L. ....................................... 30
3.6.5 Amaranthus viridis L. ........................................... 32
3.6.6 Amaranthus graecizans L. .................................... 33
3.7 Nutritive Value of Vegetable Amaranths ............................. 34
3.8 Uses of Amaranths .............................................................. 35
3.8.1 Uses of Vegetable Amaranths ............................... 35
3.8.2 Uses of Grain Amaranths ..................................... 36
3.9 Culinary Properties of Amaranths ...................................... 39
3.10 Bioactive Components and Medicinal Properties ............... 39
3.11 Weed Amaranths ................................................................. 42
vii
viii Contents
4 Infrageneric Classification of Amaranths .................................. 49

4.1 General ................................................................................ 49
4.2 Synopsis of Infrageneric Classifications
of Amaranths ....................................................................... 49
4.2.1 Amaranthus Subgen. Acnida
Aellen ex K.R. Robertson..................................... 50
4.2.2 Amaranthus Subgen. Amaranthus ........................ 50
4.2.3 Amaranthus Subgen. Albersia
Grenier & Godron ................................................ 51
4.3 Infrageneric Classification After the Modification
by Mosyakin and Robertson (1996) ................................... 51
4.3.1 Amaranthus Subgen. Acnida (L.) Aellen
ex K.R. Robertson ................................................ 52
4.3.2 Amaranthus Subgen. Amaranthus ........................ 53
4.3.3 Amaranthus Subgen. Albersia .............................. 54
4.4 Provisional Key to Some Edible Species
of Amaranthus..................................................................... 55
5 Taxonomy and Phylogeny of Grain Amaranths ........................ 57
5.1 General ................................................................................ 57
5.2 Morphological Approach in Solving
Taxonomic Disputes ........................................................... 58
5.3 Biochemical and Molecular Approach
in Understanding Systematics and Taxonomy .................... 65
5.4 Cytogenetical Approach in Understanding
Species Relationship ........................................................... 75
5.5 Taxonomic Delimitation in Vegetable Amaranths .............. 80
5.6 Key to the Varieties of Amaranthus tricolor L.................... 80
5.7 Taxonomic Delimitation in Weed Amaranths ..................... 88
5.8 Different Phylogenetic Concepts on Grain Amaranths....... 88
6 Weed and Herbicide Resistance .................................................. 95
6.1 General ................................................................................ 95
6.2 Resistance to Different Types of Herbicides....................... 95
6.2.1 Resistance to Photosystem II Inhibitors ............... 96
6.2.2 Resistance to Acetolactate Synthase
(ALS) Inhibitors ................................................... 96
6.2.3 Resistance to Protoporphyrinogen
Oxidase (PPO) Inhibitors ..................................... 96
7 Distribution and Maintenance of Amaranth
Germplasm Worldwide ................................................................ 99
7.1 General ................................................................................ 99
7.2 Germplasm Collection Centres in the World ...................... 99
7.3 Genome Resource Development of Amaranthus ................ 104
8 Breeding of Amaranths................................................................ 107
8.1 General ................................................................................ 107
8.2 Breeding Behaviour in Amaranths...................................... 108
8.3 Objectives in Amaranth Breeding ....................................... 112
Contents ix
8.4 Conventional Breeding by Selection .................................. 116

8.5 Hybridisation and Heterosis in Amaranths ......................... 118
8.6 Male Sterility in Amaranth Breeding .................................. 125
8.7 Wild Gene Pool, Landraces and Heritability ...................... 126
8.8 Mutation Breeding .............................................................. 132
8.9 Polyploidy in Amaranth Breeding ...................................... 136
8.10 Genetic Engineering and Biotechnological
Approach in Genetic Improvement..................................... 137
8.11 Genetic Transformation in Breeding Amaranths ................ 138
8.12 Molecular Markers and Genome Research
for Using Genetic Resources in Plant Breeding ................. 142
8.13 Molecular Breeding in Amaranths ...................................... 143
8.14 Plant Tissue Culture Techniques in Amaranth Breeding .... 144
9 Evolution of Sexuality in Amaranths ......................................... 147
9.1 General ................................................................................ 147
9.2 Trend of Sexuality in Plants................................................ 147
9.3 Sexuality in Amaranths ....................................................... 149
10 Cultivation of Amaranths ............................................................ 153
10.1 General ................................................................................ 153
10.2 Seed Dormancy and Seed Viability of Amaranthus............ 154
10.3 Agroclimatic Condition for Cultivation of Amaranths ....... 159
10.3.1 Climate ................................................................. 159
10.3.2 Soil ....................................................................... 160
10.3.3 Temperature .......................................................... 160
10.3.4 Rainfall ................................................................. 161
10.3.5 Day Length ........................................................... 161
10.4 Cultivation Practices ........................................................... 161
10.4.1 Sowing of Seeds ................................................... 161
10.4.2 Seedling Growth and Interculture ........................ 162
10.4.3 Manuring .............................................................. 163
10.4.4 Obstacles in Productivity in Grain Amaranths ..... 163
10.4.5 Harvesting ............................................................ 164
10.4.6 Pests and Diseases ................................................ 165
11 Future Prospects in Amaranth Research ................................... 167
Appendices ............................................................................................ 173

Appendix I......................................................................................... 173
Appendix II ....................................................................................... 174
References ............................................................................................. 189

Introduction
1
1.1 General yams and five oil seed types and a few beverages
(Harlan 1975), of which only three crop types,
Global food security and struggle against malnu- viz. rice, wheat and maize, supply 60 % of food
trition are going to face a strong challenge in the requirements for the world human population.
coming decades from population outburst spe- Though over 7000 species either partly or fully
cially in the developing and Third World coun- domesticated have been used so far from time to
tries. Only a handful of crops have to feed nearly time for food production, today only 30 species
nine billion people all over the world by 2030 as are bearing the herculean responsibility of pro-
against 5.7 billion at present. The condition is viding 95 % of the huge world food requirements.
much more threatening due to the declining num- Many of the neglected and underutilised species
ber of plant species and genetic erosion of overex- are of great significance as a source of food in
ploited plant species. Scientists all over the world low-income food-deficient countries (LIFDCs).
are engaged in exploring the plant biodiversity to They are extremely important because of their
broaden the crop list. There are many neglected wide adaptability to the marginal areas and con-
and underutilised species which are capable of tribution of a significant part of the local diet with
satisfying the demand for food, nutrition and useful nutritional supplements. In comparison
energy. They can function as alternative crops, not with major crops, these neglected and underuti-
competitive to other major crops, and can be lised species require relatively low input and
adapted to fragile environment and marginal areas therefore help in sustaining agricultural produc-
needing least agronomic requirements. tion. These regional traditional crops are often
low yielding and are not competitive to conven-
tional major crops, even though many of them
1.2 Agricultural Development have the potential to become economically via-
and Global Food Security ble. Very often narrow genetic diversities in
important agronomic traits as well as lack of
Food is one of the basic needs of human being. genetic improvements prevent the development
The bulk of the food consumed in the world are of these crops. Other constrains include lack of
procured from a very limited number of crop spe- adequate knowledge on the taxonomic delimita-
cies. Today by and large, 25–30 food-yielding tion, the genetics of agronomic and quality traits
species supply food to mankind, viz. wheat, rice, and reproductive biology.
maize, barley, oats, sorghum, millets, soybean, Many plant species with significant food and/
bean, pea, chickpea, peanut, banana, citrus, or industrial value are yet to be utilised properly
tomato, sugarcane, cassava, potato, sweet potato, due to lack of appropriate and adequate
© Springer Science+Business Media Singapore 2016 1

S. Das, Amaranthus: A Promising Crop of Future, DOI 10.1007/978-981-10-1469-7_1
2 1 Introduction
programmes for their evaluation and develop- reports on underutilised tropical plants with
ment and remained underutilised. Some of them promising economic value (1975). In recent
have been so neglected and erosion of their gene times NAS issued further reports (e.g. NAS 1989,
pool is so severe that they are often considered as 1996) and FAO issued many (FAO 1988;
‘lost crops’. A vast majority of these plants have Hernandez Bermejo and Leon 1994).
the capability to meet the increasing demand for An autonomous international scientific organ-
food and nutrition, healthcare, medicine and isation, International Plant Genetic Resources
industrial needs. Many of these species are Institute (IPGRI), was established by the
involved in traditional localised farming espe- Consultative Group on International Agricultural
cially in marginal remote areas, and quite often Research (CGIAR) in 1994, and it is situated in
these crops act as life-savers for millions of poor Rome, Italy, at the Food and Agriculture
people in the regions where food security and Organization of the United Nations. The prime
malnutrition are a problem. The rural community role of IPGRI is to monitor research activity and
knows very well the cultivation practices of the to promote and coordinate an international net-
underutilised crops and prepare food from them work of plant genetic resource, germplasm man-
and use them in their daily life, for health care, agement conservation, evaluation, documentation
shelter, forage and fuel particularly during and utilisation of useful plant germplasm glob-
drought, famine and the dry seasons (Campbell ally and also the collection and exchange of plant
1987). These crops include cereal, pseudocere- genetic resources. The functioning of IPGRI and
als, fruits and nuts, pulses, vegetables, root and other such institutes is monitored by CGIAR
tubers, oilseeds and other industrial, forage and which was established in 1972. The joint efforts
fodder species. of Food and Agriculture Organization (FAO), the
Global food security and economic growth is World Bank and United Nations Development
certainly going to face a stiff challenge in the Programme (UNDP) resulted in the creation of
coming few decades due to population outburst CGIAR to establish international research insti-
especially in the developing and Third World tutes and subsequently to look after their
countries. In an estimate it appears that nearly 1.2 progress.
billion people in the world are not lucky enough Global food security and economic growth is
to get adequate food to meet their daily nutri- now being threatened by declining number of
tional requirement and a further 2 billion people plant species. This decline has placed the future
are deficient in one or more micronutrients supply of food and rural income at risk. IPGRI
(Azam-Ali and Battcock 2002). The scale of food has succeeded in promoting greater awareness
shortage has been estimated by the Food and regarding the important role that minor crops can
Agriculture Organization; Jacques Diouf, its sec- play in securing and safeguarding the livelihood
retary general, says that 800 million people in the of people all over the world. Ethnobotanical sur-
world (20 % of the population of developing veys confirm the presence of hundred of such
countries and up to 37 % of sub-Saharan Africa) crops in many countries and different remote cor-
suffer from shortage of food, and 192 million ners of the globe that represent a plentiful trea-
children have chronic malnutrition. By 2030 the sure of agro-biodiversity. Such underutilised
humanity has to perform a task of feeding 9 bil- mainly ethnic crops can play a vital role to
lion people as against 5.7 billion at present. improve income, food security and nutrition. The
Nearly 30 % of the total population of Africa is development of agriculture and food security
suffering from chronic hunger and malnutrition depends partly on our ability to extend the agri-
which compelled the stakeholders in and outside cultural species range in an effective and sustain-
the region to look into possible measures to solve able manner. This requires finding of ways and
food crisis. The gradually emerging threat of means to protect and enhance cultivation of the
worldwide food shortage compelled the US locally important species so that they can be
National Academy of Science (NAS) to issue employed more widely in agriculture and
1.3 Underutilised Crop 3
environment management and finding of ways to mented and neglected for research and conserva-
explore the use of local crops in order to tap the tion. The benefits and features of these plants are
hidden potential contained in them. Today global as follows:
food security has become increasingly dependent
on few conventional crops causing their over- 1. They are of local importance in consumption
exploitation. Even if mankind has used more than and production system.
10,000 edible species since the prehistoric period, 2. They can grow well in fragile environment
today only 150 plant species are commercialised and help to stabilise the agroecosystem par-
globally in a significant scale, 12 of which are ticularly in arid, semiarid, mountainous
supposed to provide approximately 80 % dietary locality and tropical forest. They are virtu-
energy from plants and only four plant types, viz. ally adapted to any environmental condition,
rice, wheat, maize and potato are supposed to soil types and specialised agro-ecological
supply over 60 % of the global requirements for niche. They can tolerate difficult conditions
protein and calories. Moreover the gradual and environment stresses.
decrease of the crop varieties are increasingly 3. Regarding achievement of success in various
threatening the future supply of food and rural social objectives like poverty elimination
income and this has compelled research organ- and generation of employment and income
isations and scientists worldwide to retrieve, opportunities in both rural and urban envi-
research and disseminate the knowledge regard- ronment, they have a great role to play.
ing production and utilization of neglected, 4. For the sustenance of livelihood through
underutilised new plant species or the so-called safeguarding food security and widening
alternative crop (FAO 1996a, b, c, 2005). food basket, their role cannot be ignored.
5. They are the rich source of nutrients and can
add nutrients to the diet. These nutrient-
1.3 Underutilised Crop enriched foods are convenient for low-
income people group.
Underutilised crops can be defined as a class of 6. They provide a wide range of crop species to
crop that once grown more rapidly and intensely meet new market demands.
but lagged behind the conventional major crops 7. They have attracted attention of the National
in terms of cultivation and use for variety of agro- Agriculture and Biodiversity Corporation
nomic, genetic, economic and cultural reasons. policies, researches and development.
They are not properly utilised though not com- 8. They are cultivated and utilised based on
petitive with other crops. Neglected crops appear indigenous knowledge.
to be synonymous with underutilised crops. But 9. They are scarcely represented in ‘ex situ’
neglected crops are those crops which are grown collection.
mainly in their centre of origin by traditional 10. They are represented by ecotypes/or
farmers for local communities and are ill docu- landraces.
Pseudocereals: An Efficient Food
Supplement 2
2.1 General Ghats, out of the 25 biodiversity hotspots (Fig.

2.1) identified in the world (Myers 1990). These
Pseudocereals are defined as fruits or seeds of two hotspots show most of the plant diversities in
non-grass species that are consumed in very simi- India. As far as species diversity is concerned,
lar way as cereals. Pseudocereals are effective approximately 45,000 plant species are recorded
supplements to conventional cereals. The protein in India (Khoshoo 1995; Sharma et al. 1997). In
contents of pseudocereals like quinoa, amaranths India the angiosperm flora is represented by
and buckwheat are much higher than cereals, and approximately 17,500 species of which 5725 spe-
the quality of proteins is also much improved cies belong to endemic category. In another esti-
containing higher amount of lysine which is lim- mation about 28 % of the total Indian flora and
iting in cereals. From the angle of digestibility, about 33 % of angiosperm flora occurring in India
bioavailability, available lysine and net protein are endemic (Nayar 1996). In a rough estimate
utilisation, pseudocereal proteins are definitely about 10 % of flowering plant species in India are
better when compared to cereals. The nutritive threatened and 34 plant species have been
value of pseudocereals is very much competitive declared to be extinct (Nayar and Sastry
to conventional crop, in most cases even better. 1987–1990).
India is one of the versatile centres of diver-
sity of cultivated plants. This region is character-
2.2 Floristic Composition ised with enormous landrace diversity which is
of India gifted with useful gene pools for tolerance to
physiological and ecological stresses and resis-
India is the seventh largest and tenth industri- tance to disease, pest, etc. and good quality traits.
alised country in the world with a geographical Wild relatives and progenitors of cultivated
area of about 3287,263 sq km situated between plants are of particular importance. About 326 of
804′ N to 3796′ N latitude and 6807’ E to 97025′ such plants have been identified in India. Nearly
E longitude. The Indian subcontinent is divided 1000 wild plant species which are edible have
into four climatological zones – equatorial, tropi- been widely exploited by native tribal people
cal, subtropical and warm temperate according to (Arora and Nayar 1984; Arora 1985, 1987; Arora
longitudinal variation. et al. 1991; Pandey 1998; Malik and Singh 2006).
India represents about 11 % of the world’s All India Coordinated Research Project
flora in spite of having only just about 2.4 % of (AICRP) on underutilised crop (UUC) was initi-
the total landmass. India has two biodiversity ated in 1982 with a headquarter at the National
hotspots, namely, Eastern Himalaya and Western Bureau of Plant Genetic Resources (NBPGR),

6 2 Pseudocereals: An Efficient Food Supplement
Fig. 2.1 Biodiversity hotspots identified worldwide with two biodiversity hotspot in India
New Delhi, with 15 main centres and 10 co- year 2020, the anticipated food grain demand
operating centres in different agricultural zones would escalate up to nearly 250 million tonnes,
of the country. which means an additional 72 million tonnes of
AICRP has identified the following categories food grains are to be procured.
of underutilised crops which are to be considered
for utilisation:
2.3 Pseudocereals:
1. Pseudocereals – Grain amaranths, quinoa, An Alternative Food
buckwheat, Job’s tear, etc. Resource
2. Food legumes and pulses – Rice beans,
winged beans, faba beans, etc. Cereals belong to the grass family (Poaceae) and
3. Oilseed – Perilla, paradise tree, etc. cultivated for their starchy edible seeds.
4. Vegetables – Kankora, winged bean, etc. Pseudocereals are also grown for the same pur-
5. Fodder plant – Amaranth, salt bush, etc. pose, but they do not belong to the grass family.
6. Energy, hydrocarbon and industrial plants – According to the definition proposed by Shewry
Jatropha, guayule, jojobe, etc. (2002), pseudocereals are dicotyledonous species
which are not closely related to each other or to
The bulk of India’s cereal supply (nearly the monocotyledonous true cereals. These are
75 %) is provided by three major cereal crops like grouped artificially keeping in view the use only
wheat, rice and maize, while minor cereals like rather than the biology of plants and rapidly gain-
Avena sativa, Eleusine coracana, Echinochloa ing popularity especially in the Third World
sp., Hordeum vulgare, Panicum miliaceum, countries. Cereals and pseudocereals are the pri-
Pennisetum sp., Secale cereale, Setaria italica mary suppliers of carbohydrates for the world’s
and Sorghum bicolor and pseudocereals like human population. The human population con-
Amaranthus, Chenopodium and Fagopyrum sume nearly half of the annual cereal production.
esculentum provide only the remaining 25 %. The primary cereals comprise of rice, wheat,
More than 1200 cultivated and wild herbaceous corn, sorghum, millet, oats, barley and triticale.
plants are consumed as leafy vegetable. By the The term millet refers to the small-seeded grain
2.4 Nutritive Value of Pseudocereals 7
obtained from few unrelated species. Beside con- view point, the values for pseudocereal proteins
ventional cereals human food resources also are definitely higher in comparison to cereals and
include a wide variety of other plants like minor are close to those of casein. The protein composi-
cereals and pseudocereals which are of minor tion of pseudocereals is typical for dicotyledons
significance not cultivated extensively but not at in having 2S albumins, 11S and 7S globulins and
all negligible. Such plants have some obvious therefore similar to protein of legumes, crucifers
advantages: firstly they are adapted to drought and composites (Marcone 1999). Due to lower
and stress condition, secondly they have the abil- prolamine content, the pseudocereal proteins are
ity to grow on poor soils of marginal areas unfit suitable for the person suffering from celiac dis-
for other major crops and thirdly local rural peo- ease. The fat content of pseudocereals is also
ple know well how to cultivate and use them. higher compared to most cereal species and that
Such plants provide better nutrition than the fat is characterised with a high content of unsatu-
major crops. The black fonio (Digitaria iburua) rated fatty acids (e.g. linolenic acid). The mineral
in Nigeria, white fonio (Digitaria exilis) in the content in amaranth and quinoa is about twice as
rest of the tropical Africa, Brachiaria deflexa var. high as in other cereals.
sativa and B. ramose in certain areas of Africa The genus Amaranthus comprises many edi-
and the staple cereal Teff grass (Eragrostis abys- ble species besides many weedy members which
sinica) in Ethiopia have gained great importance grow worldwide but prefer the tropical climate.
and popularity and contributed a lot towards food Amaranths are one of the earliest known crop
security specially in Africa. Few dicotyledonous plants extensively cultivated and utilised by the
members like Amaranthus caudatus, Amaranthus Aztec people, who considered it as a ‘superfood’.
cruentus and Amaranthus hypochondriacus of Its exceptionally high nutritive value was
Amaranthaceae, Chenopodium quinoa (quinoa) explored long before. In Africa the leaves of the
of Chenopodiaceae and Fagopyrum esculentum vegetable amaranths are consumed like spinach.
and F. tartaricum (buckwheat) of Polygonaceae The presence of amaranth seeds in the prehistoric
produce starch-rich seeds that can be consumed period was evidenced through archaeological
as cereals, known as pseudocereals (Fig. 2.2). excavation at a cave in Tehuacan, Puebla, Mexico.
The seeds of Amaranthus cruentus collected
from Puebla, Mexico, dates back nearly 6000
2.4 Nutritive Value years. Records from Aztec civilisation indicated
of Pseudocereals the cultivation, collection and use of grain ama-
ranths and also indicated collection of grains in
The nutritive value of pseudocereals is much huge quantities along with corn and beans as
superior than cereals. As far as protein content annual tribute to ruling elite class. The people of
and protein qualities are concerned, the pseudo- Aztec, Mayan and Inca civilisation (Fig. 2.3)
cereals are much better than the cereal species. used to grow different species of Amaranthus and
Amino acids like tryptophan, lysine particularly consume both leafy vegetable and cereal grains.
lysine which is limiting in cereals is found to be In Central America during the Mayan and Aztec
present in high amount in pseudocereals. The period, people used amaranths as principal food.
amino acids arginine and histidine both proved After the colonisation of America, the use of
essential for infant and child health present in amaranths significantly faded out and its utilisa-
significant amount in seeds, which projected tion drastically decreased. This crop is now culti-
amaranth and quinoa as an appropriate food sup- vated only in some pockets of the world as merely
plement for child nutrition. Net protein utilisa- traditional regional crop, though global interest
tion (NPU) or protein efficiency ratio (PER), in amaranths has been rejuvenated in the last cou-
protein digestibility or bioavailability of protein ple of decades.
and available lysine are some of the prime indica- Enzyme inhibitors and allergens are known to
tors of nutritional quality of a protein. From this be present in cereals. Protein isolated from wheat,
Fig. 2.2 Different types of pseudocereals (a) quinoa, (b) buckwheat, (c) Amaranthus
Fig. 2.2 (continued)
Fig. 2.3 Use of grain amaranths in ancient civilisations
rice, maize and barley may cause allergic reac- Pseudocereal quinoa (Chenopodium quinoa)
tion, a gliadin fraction isolated from wheat causes is considered as the mother of all grains by the
celiac disease. But these components are not Incas and originated in the Andean region of
available in pseudocereals and legumes such as South America. It is now produced in Bolivia,
soybean and amaranths (Kuhn et al. 1996). Peru and Ecuador. Unlike many other grains, it
Furthermore, pseudocereals contain dietary fibre has a significant amount of protein and a well-
in high proportion, which improves lipid metabo- balanced amino acid profile including high con-
lism (Gorinstein et al. 1998; Oleszek et al. 1999). centration of lysine (generally low in most other
The nutritional value of pseudocereals is mainly cereals). Its recognition as grain of high nutritive
due to its protein fraction. Natural vegetable pro- value has not been changed much since the Inca
teins of leafy amaranths are useful for its high period, and today it is sometimes considered as
biocompatibility, nutritional value and low cost. ‘supergrain’. It resembles amaranths in having a
All the pseudocereals have major group of 11S relatively high protein content of good nutritive
globulin storage protein while smaller amounts value and great tolerance to arid condition
of 2S albumin and 7-8S globulins are found in (Taylor and Parker 2002). Possible utilisation of
buckwheat and amaranths. quinoa as food was studied by several research-
ers emphasising its use in the production of mins in cereals and globulins in pseudocereals
bread and cakes (Been and Fellers 1982; Lorenz were investigated by several workers (Konishi
and Coulter 1991; Chauhan et al. 1992) and and Yoshimoto 1989; Ker et al. 1993; Gorinstein
pasta (Caperuto et al. 2001). Amaranths and qui- et al. 2002). A correlation between nutritive
noa both produce significant amount of edible value of protein and amino acid composition
grain, especially amaranth which is considered revealed a close identity between soybean and
as the grain of the twenty-first century (Oleszek amaranth. The suitability of amaranths as a
et al. 1999; Vetter 1994; Zheleznov et al. 1997; nutritive substitute for cereals is well substanti-
Gorinstein et al. 1996). Both amaranths and qui- ated by its rich protein and amino acid
noa are very rich source of minerals and vita- composition.
mins than most of the cereals (Vetter 1994; Buckwheat, not a type of wheat but another
Gorinstein et al. 1996). Qualitatively and quanti- pseudocereal, has also gained attention as a
tatively the protein of grain amaranths is more prospective crop. It also contains protein of
superior than cereals and legumes (Singhal and high nutritional value and the protein is rela-
Kulkarni 1988). The protein content of tively rich in lysine and other essential amino
Amaranthus is about 16 %, and proteins are also acids. It is enriched with high amount of pheno-
enriched with high content of arginine, lysine, lics, iron, chromium, calcium, magnesium,
tryptophan and sulphur containing amino acids selenium and polyunsaturated fatty acids.
(Oleszek et al. 1999; Vetter 1994). The lysine Buckwheat originated in the Middle Asia and
content of amaranth is twice that of wheat and was transported to Central and Eastern Europe
thrice that of maize, and the nutritive value is by the nomadic people. Within the thirteenth
about 75 compared to 44, 57 and 62 for maize, century, buckwheat gained some importance in
wheat and barley, respectively (Zheleznov et al. Germany, Austria and Italy, which was eventu-
1997). It has also been observed that the globulin ally lost due to the cultivation of other cereals.
fraction of oat and amaranth is highly homoge- Today, due to the demand of gluten-free diet,
neous and shares similarity with the legume 11S the interest in buckwheat has been revived
storage protein (Bressani and Garcia-Vela 1990; (Tables 2.1a and 2.1b).
Segura-Nieto et al. 1994; Gorinstein et al. 1999). According to the recent APG classification,
Main storage protein fractions, such as prola- both the genus Amaranthus and Chenopodium
Table 2.1a Percentage based dry weight of chemical components of amaranth, quinoa and buckwheat
Buckwheat
Amaranth (Amaranthus Quinoa (Chenpodium (Fagopyrum Wheat (Triticum
Components cruentus L.) quinoa L.) esculentum) aestivum L.)
Protein 15.2 13.3 10.9 11.7
Fat 8.0 7.5 2.7 2.0
Starch 67.3 69.0 67.2 61.0
Ash 3.2 2.6 1.59 1.8
Souci et al. (2000)
Table 2.1b Comparative account of nutritive value of grain amaranths and other cereals
Food
Crops Protein Fat Carbohydrate Calcium Iron Phosphorus energy
Amaranth 16.0 3.1 60.0 0.49 17.5 0.60 391
Rye 12.1 1.7 73.4 0.38 10.5 0.37 334
Buckwheat 11.7 2.4 72.9 0.12 15.5 0.28 335
Chenopod 12.0 5.0 68.0 0.20 12.6 0.50 342
Wheat 13.3 2.0 71.0 0.41 10.5 0.37 333
Maize 9.2 3.9 73.7 0.20 3.5 0.25 355
Rice 7.0 1.0 78.0 0.20 3.5 0.18 345
Souci et al. (2000)
are now included in the family Amaranthaceae Polygonales and Caryophyllales are closely
under the order Caryophyllales, whereas buck- linked, though Drzewiecki et al. (2003) found
wheat (Fagopyrum sp.) is included in the fam- significant genetic divergence between
ily Polygonaceae under the order Polygonales. Caryophyllales and Polygonales.
Amaranths: The Crop of Great
Prospect 3
3.1 General and A. blitum. The tender plant of grain species

A. cruentus is also consumed as leafy vegetable.
Amaranthus is a widely distributed herbaceous Green amaranths are rich source of lysine-rich
genus of herbs comprising approximately 70 spe- protein, β-carotene, various vitamins, minerals
cies collectively called amaranths or pigweeds. and dietary fibres. Anti-nutrients like nitrates and
Most of them are annual weeds; few are known oxalates are present in small amount that does not
as nutritive vegetable and graceful ornamentals, cause any nutritional problem under normal con-
while protein-rich grains of few species are condition of consumption. The underutilised ama-
sumed as pseudocereals known as grain amaranths are projected as crop of the twenty-first
ranths. The species number is still tentative due century because of their health benefits and nutri-
to misapplication of names and synonyms applied tive value. Vegetable amaranths provide energy,
to the misidentified names. Three species of help to maintain proper mineral balance, reduce
Amaranthus are familiar for grain production – bad cholesterol, improve eyesight and prevent
A. hypochondriacus, A. caudatus and A. cruen- anaemia. Compared to wheat, rye, rice and oat,
tus. According to one school of thought, all the grain amaranths are gluten free and contain 30 %
grain amaranths are of the New World origin, but more protein with complete set of amino acid.
other school of thoughts suggested that grain Amaranth grain may be processed in various
amaranths might have been cultivated in South forms like flaked, popped, extruded and ground
Asia from prehistoric period and probably have into flour and also can be used as a substitute in
domesticated there. A comparative study of grain porridge, stirred into soup. Grain amaranths have
amaranths in India and Central America indi- several health benefits like lowering of plasma
cated close similarity in species distribution, evo- cholesterol level, protection of heart, stimulation
lution, variety pattern and cultivation practices. of immune system, anticancer activity, control of
Seeds of grain amaranths are very rich in crude blood sugar level, improved condition of hyper-
protein with lysine and threonine. Average pro- tension and anaemia, anti-allergic and antioxi-
tein score is either equal or much greater than dant activity, etc., due to the presence of some
rice, wheat, soybean and maize. Seed oil contains bioactive components. Like other vegetables,
squalene, trypsin inhibitor, tocotrienols, tannins, green amaranths go through cooking such as fry-
etc. Vegetable amaranths are the most popular ing, simmering, boiling, steaming and blanching
vegetable crops in tropics especially in the tropi- before consumption. Cooking has a significant
cal humid climate of Africa and Asia. Several effect on bioactive components either positive or
species are known as vegetable amaranths of negative depending on the process. Amaranths
which two are most popularly grown, A. tricolor are also known for its weedy members known as

14 3 Amaranths: The Crop of Great Prospect
pigweed. Approximately ten Amaranthus species Amaranthus are available in Asian region. The
are regarded as weeds. They are also capable of antiquity of amaranths in Indian subcontinents
competing with other crop plants, express highly was evidenced by fossil record of Amaranthus
flexible adaptability to environmental changes pollen documented in several excavations in
and various ecoclimates and ensure their exis- India that dates back to the Holocene and late
tence producing a large number of seeds. Palaeocene periods.
The species of Amaranthus are mostly annual
weeds; few are utilised as vegetables and orna-
3.2 Grain Amaranths: A Nutritive mentals. Protein-rich grains or seeds of few spe-
Supplement to Major Cereals cies ( A. caudatus L., Amaranthus hypochondriacus
L. and A. cruentus L.) are consumed as pseudo-
Amaranthus L. is a cosmopolitan genus of herbs cereals; they are called grain amaranths. All the
of the family Amaranthaceae collectively known species fall roughly under any of the four catego-
as amaranths or pigweed. It includes about 70 ries – grain, vegetable, ornamental and weed.
species (Costea et al. 2001a, b; Iamonico 2012) Amaranth is considered as one of the few multi-
and 40 of which are considered native to America. purpose crops which supply seeds in huge quanti-
In another estimation the genus Amaranthus is ties that can be used as pseudocereals, as tasty
reported to include 87 species, of which 14 found leafy vegetable of higher nutritional quality and
in Australia, 17 found in Europe and 56 available also as food and animal feed. Some member has
in America (Jacobsen and Mujica 2001; Mujica attractive inflorescence of various colours that
and Jacobsen 2003). Among the American spe- made them valued as ornamental also. Although
cies, ten are dioecious and the remaining 46 are the crop was one of the sources of staple food in
monoecious. Dioecious species are confined in the pre-colonised South American civilisation, the
North America. Distribution of monoecious spe- cultivation and knowledge fell into oblivion, and
cies in American subcontinent is scattered: 13 thus nowadays it could be considered as a new,
species are endemic to North America and forgotten, neglected but prospective and alterna-
Mexico; 17 species are restricted to Antilles, tive crop of immense nutritive potential. The grain
Central America and South America; and the amaranths growing in the Himalayan region show
remaining 16 species are quite common to wide genetic diversity and morphological vari-
Americas. It is very difficult to decide which are ability, which surely substantiate the speculation
distinct species and which are synonyms applied about probable spread of the crop in India from
to the misidentified specimen. Same new species that region in the eighth century. Based on direct
of considerable phenotypic differences from the and indirect evidences regarding antiquity of the
existing species may turn out to be ecotypes or crop in India, Joshi and Rana (1991) suggested
natural hybrids of complete sterility or marginal that the grain amaranths were prevalent in South
fertility, but such plants may be included as new Asia from the time immemorial (Joshi and Rana
species (Chan 1996). There is no general agree- 1991). In Asia one can find a great ill-defined
ment on the taxonomy of amaranths and species grain amaranth region extending all the way from
number. Brenner et al. (2000) and Robertson Manchuria through interior China and Himalaya
(1981) mentioned about 60 species, while USDA, to Afghanistan and Persia. Wide scattering of the
ARS included 86 species under the genus crop throughout Asia and popularity and tradi-
Amaranthus. Behera et al. (1974) and Brenan and tional use of the grains in marginal areas are sup-
Townsend (1980) included 50 species under portive to its antiquity in the area.
Amaranthus, while Sauer (1993) considered 75 The word ‘amaranth’ originated from the Greek
species under the genus Amaranthus. Over 400 word ‘amarantos’ which means ‘unwithering’.
varieties are included within Amaranthus found The term was applied to amaranth to signify its
throughout the tropical and temperate regions of hearty characteristics symbolising immortality.
the globe. Approximately 25 species of Grain amaranths are grown all over India from
3.3 Different Species of Grain Amaranths 15
high slopes of Himalaya to coastlines. A large (1967) recognised three principal species of
number of varieties are grown throughout the genus Amaranthus for grain production:
country, but Himalayan region is considered to
represent the centre for diversity of grain ama- • A. hypochondriacus L. (sin. A. leucocarpus
ranths in India because varietal and morphotypic S. Watts, A. frumentaceous) - Prince’s feather
variability which is found in Himalaya is unique (Fig. 3.1);
and unparallel not comparable to other parts of the • A. cruentus L., sin. A. paniculatus L. - bush
country. The grain amaranths also grow as native greens, red amaranth (Fig. 3.2)
species in the Andean region of South America, • A. caudatus L. of two subspecies: subsp. cau-
including Argentina, Peru and Bolivia. Today in datus; and subsp. mantegazzianus Passerini
the Andes region, it is widely cultivated. This crop syn.: A. edulis Spagazzini, named love-lies-
has been termed as ‘Incan Wheat’ because it was a bleeding and Inca wheat, respectively (Fig. 3.3)
principal food for the Incas. Today the grains are Within each grain species, there are several
often familiar by the name kiwicha. grain types or races defined by their common
branching pattern, height, inflorescence size and
form, days to maturity, seed size and colour and
3.3 Different Species of Grain other morphological characteristics (Kauffman
Amaranths 1992; Espitia-Rangel 1994; Brenner et al. 2000).
Though cultivation of grain amaranths has a great
Grain amaranths are appropriate to be considered antiquity, today it is cultivated in small scale in
as an alternative and prospective crop in temper- some discrete pockets of the world like in parts of
ate climate. Due to globalisation and industriali- Mexico, Guatemala, Peru, India and Nepal. It has
sation of agriculture, the food supply gradually a bright prospect for further cultivation in the
became dependent on only a handful of plant spe- USA and tropical countries. The grain amaranths
cies. Grain amaranths may be utilised as an alter- have been described as native crops of low profile
native source of food. Sauer’s taxonomic key that could be cultivated easily by native people in
Fig. 3.1 Morphology of Amaranthus hypochondriacus L.

Fig. 3.2 Morphology of Amaranthus cruentus showing different morphotypes with massive colourful terminal
inflorescence
rural areas for several reasons: (i) They can be Although grain Amaranthus is a crop of the
harvested easily, (ii) they grow very fast and pro- Americas, Amaranthus hypochondriacus migrated
duce a lot of seeds which are used as pseudocere- to Asia. During the last century, increasing popu-
als (iii) tolerant to arid environment and suited larity in its cultivation has been observed among
for subtropical and tropical region and (iv) seeds the hill tribal areas in India, Pakistan, Nepal, Tibet
contain large amount of protein and essential and China. In India grain amaranths is an impor-
amino acids. tant crop of the Himalayan region. Besides upland
Fig. 3.3 Morphology of Amaranthus caudatus with characteristic drooping inflorescence
Asian region in the Himalayan belt, grain ama- opined that grain amaranths have been cultivated
ranths are also gaining popularity among the peo- in South Asia from time immemorial and proba-
ple in northwestern plains of India as well as in the bly originated there. This concept has been con-
hilly regions of Southern India with a common solidated by the fact that the grain amaranths
name Rajgira (‘king seed’), Ramdana (‘seed sent growing in the Old World are indistinguishable
by God’) and Keerai. It often occupies more than both taxonomically and morphologically from
half of the non-irrigated crop land of the higher few specific members of grain amaranths culti-
elevation in the hills of North-West. It is now a vated in the New World. The similarity was
prominent crop in a few local areas in India where observed both at species level and subspecific
a kind of bread made from its seed flour is a popu- level. It is speculated that the Old World speci-
lar food. Popped grain are used to make confec- mens represent nothing but a small sample of
tioners and also taken soaked in milk. diversity available in the American grain ama-
According to Sauer (1950), there are four ranths. Ancient literature of the Old World which
major grain amaranth regions in the Americas, has not been searched for evidence of grain ama-
each having its own particular cultivated species: ranths indicates short gaps in the history of grain
the Mexican centre which is dominated by A. amaranths in the Old World. The first record of A.
hypochondriacus, Guatemala with its main crop hypochondriacus in Asia was surfaced through
of A. cruentus, the Andes with A. caudatus and Linnaeus’ description of an Indian form called A.
Argentina with A. edulis. In subsequent analysis flavus. The grain amaranths in Asia were first
of the genus, it was concluded that A. edulis was studied by Francis Buchanan Hamilton. In the
a variety of A. caudatus, and thus, the latter two early nineteenth century, he found grain ama-
centres, the Andes and Argentina, could be con- ranths in South India and in the Himalayan
sidered one large region where A. caudatus domi- region. He named the white-seeded grain of
nates. Sauer concluded that all the grain South India A. frumentaceus. A later observer,
amaranths are of the New World origin. He Wight (1843), wrote that seeds of A. frumenta-
acknowledged the observations of other investi- ceus were the principal food of the wild inhabit-
gators like deCandolle (1883), Hooker (1885), ants of the hill areas in Coimbatore, Salem and
Ames (1939), Vavilov (1949), Darlington and Madurai (South India). Sauer noted that the name
Janaki-Ammal (1945) and Merrill (1950) who of the crop recorded by Buchanan Hamilton was
Kiery which is currently grown under the name Table 3.1 Comparative study of grain amaranths in India
and in Central America
A. hypochondriacus. Amaranths are an important
field crop in the foot hills and mountains of the (A) Name of the species Similarity observed
entire Himalayan region. The extensive research 1. Amaranthus Distribution from lower to
on the grain amaranths of the Himalayan region hypochondriacus higher altitude in both
regions
was carried out by Joshi (1978–1988). They are
2. A. cruentus Distributed in lower to
cultivated as a minor crop in Mexico, Guatemala, middle altitude in both the
Peru, Bolivia, Ecuador, Argentina, Sierra Leone, regions comparatively less
Nigeria, Zambia, Kenya and Egypt, Afghanistan, adaptable to higher hills
than A. hypochondriacus
Persia, China, Manchuria, Nepal, Bhutan and
and A. cruentus
India. Its maximum cultivation and distribution at
3. A. caudatus Confined to higher
present is observed in the Himalayas. elevation in both India and
On the basis of his detailed study of the Central America, indicates
Himalayan grain amaranths, Joshi concludes that specific adaptations
the cultivation of grain amaranths in India is very 4. A new genotype Found in both the regions in
close to A. edulis the plant population of
ancient. A comparison of collections and evalua- amaranth field
tion studies of grain amaranths in India by Joshi (B) Crop characteristics
(1981a, b) and in Central America by Hauptli 1. Seed colour Seven seed colour types
et al. (1979) indicated close similarity in species were collected by Hauptli
distribution, evolution, variation pattern and cul- (1979) in Central America
tivation practices in two widely separated geo- and Joshi (1981a, b) in
Himalayas
graphical regions of the world (Table3.1). The
2. Inflorescence types Similar variation in
study further suggests that either of the grain inflorescence pattern and
amaranths, i.e. A. hypochondriacus and A. cau- other plant parts in both
datus, have independently originated or were regions
introduced in the Old World before 1500 3. Harvestability Presence of circumscissile
AD. Fossil records of pollens have been found in utricle
4. Cultivation process Mixed cropping with maize
several digs in India from Holocene and Late
and French beans,
Pleistocene periods (Possehlo 1993). indicating close ecological
relationship
Joshi and Rana (1991)
3.3.1 Amaranthus hypochondriacus L.
Among the grain amaranths, it is the most robust others are tall and unbranched. The species is
and the highest-yielding grain types. It was suitable for the tropical areas, high altitudes and
probably domesticated in Central Mexico and dry conditions. It is of greatest potentiality to be
further north much after the domestication of used as food ingredient due to its excellent seed
Amaranthus cruentus and reached the USA in quality. The grain mills and pops well that tastes
prehistoric time, but later became extinct there. and smells pleasant. The grain reached Europe
The pale seeds of A. hypochondriacus of 1500 with the help of the Spanish who took seeds to
years back were discovered in the Tehuacan Europe, and by the sixteenth and seventeenth
caves. Though it originated in Central America, century, the plants spread through European gar-
today it is mostly cultivated in India, particularly dens as graceful ornamentals. Around 1700, it
in the Garhwal and Kumaon regions of Uttar reached Central Europe and Russia as a minor
Pradesh and in the Sutlej Valley of Himachal grain crop and consumed as mush and groats. By
Pradesh. It shows diversity in habits, and some the early nineteenth century, it was taken to
types of Amaranthus hypochondriacus are bushy; Africa and Asia, and at present it is cultivated as
grain crop in widely scattered regions of the or spikes, dark red or deep beet-red, purple, less
world like the mountains of Ethiopia, the hills of commonly yellowish or greenish, leafless at least
South India, the Nepal Himalaya and the plains in distal part, prickly and stiff. Bracts are lanceo-
of Mongolia (Fig. 3.4). late to linear-subulate, spinescent, two times as
Amaranthus hypochondriacus is known vari- long as tepals and style branches, texture rigid.
ously as Prince’s feather, Prince-of-Wales-feather Pistillate flowers have five tepals and tepals are
and Prince’s feather amaranth. It is an annual, slightly recurved, proximal ones lanceolate, distal
erect, herbaceous plant, glabrous or moderately ones narrowly ovate-elliptic to elliptic, unequal to
pubescent in distal portion, at maturity often occasionally subequal, inner tepals shorter than
becoming glabrescent, attaining a height up to outer tepals, apex acute to acuminate; tepals are
3 m. Stems are green or reddish purple in colour, comparatively larger than other grain species.
branched, mainly in inflorescences, to nearly sim- Style branches are spreading recurved, meeting in
ple proximally, coarse. Leaves are simple, petio- a sharp cleft at thick bases, stigmas 3. Staminate
late, petioles of the distal leaves equaling or flowers are clustered at tips of inflorescence
slightly shorter than blade, become longer proxi- branches, tepals 5, stamens 5. Fruits are com-
mally. Leaf blades are rhombic-ovate to broadly pressed, elongate-ovate to ovoid, circumscissile
lanceolate or elliptic or ovate-oblong, 4–14 × 2–8 utricles with broad cap, (1.5 -) 2–3 mm long,
cm, base cuneate to broadly cuneate, narrowly equaling tepals or nearly so, smooth or lid slightly
cuneate in distal leaves, margins entire, apex acute rugose or minutely verrucose, dehiscence regular.
or acuminate or indistinctly emarginate, mucro- Seeds are white, pinkish white or black to dark
nulate. Inflorescences are enormous, robust, pre- reddish brown, subglobose to lenticular, 1–1.4 mm
dominantly terminal or lateral thick erect panicle in diameter, smooth and shiny (Fig. 3.5).
Fig. 3.4 Worldwide distribution of Amaranthus hypochondriacus L.

Fig. 3.5 A.
hypochondriacus L.
(a) Habit, (b) bract,
(c) tepal, (d) male
flower, (e) female flower
3.3.2 Amaranthus cruentus L. Southwestern USA, light brown grain type of A.

cruentus is used as a source of dye for colouring
The species was domesticated in Central America corn-based food in the Indian Pueblos where it
(Mexico and Guatemala) much earlier than A. probably became established in prehistoric times.
hypochondriacus. It is useful both as a source of Among the Amaranthus species, it is probably
pseudocereal and leafy vegetable and consists of the most adaptive and can bear flowers under a
two grain types – the white-seeded type used as wider range of day lengths. Amaranthus cruentus
pseudocereal and brown-seeded vegetable type L. with yellowish white or pale brown seed is tra-
used as vegetable. The latter type is also used to ditionally grown as pseudocereal crop in Latin
extract red dye. The remains of pale grains and American countries like Mexico, Guatemala,
bundle of plants for threshing at a dozen levels Ecuador and Columbia. It is also cultivated for
which date back to 5500 years have been dug up commercial purpose in dry and hot regions of the
through archaeological excavation from the USA, Argentina and China (Fig. 3.6).
renowned Tehuacan caves in Central Mexico. In Amaranthus cruentus is commonly known as
a few Indian village of Guatemala and Southern blood amaranth, purple amaranth and caterpillar
Mexico, A. cruentus is still cultivated as grain amaranth. It is an erect, annual herb and almost
crops, and seedcakes made up of popped ama- glabrous or slightly pubescent at distal part,
ranth seeds are sold on streets. In the arid especially when young, attaining a height of
Fig. 3.6 Worldwide distribution of Amaranthus cruentus L.
2.0 mt, smaller than A. hypochondriacus. Stems erect, stigmas 3. Staminate flowers are situated
are erect, branched distally, mostly in inflores- at tips of inflorescences, tepals 5, stamens 5.
cence, to nearly simple, green or reddish purple. Fruits are obovoid to elongate-obovoid, circum-
Leaves are petiolate, petiole one-half as long as scissile utricle, tapering into a tower at apex,
to nearly equaling blade, rhombic-ovate or ovate 2–2.5 mm long, smooth or distally slightly
to broadly lanceolate, 3–14(−20) × 1.5–8(−15) rugose, dehisce regularly. Seeds are broadly len-
cm; in robust plants it is occasionally larger, ticular to elliptic-lenticular, showing various
cuneate to broadly cuneate, entire, acute to acu- colour morphs like white or ivory with reddish or
minate or subobtuse to slightly emarginate with yellowish tint, occasionally dark brown to dark
small mucro. Inflorescences are both terminal reddish brown, 1.2–1.6 mm in diameter, smooth
and axillary, erect, reflexed or nodding, usually or indistinctly punctate (Fig. 3.7).
dark red or deep beet-red, purple less commonly,
almost green or greenish red, leafless at least dis-
tally, huge and robust, lower inflorescence form- 3.3.3 Amaranthus caudatus L.
ing lax and soft spikes and higher-forming
panicles. Bracts are spatulate, 2–3 mm long, This species originated in the Andean highlands
equaling or slightly longer than tepals, with of Argentina, Peru and Bolivia where common
short-spinescent apex, nerve as long as style potato originated. The Spanish conquerors
branches. Pistillate flowers have five tepals: termed it as ‘Inca wheat’, but it appeared and
tepals straight, oblong to oblong-obovate or lan- domesticated much earlier than Inca civilisation.
ceolate, not clawed, equal or subequal, the inner Pale seeds of A. caudatus more than 2000 years
tepals shorter than outer tepals, 1.5–3 mm long, old were discovered from the tombs, where it
with acute apex. Style branches are slender, was kept as food for the dead. Continuing the
Fig. 3.7 A. cruentus L.

(a) Habit, (b) a small
part of inflorescence,
(c) bract, (d) bracteole,
(e) tepal, (f) male flower,
(g) female flower
tradition, the plants are still grown in the Andean monly reddish or purplish throughout. Stems are
highlands mostly by the Indians to maintain the rather stout, branched sparsely and glabrous or
traditional custom. The plants are grown not in a finely covered with rather long, multicellular
large scale as a staple crop but in a small patch hairs which are increasingly numerous upwards.
adjacent to houses. The species is characterised Leaves are simple, long petiolate (petiole up to
with pendulous, blazing-red, elephant hoodlike 8 cm long but not longer than the lamina), lamina
inflorescences, commonly sold in European and broadly ovate to rhomboid-ovate or ovate-elliptic,
American countries as an ornament with a nick- lanceolate, 3.0–15 × 1–8 cm, apex acute to sub-
name ‘love-lies-bleeding’ or ‘red-hot-cattail’ . acute or obtuse, base shortly cuneate to attenuate,
Other forms of the species give much better glabrous, sparingly pilose along the margins and
grain yields. The crop shows a great deal of lower surface of the primary venation.
genetic variability in South America, only a Inflorescences are extremely long, soft and lax
small sample has been introduced to other conti- (up to 1.5 m.) of drooping spikes or panicles,
nents (Fig. 3.8). The grains are ground into flour, with knobby appearance due to large glomerules
or boiled for gruel, toasted and popped to be spaced relatively far apart. Flowers are arranged
used in seedcakes. It is considered useful for in axillary and terminal spikes formed of increas-
children and invalids. In India, germplasms col- ingly aggregated cymose clusters; the terminal
lected from the Northern part showed much inflorescences vary from a single, elongated, tail-
genetic diversity. like, pendulous spike to a panicle with the ulti-
Amaranthus caudatus L. is an annual, erect mate spike so formed. Male and female flowers
herb, attaining a height of 2.0 mt, and is com- are intermixed throughout the spikes. Bracts are
3.4 Potentiality of Grain Amaranth as Food 23
Fig. 3.8 Worldwide distribution of Amaranthus caudatus L.
deltoid-ovate, pale-membranous, acuminate with 3.4 Potentiality of Grain

a long, pale or reddish, rigid, erect arista formed Amaranth as Food
of excurrent midrib, the longest one up to twice
as long as the perianth, as long as style branches. The seeds of the grain amaranths on average are
In perianth with five tepals, those of the male composed of 13.1–21.0 % of crude protein, 5.6–
flowers are oblong-elliptic, acute, aristate, 2.5– 10.9 % of crude fat, 48–69 % of starch, 3.1–5.0 %
3.5 mm long, and those of the female flowers are of dietary fibre and 2.5–4.4 % of ash. The ‘pro-
broadly obovate to spatulate, abruptly narrowed tein component of grain amaranth and its quan-
to a blunt or sometimes faintly emarginate, tity and quality (as far as amino acid composition
mucronate tip, 1.75–2.5 mm long, inner tepals is concerned) are very close to the levels recom-
shorter than outer tepals. Staminate flowers are mended by FAO/WHO (Table 3.2). Protein effi-
mostly at the tips of inflorescence, stamens 5. ciency ratio (PER) of grain amaranths ranges
Style branches spreading meeting in a saddle at from 1.5 to 2.0 (value is 2.5 for Casein); for
base, stigmas 3. Fruits are ovoid-globose, cir- cooked grains, proteins have high digestibility
cumscissile and utricle, not forming tower at the (approx. 90 %) and are rich with lysine (0.34 g
apex; the lid is smooth or furrowed below, Lys/g N). Extremely balanced amino acid com-
abruptly narrowed to a short, thick neck, 2.0– position is due to the fact that in amaranth, 65 %
2.5 mm long. Seeds are compressed, shiny, of the proteins are found in the embryo and only
creamy white or grey coloured with a thick yel- 35 % in the perisperm, while in other grains
lowish or pink rim and translucent centre, 0.75– amino acids are mainly concentrated in the endo-
1.25 mm in diameter (Fig. 3.9). sperm (85 % in average) and embryos are poorer
Fig. 3.9 A. caudatus L.

(a) Habit, (b) bract,
(c) tepal, (d) male
flower, (e) female flower
Table 3.2 Seed composition in different grain amaranths

Components A. cruentus A. hypochondriacus A. caudatus
Crude protein 13.8–21.5 15.0–16.6 13.1–21.0
Lysine (g100 g−1 on dry basis) 4.9–6.1 4.9–6.0 5.9
Crude fat 5.6–8.1 6.1–7.3 5.8–10.9
Crude fibre 3.1–4.2 4.9–5.0 2.7–4.9
Carbohydrate 63.1–70.0 67.9 63.7–76.5
Ash 3.0–3.8 3.3–3.4 2.5–4.4
Squalene (% in oil) 2.2–6.9 1.9–4.6 3.8–6.7
Sources: Becker et al. (1981), Sanchez-Marroquin et al. (1986), Lyon and Becker (1987), Pederson et al. (1987), Singhal
and Kulkarni (1988), Ayorinde et al. (1989), Gorinstein et al. (1991), Prakash and Pal (1992), Bressani (1993), Dodok
et al. (1994), Zheleznov et al. (1997), Marcone and Yada (1998) and Leon-Camacho et al. (2001)
3.5 Vegetable Amaranths 25
Table 3.3 Distribution of essential amino acids in seeds of different grain amaranths and other cereals (g 100−1 of
protein)
Amino acids
Protein sources Trp Met/Cys Thr Isl Val Lys Phe/Tyr Leu LAAA EAAB
FAO/WHO (1973) 1.0 3.5 4.0 4.0 5.0 3.5 6.0 7.0 – –
Amaranth (average)a 1.3 4.4 2.9 3.0 3.6 5.0 6.4 4.7 67 87
A. cruentusb – 4.1 3.4 3.6 4.2 5.1 6.0 5.1 84 89
A. cruentusc 0.9 4.6 3.9 4.0 4.4 6.0 7.9 6.2 88 95
A. cruentusc – 4.6 3.9 4.0 4.5 6.1 8.5 6.1 87 96
A. caudatusc 1.1 4.9 4.0 4.1 4.7 5.9 8.1 6.3 90 98
A. hypochondriacus 1.82 0.6 3.3 2.7 3.9 5.95 8.42 4.2 34 78
A. cruentuse 1.4 4.1 3.4 3.6 4.2 5.1 6.0 5.1 73 91
Amaranth (average)a-e 1.3 4.5 3.5 3.6 4.2 5.6 7.3 5.4 75 94
Barleya 1.2 3.2 3.2 4.0 4.7 3.2 8.2 6.5 83 97
Buckwheata 1.4 3.7 3.9 3.8 5.2 5.9 5.8 5.8 83 97
Maizea 0.6 3.2 4.0 4.6 5.1 1.9 10.6 13.0 35 86
Oata 1.2 3.4 3.1 4.8 5.6 3.4 8.4 7.0 62 92
Rice 1.0 3.0 3.7 4.5 6.7 3.8 9.1 8.2 69 94
Soyaa 1.4 3.1 3.9 5.4 5.3 6.3 8.1 7.7 89 98
Wheata 1.2 3.5 2.7 4.1 4.3 2.6 8.1 6.3 47 86
Sources: aSenft (1979), bBetschart et al. (1981), cBecker et al. (1981), dDodok et al. (1997) and eSanchez-Marroquin
et al. (1986)
with essential amino acids (Table 3.3). The com- 3.5 Vegetable Amaranths
pilation of maize and amaranth grain flour in
50:50 ratio nearly reaches the perfect score of Vegetable amaranths are considered as the most
100 on the nutritionist’s scale (Segura-Nieto et al. popular vegetable crops grown in the tropics for
1994; Saunders and Becker 1984; Grobelnik their protein, vitamin and mineral-rich leaves and
et al. 2009a, b). In grain amaranths the saturated stems. Vegetable amaranths are grown in the hot,
and unsaturated fatty acid ratio ranges from 0.29 humid regions of Southeast Asia (especially
to 0.43. The shares of linoleic acid, oleic acid, Malaysia and Indonesia), Africa, Southern China,
palmitic acid, stearic acid and linolenic acid are India and Caribbean islands. Leaves of most
25–62 %, 19–35 %, 12–25 %, 2–8.6 %, and 0.3– Amaranthus species are edible, but few are very
2.2 %, respectively. Amaranth seed oil has been popular, e.g. vegetable amaranths such as A. tri-
reported to contain large amount (7–8 % and color L., A. blitum L., A. dubius, A. cruentus and A.
11 %) of squalene which is often used in cosmet- viridis L., the first two being the most popularly
ics and medicine, where olive oil contains only grown. Mild spinach-like flavour of vegetable ama-
1 % of squalene. Amaranth oil is also a rich ranths, high yields, ability to grow in hot weather
source of tocotrienols which is very effective to and high nutritive value are few reasons for their
lower the LDL cholesterol (Becker et al. 1981; popularity. They are probably the most widely con-
Plate and Areas 2002). Anti-nutritive components sumed leafy vegetables in the humid tropical low-
like saponins, trypsin inhibitor and tannin are land of Africa and Asia (Schnetzler and Breene
present in amaranth grain keeping parity with 1994). Amaranthus tricolor is very rich in morpho-
legumes and some other grains like sorghum. logical diversity, represented in a number of differ-
Now these components are not considered as ent morphotypes (Fig. 3.10). Vegetable amaranths
nutritional hazard. are very rich in protein; calcium; iron; vitamin A, C
Fig. 3.10 Different

varieties of Amaranthus
tricolor L.
3.6 Species of Vegetable Amaranths 27
and K; riboflavin (B2); niacin (B3); vitamin B6; 2006b). This underutilised plant with significant
and folate which have attributed to their high nutri- nutritive value has been recognised by the US
tive value. Due to the presence of a number of male National Academy of Science (1984). This popular
flowers per glomerule, terminal inflorescence and vegetable is supposed to have originated in tropical
development of axillary glomerules (Rajan and South Asia (Grubben and Van Sloten 1981) and
Markose 2007), self-pollination is the predominant spread throughout the tropical and temperate
phenomenon in vegetable amaranth. In spite of the regions of the globe (Martin and Telek 1979). In
fact that vegetable amaranth is used as a cheap South and Southeast Asia, it is a major leafy vege-
source of protein and staple food crop in many table. It is occasionally cultivated in East, West and
parts of the world, negligible efforts have been Southern Africa. It was domesticated in prehistoric
made for its genetic improvement. times from the wild progenitor which is not clearly
known. Weedy plants of Amaranthus tricolor can
be found occasionally as an escape from cultiva-
3.6 Species of Vegetable tion. It is far from competitive with true weeds.
Amaranths Amaranthus tricolor occurs as a rare exotic vegeta-
ble in several African countries, thought to have
3.6.1 Amaranthus tricolor L. been introduced by Indian immigrants and occa-
sionally cultivated especially in East and Southern
It has been recognised as good as or superior in Africa. Its cultivation has been reported from
taste to spinach and considerably rich in carotenoid Benin, Nigeria, Kenya and Tanzania. In many trop-
(90–200 mg/kg), protein (14–30 % on dry weight ical regions of India, Southeast Asia and South
basis) and ascorbic acid (nearly 28 mg/100 g) Pacific Islands, Amaranthus tricolor is extensively
(Makus 1990; Prakash and Pal 1991; Shukla et al. grown (Fig. 3.11).
Fig. 3.11 Worldwide distribution of Amaranthus tricolor L.

Amaranthus tricolor is an annual, ascending or Male and female flowers are intermingled. Bracts
erect herb, attaining a height of 1.0–1.25 m or and bracteoles are deltoid-ovate, bracteoles sub-
more in cultivation. Stem is usually much- equalling or shorter than the perianth, pale-mem-
branched and stout and the branches are angular, branous, broadest near the base and narrowed
glabrous or furnished in the upper parts with sparse upwards to the green midrib, which is excurrent to
(or denser in the inflorescence), more or less form a long, pale-tipped awn usually at least half
crisped hairs. Leaves are simple, petiolate, the as long as the basal portion and not rarely equal-
lamina broadly ovate, rhomboid-ovate or broadly ling it. Perianth consists of three tepals: tepals
elliptic to lanceolate-oblong, lamina tip emargin- 3–5 mm long, elliptic or oblong-elliptic, narrowed
ate to obtuse or acute, base shortly cuneate to above, pale-membranous, the green midrib excur-
attenuate, decurrent along the petiole, glabrous or rent into a long, pale-tipped awn. Male flowers
thinly pilose on the lower surface of the primary have three stamens, female flowers with the tepals
venation, green or purplish, very variable in size slightly accrescent in fruit, stigmas 3, erect or
(up to 18 cm long). Flowers are green to crimson recurved, 2 mm long. Fruits are ovoid-urceolate
in more or less globose clusters of 4–25 mm in utricle with a short neck below the style-base,
diameter; all or only the lower parts are axillary 2.25–2.75 mm, circumscissile, membranous,
and distant, with the upper sometimes without obscurely wrinkled. Seeds are blackish- brown,
subtending leaves and increasingly approximate to lenticular, shiny, 1–1.5 mm in diameter, spermo-
form a thick terminal spike of variable lengths. derm very faintly reticulate (Fig. 3.12).
Fig. 3.12 Amaranthus

tricolor L. (a) Habit, (b)
a part of inflorescence,
(g) pistillate flower
Fig. 3.13 Worldwide distribution of Amaranthus blitum L.
3.6.2 Amaranthus blitum L. base, glabrous, attaining a height of 10–30 cm.

Leaves are simple, petiolate, petiole 1–3.5 cm,
Though it is supposed to have originated in the leaf blade ovate or ovate-rhombic, with notched
Mediterranean region, now it is found worldwide apex with a mucro, cuneate base, entire or slightly
ranging from the tropical to temperate region. It undulate margin, dimension 4.0–7.5 × 3.0–6.0 cm.
is one of the most popularly cultivated vegetables Flowers are arranged in slender terminal spikes or
in India, and its cultivation has been reported panicles and also in axillary clusters; spikes are
from East and Central Africa. It has also been erect or sometimes reflexed. Bracts and bracteoles
recorded from many African countries and prob- are oblong, shorter than 1 mm. Perianth has three
ably occurs throughout tropical Africa, from tepals, tepals are elliptic, spatulate, oblong or lan-
Senegal to Ethiopia, South Africa and the Indian ceolate, with an adaxial midvein, equal or sub-
Ocean islands. It is generally a protected weed in equal, apex acute, light green, length 1.5–2.0 mm.
backyards and home gardens and occasionally Staminate flowers are clustered at the tip of spikes,
produced for sale in the market. In Greece, Japan stamens 3, slightly shorter than tepals. Style
and Western Europe, it is a popular green vegeta- branches are erect, stigmas 3, fall off when utricle
ble and is used as a substitute for spinach matures. Fruits are compressed-ovoid, utricle,
(Spinacia oleracea) during the hot summer sea- exceeding tepals, indehiscent, slightly rugose to
son (Fig. 3.13). nearly smooth, 3 mm long. Seeds are black to
Amaranthus blitum is an annual herb with light brownish black, circular in outline, 1.2 cm in
green or purple, ascending stem, branched from diameter (Fig. 3.14).
in distal portion. Stems are green and branched.

Leaves are simple, petiolate, petioles of proximal
leaves are equal or longer than blade, becoming
shorter distally, lamina rhombic-ovate or ovate to
elliptic, 3–12 × 2–8 cm, with entire margin,
slightly acuminate to obtuse and faintly emargin-
ate, mucronate tip and broadly cuneate base.
Inflorescences occur as dense terminal panicles
and axillary spikes, and panicles are erect or
often drooping, green, branched, leafless at least
distally. Bracts are lanceolate, less than 2 mm,
shorter than tepals with spinescent tip. Pistillate
flowers have five tepals; tepals are oblong-
spatulate to oblong, with acute apex often shortly
mucronate, 1.5–2 mm long. Style branches are
strongly spreading, shorter than fruit, stigmas 3.
Staminate flowers are generally clustered at tips
of inflorescence branches, sometimes aggregated
in proximal glomerules (as in A. spinosus) with
five tepals equal or subequal and five stamens.
Fruits are ovoid or subglobose, circumscissile
utricle,1.5–2 mm long, slightly shorter than
tepals, smooth to irregularly wrinkled, dehisce
regularly. Seeds are dark reddish brown to black,
subglobose or lenticular, 0.8–1 mm in diameter
Fig. 3.14 Morphology of vegetable amaranths, A. blitum
L. with shiny, smooth spermoderm (Fig. 3.16)
3.6.3 Amaranthus dubius L. 3.6.4 Amaranthus cruentus L.
It may have evolved from weedy ancestors in The vegetable form of Amaranthus cruentus was
tropical Asia (Indonesia and India) and may have probably introduced in the tropics and subtropics
been introduced later to Africa and Central of the Old World during the colonial period. Now
America by immigrants. This weedy species is Amaranthus cruentus is familiar as a widespread
occasionally used as a green leafy vegetable in traditional vegetable in all countries of tropical
West America and the Caribbean islands and is Africa. It is the principal leafy vegetable in Benin,
found in Java and other parts of Indonesia as a Togo and Sierra Leone and very important in
home garden crop (Fig. 3.15). One of the best many lowland areas like in Southern Nigeria, DR
varieties of this species, known as the cultivar Congo, Kenya and Tanzania. It is more popular in
‘claroen’, is particularly popular in Benin and humid lowland than in highland or arid areas. It is
Suriname. Its seeds are extremely small (4500 also an important leafy vegetable in many tropical
seeds per g). It is a fast-growing, high-yielding countries outside Africa, e.g. in India, Bangladesh,
plant with considerable morphological variabil- Sri Lanka and the Caribbean islands. The
ity. It has characteristic dark green, broad, ridged Bangladesh type has big fleshy stems and is con-
leaves. This is the only known tetraploid (2n = 64) sumed with the leaves. Amaranthus cruentus is
species in the genus Amaranthus so far. grown throughout Southeast Asia as leafy vegeta-
Amaranthus dubius (Spleen amaranth) is an ble, although to a lesser extent than Amaranthus
annual erect herb, glabrous or slightly pubescent tricolor L. During thinnings of young seedlings
Fig. 3.15 Worldwide distribution of Amaranthus dubius Mart. ex Thell
in the cultivation field of the grain crop, uprooted

seedlings are frequently used as a vegetable. A
very deep red coloured, dark-seeded morphotype
of the species, sometimes known as ‘blood
Amaranth’, is often sold as ornamentals. During
the nineteenth century, this deep-red vegetable
form was adopted as green vegetable by garden-
ers throughout the tropics. It soon appeared as a
more important crop in tropical Africa than any-
where else. Like corn, sweet potatoes, peanuts
and other American Indian crops, Amaranthus
cruentus was evidently introduced in Africa by
Europeans. After being introduced it was quickly
transferred from tribe to tribe, probably as a weed
in millet and sorghum seed. Now, it has colonised
a large area in Africa and is being planted and
gathered year-round in the humid regions of
America. The tender young seedlings are pulled
up by the roots and sold in town markets by the
thousands of tons every year in parts of West
Fig. 3.16 Plant morphology of Amaranthus dubius Mart.
America.
ex Thell
3.6.5 Amaranthus viridis L. terminal, often paniculate spikes, 2.5–12 cm long

and 2–5 mm wide, or in the lower part of the stem
Amaranthus viridis is possibly of Asian origin, but is in dense axillary clusters. Male and female
now it has become a cosmopolitan weed in the trop- flowers are intermixed but the latter is more
ical and subtropical regions of the world, also reach- numerous. Bracts and bracteoles are deltoid-
ing far into temperate regions (e.g. in Europe, North ovate to lanceolate-ovate, white-membranous
America, Asia and Australia). In tropical Africa it is with a very short, pale or reddish awn formed by
a widespread common weed. It is occasionally cul- the excurrent green midrib, bracteoles are shorter
tivated in Nigeria, Gabon and DR Congo. than the perianth (c. 1 mm.). Tepals are three in
It is an annual herb, erect or more rarely number, very rarely four, those of the male flow-
ascending, attaining a height of 75–100 cm. ers are oblong-oval, acute, concave, 1.5 mm long,
Stems are rather slender, sparingly to consider- shortly mucronate, while those of the female
ably branched, angular, glabrous or more fre- flowers are narrowly oblong to spatulate, 1.25–
quently increasingly hairy upwards (especially in 1.75 mm long, minutely mucronate or not, with
the inflorescence) with short or longer and rather white-membranous borders, midrib green and
floccose multicellular hairs. Leaves are glabrous often thickened above. Male flowers are incon-
or shortly to fairly long-pilose on the lower sur- spicuous, mostly at the tips of inflorescence, with
face of the primary and most of the venation, three stamens. Stigma 2–3, short, erect or almost
long-petiolate (petioles up to 10 cm long and the so. Fruits are subglobose utricle, not or slightly
longest commonly longer than the lamina), lam- exceeding the tepals, indehiscent or rupturing
ina is deltoid-ovate to rhomboid-oblong, the mar- irregularly, very strongly rugose throughout,
gins entire, occasionally obviously sinuate, apex 1.25–1.5 mm. Seed are round, slightly com-
obtuse and obscurely to clearly emarginate at the pressed, black with an often paler thick border,
tip, minutely mucronate, base shortly cuneate to 1–1.25 mm in diameter, more or less shiny, retic-
sub-truncate below, 2–7 × 1.5–5.5 cm in length. ulate and with shallow scurfy verrucae on the
Flowers are green, arranged in slender, axillary or reticulum (Fig. 3.17a).
Fig. 3.17 General plant morphology of (a) A. viridis L. (b) A. graecizans L.

In Asia, domestication of amaranths took short, gland-like hairs. Flowers are arranged all
place for use as pot herbs, and a variety of culti- in axillary cymose clusters, male and female
vated amaranth races belonging to A. lividus and flowers are intermixed, and male flowers are gen-
A. tricolor are common pot herbs in Eastern and erally situated mostly in the upper whorls. Bracts
Southern Asia which subsequently have been and bracteoles are narrowly lanceolate-oblong,
introduced into the New World. A. tricolor might membranous, acuminate, with excurrent green
have originated in the Indian subcontinent espe- midrib forming a pale or reddish arista, bracte-
cially in India and have spread to neighbouring oles are subequalling or usually shorter than the
countries and other parts of the world by the tepals, bract length 1.5–2 mm. Perianth consists
immigrants. Vegetable amaranths like A. tricolor, of three tepals, all 1.5–2 mm; those of the male
A. melancholicus, A. gangeticus, etc., are impor- flowers are lanceolate-oblong, acute or subacute,
tant group of cultivated amaranths which are pale-membranous with a narrow green midrib
obviously natives of Asia. However, these are excurrent in a short, pale arista, and those of the
grown exclusively as pot herbs or ornamentals, female flowers are lanceolate-oblong to linear-
apparently never for seeds. This entire Asiatic oblong, gradually to abruptly narrowed to a very
potherb group is clearly distinguishable from the short to rather long mucro, the midrib often bor-
grain group. dered by a green vitta above and apparently
thickened, the margins pale whitish to greenish.
Staminate flowers are intermixed with pistillate
3.6.6 Amaranthus graecizans L. flowers, male flowers have three stamens and
female flowers have three stigmas; stigma is slen-
Amaranthus graecizans occurs scattered through- der, usually pale, flexuose, 0.5 mm long. Fruits
out tropical Africa. It is well distributed in are subglobose to shortly ovoid utricle, 2–2.25
Southern Europe and in tropical and subtropical mm, usually strongly wrinkled throughout with a
Asia and has also been introduced to the USA. It very short, smooth neck, slightly exceeding the
is very popular as a vegetable in parts of Kenya, perianth, circumscissile or sometimes not, even
Uganda, Tanzania, Malawi and elsewhere in on the same plant. Seeds are shiny, compressed,
Southern Africa and sometimes appears as a pro- black and 1–1.25 mm in diameter; spermoderm is
tected weed in backyards, cultivated locally on faintly reticulate (Fig. 3.17b).
small scale in home garden and for market sale in Amaranthus graecizans is mainly used as
Nebbi, Uganda and in Tanzania. cooked leaf vegetable. In many countries it is col-
Amaranthus graecizans is an annual erect, lected from the wild as a potherb. Though it tastes
decumbent or prostrate herb, branched, pubes- slightly bitter, such slight bitterness is liked by
cent in distal parts or become glabrous at matu- the older people. A major drawback is that the
rity, attaining a height up to 45 cm (rarely to 70 leaves are small, so the collection is time taking.
cm). Stems are slender to stout, angular, glabrous The plant has axillary clusters of flowers; the
or finely to moderately hairy; hairs are short or people don’t like to cook the whole shoot, but
long, often crisped multicellular, which increase they have to pluck the individual leaves which is
upwards, especially in the inflorescence. Leaves time taking. This is one of the reasons of its low
are simple, long-petiolate (petiole length 3–4.5 market value.
cm, sometimes longer than the lamina), lamina In some regions it is consumed by mixing
broadly ovate or rhomboid-ovate to elliptic-ovate with other leaf vegetables collected from the
or narrowly linear-lanceolate or linear, wild, as, for example, the Okiek people in
4–55 × 2–30 mm, with generally entire margins, Western Kenya, who mix it commonly with
subacute to obtuse or emarginate, mucronulate Solanum or Rumex species and Urtica massaica
apex and cuneate to broadly cuneate base, gla- Mildbr. Amaranthus graecizans is also used as a
brous or occasionally the lower surface of the fodder for livestock. In Mauritania the seed is
principal veins is sparingly covered with very baked into thin cakes, while in the Western USA,
it is ground into flour. The whole plant of According to different authors, oxalate and
Amaranthus graecizans is used in East and West nitrate concentrations in fresh matter vary from
Africa to manufacture a local salt. For this pur- 4.1 to 9.2 g kg−1 and from 3 to 16.5 g kg−1, respec-
pose, the plants are first dried and then burned to tively. Compounds are generally associated with
ashes, the filtrate is evaporated and the residue is forage and vegetable application, and they
used as a substitute for common salt. In Uganda, strongly depend upon genotype and fertilisation
the leaves are chewed and the liquid extract is practice (Williams and Brenner 1995).
swallowed for the treatment of tonsillitis. In The vegetable amaranths are cultivated or col-
Senegal, the leaves are used as an anthelmintic. lected from wild in so many regions of the world,
but few references are available regarding their
culture. This indicates the facts that their cultiva-
3.7 Nutritive Value of Vegetable tion practice is easy and optimal conditions for
Amaranths maximum yields are unknown due to their wide
adaptability. Often seeds are not at all sown, fall
Prakash and Pal (1991) reported high nutritive naturally in the cultivation field and raise the crop
value of vegetable amaranth. It is rich in protein of the next year. The plants grow rapidly, so the
(14–43 g kg−1 in fresh matter), lysine (40–56 g time interval between planting and harvest of the
kg−1), carotenoids (60–200 mg kg-1) and different tender foliage and stems is short (generally only
types of vitamins and minerals (Table 3.4). 3–6 weeks). In Tamil Nadu (South India), plants
Table 3.4 Nutritional value of raw and cooked amaranth leaves, in comparison with raw cabbage, Chinese cabbage
and spinach
Cabbage raw Chinese cabbage Spinach raw Amaranth raw Amaranth cooked
Nutrients (value/100 g) raw (value/100 g) (value/100 g) (value/100 g) (value/100 g)
Proteins (g) 1.28 1.20 2.86 2.46 2.11
Minerals
Calcium(mg) 40 77 99 215 209
Iron (mg) 0.47 0.31 2.71 2.32 2.26
Magnesium (mg) 12.0 13.0 79.0 55.0 55.0
Phosphorus (mg) 26.0 29.0 49.0 50.0 72.0
Potassium (mg) 170.0 238.0 558.0 611.0 641.0
Sodium (mg) 18.0 9.0 79.0 20.0 21.0
Zinc (Zn) 0.18 0.23 0.53 0.90 0.88
Copper (mg) 0.019 0.036 0.136 0.162 0.158
Manganese (mg) 0.160 0.190 0.897 0.885 0.861
Vitamins
Vitamin C (mg) 36.6 27.0 28.1 43.3 41.1
Riboflavin (mg) 0.040 0.050 0.189 0.158 0.134
Niacin (mg) 0.234 0.400 0.724 0.658 0.559
Vitamin B6 0.124 0.232 0.195 0.192 0.177
Folic acid (mcg) 43.0 79.0 194.0 85.0 57.0
Vitamin A RAE 1 5.0 16.0 469.0 146.0 139.0
(mcg)
Vitamin K 76.0 42.9 482.9 1140.0 –
Lipids
Total saturated fatty 0.034 0.043 0.063 0.091 0.050
acids (g)
Source: USDA National Nutrient Database for standard Reference, Release 23 (2010) http://www.nal.usda.gov/fnic/
foodcomp/search
3.8 Uses of Amaranths 35
are pulled out 3 weeks after sowing and used as 1. Provides energy: Vegetable amaranths are
‘tender greens’. Certain varieties of vegetable very rich in carbohydrates, proteins, vitamin
amaranths remain succulent for longer period and K, folate, riboflavin, vitamin A, vitamin B6
can be harvested even up to 5 weeks of growth. and vitamin C. Amaranth leaves boost energy
Certain varieties are suitable for periodical cutting in the body. The crude protein content in the
(clipping). The first cutting is done at 20 days leaves ranges from 20 to 32 %, on a dry
after sowing and subsequently followed by weight basis.
weekly cuttings up to 10 cuttings. At a later age 2. Prevents electrolyte imbalance: Amaranth
generally after flowering, the foliage and stems leaves are very good source of elements like
become fibrous, brittle, pithy and unpalatable. manganese, iron, copper, calcium, magne-
sium, potassium and phosphorus necessary
to maintain adequate electrolyte balance in
3.8 Uses of Amaranths the body.
3. Excellent gluten-free diet: Vegetarians with
Amaranths are the source of highly nutritious gluten intolerance or those suffering from
food. With the ever-increasing population and celiac diseases can get daily recommended
fast-growing depletion of natural resources, it dose of protein from amaranth greens.
became necessary to explore the possibilities of Compared to other plant sources, namely,
using newer plant resources, new crop and new wheat, rye, rice and oats, amaranths are glu-
usage of old ones that hold promise to restore the ten-free and contain 30 % more protein with
balance of trade and meet the growing needs of complete set of amino acids.
food, clothes and industrial products for human 4. Improves digestion: It can improve the diges-
population. The underutilised amaranths are tive system and reduce constipation due to
good options and projected as a future crop of the the high content of dietary fibre which is
twenty-first century. Amaranths are categorised three times that of wheat. It is easily digest-
into three – vegetable, grain and weed. Vegetable ible, so it is good for both the young ones and
amaranths are widely consumed as leafy vegeta- elder people.
bles in India and other Asian and Southeast Asian 5. Aids in weight management: The protein in
countries, also in African countries but not so the leaves helps to reduce insulin levels in
familiar in North and South America, whereas the blood and also releases a hormone that
grain amaranths are widely consumed as highly lessens hunger pranks and prevent ‘binging
nutritious pseudocereals. catastrophe’.
6. Reduces bad cholesterol: One of the key
health benefits of vegetable amaranth leaves
3.8.1 Uses of Vegetable Amaranths is their cholesterol-lowering capacity. Due to
the high fibre content, this leafy vegetable is
The leaves, shoots and juicy tender stems of effective in reducing LDL levels in the blood
many cultivated species of vegetable amaranths and promotes weight loss. Tocotrienols, a
are consumed as a potherb in sauces or soups, type of vitamin E available in vegetable ama-
cooked with other vegetables and taken with a ranths, also contributes to its cholesterol-
main dish or by itself. Young leaves of grain ama- lowering ability.
ranths are also used as leafy vegetable. Amaranth 7. Good for anaemic patients: Iron-rich (five
leaves are a good source of high amount of pro- times that of wheat) red amaranth leaves pro-
tein, vitamins, minerals and dietary fibre. mote coagulation and increase haemoglobin
Chopped plants can also be used as forage for content and red blood cell counts. It is also
livestock. an excellent source of folic acid which is
Vegetable amaranths have the following ben- necessary to increase the blood haemoglobin
eficial role on human health: level.
8. Decreases risk of cardiovascular disease: ous/carcinogenic nitrosamines in the digestive

Amaranth leaves are excellent dietary source tract, though evidence supporting this is not
of phytosterols that lowers blood pressure available at present. Boiling of the amaranth
and prevents heart ailments including stroke. leaves like spinach or chard for 5–10 min and
9. Fights off cancer: The presence of lysine (an then discarding the water are proven effective to
essential amino acid) along with vitamin E, remove both oxalates and nitrate problems,
iron, magnesium, phosphorus, potassium though research has shown that consumption of
and vitamin C helps to fight against free radi- 200 g of cooked amaranth per day does not create
cals responsible for ageing and formation of any health problem.
malignant cells. The present level of nitrate and oxalate in
10. Ayurvedic treatments: Juice extracted from amaranths does not create a nutritional problem
fresh amaranth leaves is good for treating under normal condition of consumption, but con-
diarrhoea and haemorrhage. sidering this aspect, Devadas et al. (1984) anal-
11. Stops hair loss and greying: Amaranth leaves ysed 25 amaranth genotypes for oxalate and
have wonderful cosmetic benefit. Besides reg- nitrate content. The oxalate content ranged from
ular consumption, application of leaf juices 0.94 to 1.29 % and the nitrate content from 0.55
prevents brittle hair from falling. This wonder- to 1.0 %. The soluble oxalate content of A. gange-
ful cosmetic benefit of amaranth leaves also ticus was reported as 4.4 % and 7.4 % in the
retards the onset of premature greying. leaves and stems, respectively, on a dry weight
12. Prevents calcium-deficiency basis. The mean percentage of nitrate over two
ailments:Calcium present in amaranth leaves growing seasons were 0.51 %, 0.19 %, 0.39 %,
(two times that of milk) is helpful to reduce 0.54, 0.29 % and 0.65 % and those for oxalates
risk of osteoporosis and other calcium were 5.37 %, 5.59 %, 3.52 %, 6.95 %, 2.45 % and
deficiency-related disorders. 4.33 %, respectively, in the leaves of A. gangeti-
13. Improves eyesight: Amaranth leaves are cus, A. blitum, A. dubius, A. cruentus, A. cauda-
excellent source of β-carotene. Daily inclu- tus and A. hypochondriacus.
sion of vegetable amaranth in the diet can
help to prevent vitamin A deficiency which is
responsible for blindness. It is reported that 3.8.2 Uses of Grain Amaranths
the incidence of blindness in children due to
malnutrition has been reduced with the con- Amaranth grains contain more protein than corn,
sumption of 50–100 g of amaranth leaves per and the protein is also of an unusually high qual-
day. ity, rich in amino acid lysine, which is the limit-
ing amino acid in cereals like maize, wheat and
Amaranth leaves, like some other vegetables, rice, and sulphur-containing amino acids, which
contain rather high amounts of oxalic acid and are normally limiting in the pulse crops. The
nitrates. The amount of oxalic acid is roughly the total ‘protein package’ of amaranth grain is very
same as that found in spinach (Spinacia olera- near to the levels recommended by FAO/
cea) and chard (Beta vulgaris var. cicla). WHO. Grain amaranths have high protein scores.
Excessive amounts of oxalic acid may reduce the Protein scores are determined by taking the ratio
availability of certain minerals in the body, par- of the essential amino acids to the level for those
ticularly calcium. This could be of great concern amino acids recommended by FAO/WHO and
especially when the calcium intake levels are low multiplying by 100. The protein score of grain
or if foods containing high amount of oxalic acid amaranths is 67–87, while those of wheat (con-
are consumed on a regular basis for a long period taining 14 % protein), soybean (containing 37 %
of time. Nitrates present in amaranth leaves are protein), rice (containing 7 % protein) and maize
also of concern because it is speculated that (containing 9 % protein) are 47, 68–89, 69 and
nitrates may be chemically changed into poison- 35, respectively. Although amaranth is theoreti-
3.8 Uses of Amaranths 37
cally very close to the ideal level, combining it made a poppy amaranth by the thermal shock.
with another grain will increase its quality very Poppy amaranth is used in the cereal muesli, for
close to the FAO/WHO standards. However, the the muesli bars and also for bakery products to
actual nutritive value is less than expected from cover bread and rolls. The most positive proper-
the above considerations due to the presence of ties of poppy amaranth seeds are the low weight
anti-nutritional factors in the raw amaranth and the water absorption capacity. So the breads
grain. Cooking procedure can reduce the toxic and rolls remain soft and supple for longer time.
effects. The problems of unpalatability are The defatted amaranth grain flour was the most
caused by saponins and phenolic compounds in often used raw material in food industry in the
the amaranth grain (Cheeke et.al. 1981). It is a Czech Republic in the year 2008.
very nutritious animal feed, but the raw ama- Amaranth can be used as a substitute in por-
ranth grains contain toxins and anti-nutritional ridge, stirred into soups; Amaranth grains can be
factors that can lower its acceptability as an ani- cooked whole in a pot, rice cooker or pressure
mal feed. Protein-rich grains of grain amaranths cooker to prepare breakfast porridge or savoury
are easy to digest and are commonly given to ‘polenta’. The grain flour or flaked grains are
those who are recovering from illness or fasting combined with wheat or other flours to make
period. In India A. hypochondriacus is known as cereals, cookies, bread and other baked goods. As
the ‘king grain’ and is often popped to be used in per general recommendation, amaranth grain
confections. In the Third World countries, the flour should contribute only 10–20 % of the
seeds are generally ground into gluten-free flour mixed flour blended with wheat flour. But studies
and used for bread making with wheat or rye have shown that if amaranth grain flour blended
flour proportionately. Considering its signifi- up to 50–75 % of the mixed flour, it will still
cantly high nutritional properties, it is called the retain functional properties as well as flavour.
‘third millennium grain’. However, it has a high Coarsely ground amaranth grain makes a tasty
glycemic index due to its readily digestible and nutritious porridge cooked by itself or by
starch. To obtain maximum benefits, it should be mixing with other grains and pseudocereals such
best combined with nuts, seeds, pulses, legumes as oats (Avena spp.), wheat (Triticum spp.),
and vegetables. In addition, amaranth is rela- milled flax seed (Linum usitatissimum), wheat
tively a good source of cholesterol-lowering germ and cañihua (Chenopodium pallidicaule).
soluble fibre, calcium, iron, magnesium, zinc, Other seed components with significant potential
vitamins A and C and several B vitamins. include anthocyanin (red) pigments to produce
Flavonoids (such as rutin) and some phenolic nontoxic natural dyes, microcrystalline starch for
acids (such as gallic acid, p-hydroxybenzoic food and industry and squalene, specialised oil
acid and vanillic acid) with antioxidant effects used in skin cosmetics, computer and pharma-
are also found in amaranth seeds and sprouts. ceutical industries. Lipid fractions from amaranth
Many consumers prefer amaranth because they seeds contain high levels of unsaturated fatty
want a wheat- and gluten-free product, like the acids and possess a high antioxidative activity.
nutritional profile of amaranth, or enjoy ‘exotic’ Amaranth oil also contains a unique squalene
foods in their diet (Brenner et al. 2000). component, an intermediate of steroid synthesis,
Amaranth grain may be processed in various which is discussed as immunomodulator and is
ways, like grains can be popped, flaked, extruded supposed to play a role in the rate of cholesterol
and ground into flour. In Mexico Alegria, a con- synthesis (prevention of cardiovascular diseases).
fection made from popped amaranth grains is a Amaranth starch is of promising use. The fea-
popular and favourite food item among local peo- tures of starch like high solubility and digestibil-
ples and tourists. Popped amaranth can be enjoyed ity are due to its uniquely small size which is
on its own or can be served with milk or soymilk about one-tenth the size of cornstarch and there-
and fruit for a healthy breakfast. Amaranth grain fore offer new possibilities for food processing,
is very often used in the poppy machine, which pharmacology and cosmetics (Resio et al. 2000).
Amaranth starch shows unique gelatinisation and the antiretroviral drugs function poorly or not at
freeze/thaw properties which could be beneficial all on a poor diet. When the diet is poor in nutri-
to the food industry. Furthermore, amaranth- ents, often the drug even becomes a toxin in
originated products may reduce dietary fibre itself. Amaranth grain porridge (1 cup) combined
insufficiency, vitamin deficiency as well as defi- with Moringa leaf powder (1 Tbsp) from Moringa
ciency of bioactive compounds (antioxidants, leaves (Moringa oleifera) provides an excellent
folic acid). Amaranth grain oil is very special in nutritious food not only for the AIDS sufferer but
the content. The amaranth oil is used in cosmetics also for those who are taking antiretroviral drugs
in the pure form or with added tocopherol and it without any complications.
is given under argon for stabilisation. The advan- Several health benefits like decrease in plasma
tage of amaranth oil for cosmetic industry is due cholesterol levels, protection of heart, stimulation
to its squalene content. It is nonallergic on the of the immune system, anticancer activity, reduc-
skin and has a protective function for very sensi- tion of blood glucose levels, improvement of
tive skin and hair against sun and radiation. So, it hypertension and anaemia, etc., are provided by
can be utilised in various sun cream and lotions grain amaranths. In addition, it has been reported
Food specialists have evaluated the amaranth to have anti-allergic and antioxidant properties.
seed flour as an additive to wheat flour. To Most of these properties are due to the presence of
increase the palatability, different amounts of bioactive compounds. The cholesterol-lowering
amaranth grain flour were mixed with the wheat effects in amaranth may be due to unsaturated
flour and baking ingredients (1 % salt, 2.5 % fat, fatty acids. Being a good source of magnesium
1.5 % yeast, 10 % sugar and 52–74 % water), fer- which is effective to relax blood vessels and pre-
mented, moulded, pan-proofed and baked. The vent constriction and rebound dilation, it helps to
baked products were evaluated for loaf volume, fight migraines. Both the vegetable and grain
moisture content, colour, odour, taste and texture. amaranths are equally significant from nutritional
The bread containing amaranth grain flour prod- point of view (Table 3.5)
ucts were then compared with the bread made
exclusively from wheat flour. The loaf volume
decreased by 40 % and the moisture content Table 3.5 Comparison of nutritive components of vege-
table and grain amaranths (100 g portion)
increased from 22 to 42 % with the increase in
amaranth grain flour. The study showed that the Vegetable Grain
Components amaranths amaranths
sensory levels of taste, odour, colour and texture
Moisture 86.9 g 9.0 g
decreased or degraded with increasing amounts
Protein 3.5 g 15.0 g
of amaranth flour. Significant deviation in sen-
Fat 0.5 g 7.0 g
sory qualities was detected when more than 15 %
Total carbohydrate 6.5 g 63.0 g
of amaranth grain flour was added, and products
Fibre 1.3 g 2.9 g
containing high amount of amaranth grain flour
Calories 36 391
were found to be of unacceptable palatability to Phosphorous 67 mg 477 mg
the population as appeared from the sample sur- Iron 3.9 mg –
vey of baked product. The Organic Farming and Potassium 411 –
Research Center (Rodale) has successfully used a Vitamin A 6100 IU –
50:50 ratio of wheat flour and amaranth grain Riboflavin 0.16 mg 0.31 mg
flour and suggested that the percentage of ama- Niacin 1.4 mg 1.0 mg
ranth could be increased. As per their view, ‘ama- Ascorbic acid 80 mg 30 m
ranth flour contributes to the sweetness and Thiamin 0.08 mg 0.14 mg
moistness of a baked good’. In a number of Ash 2.6 g 2.6 mg
African nations, amaranth is gaining prominence Calcium 267 g 490 g
as an important nutritious food especially for Compiled from J. N. Cole Amaranth: from the past, for
those suffering from HIV/AIDS. It is known that the future, Rodale Press, Emmus, PA, USA (1979)
3.10 Bioactive Components and Medicinal Properties 39
3.9 Culinary Properties The health beneficial antioxidant activities are

of Amaranths related to their bioactive components.
The word culinary means ‘related to cooking’; it

is the art of the preparation, cooking and presen- 3.10 Bioactive Components
tation of food, usually in the form of meals. and Medicinal Properties
People working in this field are also known as
‘culinary artist’ and ‘culinarian’. It is widely accepted that vegetables play an
Amaranth is one of the most commonly con- important role in preventing the development of
sumed green leafy vegetables in Asia and cardiovascular diseases, ageing-related diseases,
Southeast Asia. Most people eat vegetables with- obesity and cancers and improving human mem-
out getting the proper amount of nutrients from ory (Wayne et al. 2000). The health-promoting
them. This is primarily due to the differences in effects of vegetables are attributed to their natural
the methods of preparation. It should be cooked dietary antioxidants. Dietary antioxidants prevent
in a proper way so that all the nutrients remain free radicals related to ageing such as reactive
preserved. A few precautions should be adopted oxygen species in the human body (Nilsson et al.
to reduce the cooking-related losses. Amaranth 2004). The free radical theory of ageing involves
leaves should be consumed rapidly and washed cumulative damage through natural free radical
quickly in freshwater before cutting. While cook- oxidative changes, which over time results in
ing, steaming or stewing is advisable rather than increasing antigenicity, protein changes and oxi-
boiling or blanching. The water produced from dative DNA damages (Edelstein and Sharlin
steaming should not be discarded, but instead 2009). Vitamin C is a powerful antioxidant con-
used in cooking to make soup or sauce. Generally, tributing to the normal function of the immune
cooking for a long time or keeping vegetables hot system (Melvin 2010). Polyphenolic compounds
for a long time can destroy all the vitamins. have most antioxidant function acting as electron
During cooking all these parameters should be donors, electron acceptors, decomposer of perox-
kept in mind to promote nutrition and communi- ides and hydroperoxides, metal activators and
cated among women (Funke 2011). deactivators and UV absorber (Svobadva et al.
Most vegetables including amaranth go 2003). Anthocyanin is demonstrated to have pow-
through cooking such as frying, simmering, boil- erful antioxidant properties against low-density
ing, steaming and blanching before consumption. lipoprotein oxidation to reduce the risk of the
Cooking has a significant effect on chemical coronary heart disease (Wallace 2011). The carot-
compositions such as bioactive components, anti- enoids are yellow to red pigment available in the
oxidant activities and physical characteristics in diet mostly as α-carotene, β-carotene, lutein,
terms of colour, texture and flavour. However, the cryptoxanthin, zeaxanthin and lycopene. The
effect can be either positive or negative depend- carotenoids are effective in scavenging reactive
ing on the processing methodologies, vegetable oxygen species or ROS formed in physiological
species and shapes (Bernhardt and Schich 2006). processes. Biological molecules such as DNA,
Amaranth is one of the commonly consumed protein, lipids and carbohydrates can be damaged
leafy vegetables in Guangdong Province, China. by ROS (Kumpulainen and Salonen 1999).
The vegetables are consumed after home cooking As a kind of vegetable, Amaranth is ranked as
procedures such as simmering, frying, boiling, one of the top five vegetables in antioxidant
steaming and blanching. Nevertheless, knowl- capacities (Walter 2001). It contains plenty of
edge about its bioactive components, for instance, bioactive components, such as L-ascorbic acid,
ascorbic acid, polyphenol, anthocyanins, carot- betacarotene, polyphenol, anthocyanins and
enoids and antioxidant activities, is limited. It is lutein (Walter 2001). It has been used as an anti-
not clear how home cooking affects these bioac- pyretic to reduce labour pain in Indian and
tive components and their antioxidant capacities. Nepalese traditional medicine, as astringent,
diuretic, haemorrhage and hepatoprotective agent cancer, Amaranthus paniculatus is the one of
(Kirtikar and Basu 1987). Amaranths have also them.
been used to treat bladder distress, piles, tooth- Several studies have shown that the possible
ache, blood disorders and dysentery (Madhav benefits of antioxidants from plant sources lie in
et al. 2008). The health beneficial antioxidant changing, reversing or forestalling the negative
activities are related to their bioactive compo- effects of oxidative stress. Antioxidant-rich foods
nents. Cooking had no deleterious effect on total or natural antioxidants may be used as effective
bioactive component except for the reduction of substances to prevent diseases of higher age
anthocyanin content. Home cooking increases (Paredes-Lopez et al. 2010). Adequate amounts
the antioxidant activities and the contents of of phenolics, flavonoids and ascorbic acid are
carotenoids, especially by steaming. Steaming found in the leaves of A. paniculatus (Linn.).
has a positive effect on the polyphenol and Intake of food containing high flavonoid content
L-ascorbic acid, which are degraded seriously is helpful to prevent diseases. The leaf extracts
after simmering procedure. Both simmering and are very rich in antioxidant phytochemicals like
blanching increased the betacarotene and lutein polyphenols, ascorbates and flavonoids which
in the cooked amaranth (Han and Xu 2014). have free radical scavenging potentials. Amaranth
A significant part of the current pharmacologi- leaves can be used as a rich source of natural anti-
cal research is devoted to anticancer drug design- oxidants with health protective potentials. Earlier
ing customised to fit new molecular targets. The studies have also identified strong antioxidant
identification of new antioxidants represents a and free radical scavenging activities of
highly emphasised research area, because antioxi- Amaranthus paniculatus seeds (Bhatia and Jain
dants have the capacity to reduce the risk of vari- 2003). Antioxidant-containing foods may exhibit
ous chronic diseases caused by ROS. Plants are the carcinogenic potential by modulating carcin-
some of the most attractive sources of new drugs ogen detoxification, by inhibiting lipid peroxida-
and have been proven effective in the treatment of tion or by enhancing antioxidant defence
a number of disorders. Plants contain a wide array mechanism (Davis and Kuttan 2001). The pre-
of natural antioxidants that might serve as a prom- liminary phytochemical studies indicated the
ising source for the development of new drugs presence of flavonoids, saponins and tannins in
(Badami et al. 2003). Secondary metabolites of the Amaranthus paniculatus leaf extract. Many
plant such as flavonoids, terpenes and alkaloids such compounds are known to have antitumor
have received considerable attention of research properties (Kintzios 2006). Any of these phyto-
in recent years due to their diverse pharmacologi- chemicals especially the flavonoids may have
cal properties including cytotoxic and cancer che- attributed to the cytotoxicity and antitumor prop-
mopreventive effects. The growing intention in erties of the Amaranthus paniculatus. The
the gradual substitution of synthetic food antioxi- extracts of Amaranthus paniculatus leaves pos-
dants by natural antioxidants and wide health sess chemopreventive potential and can therefore
implications of antioxidants in nutraceuticals has be exploited as antitumor agents.
accelerated the research on vegetable sources and The amazing health benefits of amaranths
the screening of raw materials like vegetable ama- have always been recognised by homoeopathic
ranths for identifying antioxidants. The antioxi- and Ayurvedic experts. Both the seeds and leaves
dants can terminate the oxidative damage of a of amaranth that are used as herbal remedies have
tissue indirectly by triggering the natural defences nutraceutical value. According to Manikandaselvi
of the cell and/or directly by scavenging the free and Nithya (2011), nutraceuticals are nonspecific
radical species (Xia et al. 2004). One of the best biological agents to promote wellness, prevent
approaches in search for anticancer agents from malignant processes and control symptoms. In
plant resources is the selection of plants based on Ayurvedic medicine amaranth seeds and leaves
their ethnomedical claims. Among the huge num- have been used effectively as an astringent to
ber of medicinal plants that are claimed to be anti- arrest diarrhoea, haemorrhagic problems like
3.10 Bioactive Components and Medicinal Properties 41
bloody stools and urine and excessive menstrua- internally for the treatment of internal bleeding,
tion. It is an excellent wash for skin problems diarrhoea and excessive menstrual flow and snake
such as acne and eczema to psoriasis and hives. It bites. Externally, it is used to treat mouth ulcers,
is used as a mouthwash for sore mouths, gums, nosebleeds and wounds. The plant can be used
teeth and throat and as an enema for colon inflam- fresh or it can also be harvested before com-
mation and rectal sores (Vietmeyer 1983). mencement of flowering and dried for later use.
Amaranth is used to fight against gastroenteritis, The root is emmenagogue and galactagogue; a
stomach flu and contraception. It can reduce tis- paste of the root is used in the treatment of men-
sue swelling from sprains and tick bites when orrhagia, gonorrhoea and eczema and helps to
applied externally, but it should not to be used by remove pus from boils. In Nepal the root juice is
pregnant or lactating women. The amaranth oil used for the treatment of fevers, urinary troubles,
has been shown to prevent and treat those affected diarrhoea and dysentery. The root juice can also
with hypertension and cardiovascular disease. be used to treat indigestion and vomiting due to
Regular consumption of amaranth can reduce eating of unusual foods and in combination with
cholesterol levels and lower blood pressure. root juice of Dichrophela integra, and Rubus
Amaranth has been found to boost the body’s ellipticus is often used to treat stomach disorders.
immune system. Amaranth leaves have been used A. blitum is used as stomachic, as good emollient
as a good home-made remedy for hair loss and also in the treatment of roundworm, in bilious-
premature greying. Application of the fresh leaf ness, haemorrhagic-diathesis and blisters.
juice of amaranth helps hair to retain its colour Another weed A. viridis is used as cooling, appe-
and softness. tiser, laxative, stomachic, alexiteric, and anti-
Amaranthus paniculatus is considered as the pyretic and is used in burning sensation,
world’s most nutritious plant. It is best for eye- hallucination, bronchitis, leprosy, piles, constipa-
related problems. The aqueous extract of leaves tion and leucorrhoea. The leaves are used as an
of Amaranthus paniculatus was detected to have emollient and the root as a warming agent and
radioprotective effect in whole body gamma radi- expectorant that help to lessen the menstrual flow
ation (Krishna and Kumar 2005). It can also be and treat leucorrhoea and leprosy. A fairly high
used to overcome the psychological stress-related concentration of potassium, copper and iron
problems, and this health benefit has been exam- makes it a potential nutraceutical suitable for for-
ined in stress-induced memory dysfunction prob- tification of foods. These plant organs might be
lem and found to improve learning after radiation evaluated as a significant supplement and rich
stress (Bhatia and Jain 2003). Amaranthus spino- source of dietary minerals in human food to fight
sus, a weedy amaranth, is used as cooling, laxa- various diseases. It is rich in ascorbic acid which
tive, diuretic, antipyretic, stomachic, astringent, is a known antioxidant. A popular vegetable A.
diaphoretic, emollient, febrifuge and galacta- tricolor is used as cooling, alexiteric, laxative,
gogue. It can improve appetite and reduce burn- stomachic, appetiser and antipyretic and also
ing sensation and is effective in treatment of used in burning sensation, leprosy, bronchitis,
hallucination, leprosy, piles, bronchitis, leucor- piles, hallucination, leucorrhoea and constipa-
rhoea, constipation and flatulence. Herb decoc- tion. Many compounds and extracts from ama-
tion can be used as a mouthwash for toothache. ranths showed antidiabetic, anti-hyperlipidemic,
The root is used as heating and expectorant and is spermatogenic, anti-cholesterolemic effects
useful in the treatment of leucorrhoea, leprosy (Sangameswaran and Jayakar 2008; Girija et al.
and eczema, considered specific in gonorrhoea. 2011), antioxidant and antimicrobial activity
Amaranthus spinosus Linn. leaves significantly (Alvarez-Jubete et al. 2010; Tironi and Anon
reduced aspirin-induced ulcer index. A gastric 2010).
anti-secretary effect is created by the leaves by A future promise of vegetable amaranths is
reducing gastric volume and acidity. The seed is the preparation of leaf-protein concentrates.
used as a poultice for broken bones. It is used Compared with most other species, amaranth leaf
protein is highly extractable in comparison with not valued for use and beauty, growing wild and
other species. In one experiment, amaranth had rank and regarded as cumbering the ground or
yielded the highest amount of extractable protein hindering the growth of superior vegetation.
among 24 plant species tried. During the extrac- Generally, a weed can be defined as a member of
tion of leaf protein, other nutrients are also undesirable, useless plant communities, thriving
extracted, for example, provitamin A (betacaro- in a habitat disturbed by human, possessing com-
tene), polyunsaturated lipids (linoleic acid) and petitive behaviour, capable of mass movement
iron. Heating or treating the extract with acid pre- from one place to another and can interfere with
cipitates the nutrients as leaf-protein concentrate the activities or welfare of human being.
and most of the harmful compounds are elimi- The genus Amaranthus not only includes crop
nated, as they remain in the soup. The green and ornamental species; it also includes many
cheeselike coagulum is washed with water and weedy species, known as pigweed. Pigweed is
slightly acidified with dilute acetic acid (vinegar) the common name applied for several closely
to further minimise the amounts of anti-nutritive related summer annuals that behave as major
factors if any. The prepared leaf-nutrient concen- weeds of the vegetable and row crops and
trate is especially useful for young children and acknowledged among most damaging weeds in
other persons requiring particularly high protein, agricultural field. They are highly competitive
vitamin A and iron. The fibrous pulp left after with crop plants, express high flexibility in
extraction of leaf-protein concentrate is a suitable response to environmental changes and adapt-
feed for animals. The protein quality of the ama- ability to diverse ecoclimate and ensure their
ranth leaf-nutrient concentrate in terms of amino existence by producing a huge number of seeds.
acid composition, digestibility and nutritional Pigweeds are the natural inhabitants in parts of
effectiveness is excellent. The quality is however North and Central America. Cultivation of crop
species dependent probably due to the presence and commercial germplasm exchange has opened
of secondary metabolites present in varying new niches, allowing pigweed to invade agricul-
amounts in some species (Carlsson 1982). tural ecosystem throughout the America and
Sanchez-Marroquin (1983) has given in detail parts of Africa, Europe, Asia and Australia.
agro-industrial uses of grain amaranths. A new Approximately ten Amaranthus species are rec-
food product called Amarlac and Amarmeal of ognised as weedy member. These are either mon-
high nutritional quality has been released in oecious (male and female flowers are present on
Guatemala by the Amaranth Food Company. It the same plant) or dioecious (have separate male
represents the first output of research towards and female plants) species. Monoecious category
processing and utilisation of amaranth grain and comprises redroot pigweed (A. retroflexus),
its conversion into high-quality human food par- smooth pigweed (A. hybridus), Powell amaranth
ticularly for young children and pregnant women (A. powellii), tumble pigweed (A. albus), pros-
(Amaranth news Letter 1987). These products trate pigweed (A. blitoides) and spiny amaranth
can be tried in India. (A. spinosus), and dioecious category includes
the common waterhemp (A. rudis), tall water-
hemp (A. tuberculatus), Palmer amaranth (A.
3.11 Weed Amaranths palmeri) and sandhills amaranth (A. arenicola)
(Fig. 3.18). Weedy Amaranthus species (pig-
Human interests, mostly in economic perspec- weeds) are regarded as the major problem in agri-
tives, determine whether a plant is a weed or not. culture and the notable weed A. retroflexus is
The weed can be defined in many ways, like any considered one of the world’s worst weeds.
plant that is objectionable and interferes with the Amaranthus retroflexus is thought to be a
activities or welfares of man (Anonymous 1994), native riverbank pioneer of the central and east-
a plant out of place or growing where it is not ern USA and adjacent regions of southeastern
wanted (Blatchley 1912) or a herbaceous plant, Canada and northeastern Mexico (Sauer 1967). It
3.11 Weed Amaranths 43
Fig. 3.18 Few notable pigweeds (a). A. retroflexus L. (b) A. tuberculatus (Moq.) J. D. Sauer (c) A. quitensis Kunth. (d)
A. palmeri S. Watson (e) A. powellii S. Watson (f) A. blitoides S. Watson (g) A. arenicola I.M. Johnston (h) A. albus L.
Fig. 3.18 (continued)
has become naturalised throughout the temperate rent in a short mucro. Number of stamens 5 and
regions of the Northern and Southern stigmas 2–3, flexuose or erect, 1 mm long.
Hemispheres. It is an annual herb, erect or with Capsules are subglobose, usually shorter than the
ascending branches, simple or branched (espe- perianth, circumscissile, with an indistinct neck,
cially from the base to about the middle of the rugose below the lid, 2 mm in length. Seeds are
stem). Stems are stout, sub-terete to angled and black, compressed, 1 mm in diameter, almost
densely covered with multicellular hairs. Leaves smooth centrally, faintly reticulate around the
are long-petiolate (petioles up to 6 cm, in robust margins (Fig. 3.18a)
plants not rarely equalling lamina), lamina ovate Spiny amaranth (A. spinosus) is a native of the
to rhomboid or oblong-ovate, with obtuse to subtropical Americas, but now it is available on most
acute, mucronulate tip, shortly cuneate or attenu- of the continents as an introduced noxious weed
ate base, multicellular hairs along the lower species. It can be a serious weed in the ricefield in
surface of the primary venation and often on the Asia. Spiny amaranth has spread through tropical
lower margins, 3–6 × 5-11 cm. Flowers are and subtropical latitudes around the world and
arranged in greenish or rarely somewhat pink- found in cultivated fields, waste places, road-
suffused, stout, axillary and terminal spikes, sides, garbage heaps and abandoned fields. It
which are usually shortly branched to give a grows both in wet or dry sites, but grows best
lobed appearance, more rarely with longer when soil moisture levels are below field capac-
branches. The terminal inflorescence is panicu- ity. It originated probably in tropical lowland of
late and very variable in size, with intermixed South and Central America and from their intro-
male and female flowers. Bracts and bracteoles duced into other warmer parts of the world.
are lanceolate-subulate, pale-membranous with a Amaranthus spinosus Linn. (sp. Pl. ed. 1: 991.
prominent green midrib excurrent into a stiff, 1753) is an annual erect or slightly decumbent
colourless arista, longer bracteoles subequalling herb, simple or much-branched and bushy, attain-
to twice as long as the tepals. Perianth have five ing a height of up to 1.5 m. Stems are stout,
tepals; tepals of the male flowers are lanceolate- sometimes reddish and usually branched, angu-
oblong, blunt to subacute,1.75–2.25 mm long; lar, glabrous or increasingly furnished above
those of the female flowers are narrowly oblong- (especially in the inflorescence) with long, multi-
spatulate to spatulate, obtuse or emarginate, cellular hairs. Leaves are long-petiolate (petioles
2–3 mm long, more or less green-vittae along the up to 9 cm, sometimes longer than the lamina),
midrib, which ceases below the apex or is excur- lamina ovate to rhomboid-ovate, elliptic,
lanceolate-oblong or lanceolate, with subacute or

more commonly blunt or retuse apex having a
distinct, fine, colourless mucro, entire margin and
cuneate or attenuate base, surface glabrous or
thinly pilose on the lower surface of the primary
nervation, having a dimension of 0.8–6 × 1.5–
12 cm. Each leaf-axil bears a pair of fine and
slender to stout and compressed spines up to
2.5 cm long. Flowers are green, in the lower part
of the plant arranged in axillary clusters,
6–15 mm in diameter; towards the ends of the
stem and branches, the clusters are leafless form-
ing simple or sometimes (especially the terminal)
branched spikes usually up to 15 cm long and
1 cm wide. Lower clusters are entirely female, as
are the lower flowers of the spikes, while upper
flowers of the spikes are male, mostly covering
the apical one-fourth to two-thirds of each spike.
Bracts and bracteoles are deltoid-ovate, pale-
membranous, with an erect, pale or reddish awn Fig. 3.19 General plant morphology of spiny amaranth
(Amaranthus spinosus L.)
formed of excurrent green midrib. Bracteoles are
shorter than, subequalling or little exceeding the
perianth, commonly smaller than the bracts. presence of this species in the Western North
Number of tepals is five; those of the female America, Eastern Asia, Australia and South
flowers are narrowly oblong or spatulate-oblong, Africa has been reported since the mid-1900s.
obtuse or acute, mucronulate, frequently with a Today, A. hybridus is distributed worldwide as a
greenish dorsal vita, 1.5–2.5 mm long, while weed of agricultural fields and other disturbed
those of the male flowers are broadly lanceolate habitats, and it ranks among the 18 most noxious
or lanceolate-oblong, acute or acuminate, with weeds in the world (Holm et al. 1991) (Fig. 3.20).
green midrib. Number of stamens 5 and stigmas Amaranthus hybridus L. is an annual, erect or
2–3, flexuose or reflexed, 1–1.5 mm in length. less commonly ascending herb, attaining a height
Capsule are ovoid-urceolate with a short inflated up to 2 m. not infrequently reddish-tinted
neck below the style base, 1.5 mm in length, reg- throughout. Stems are stout, branched, angular,
ularly or irregularly circumscissile or rarely inde- glabrous or thinly to moderately hairy with short
hiscent, the lid rugulose below the neck. Seed are or long multicellular hairs (increasingly so above,
black, shiny, compressed, 0.75–1 mm in diameter especially in the inflorescence). Leaves are long-
(Fig. 3.19). petiolate (petioles up to 15 cm but even then
As reported by Sauer (1967), A. hybridus orig- scarcely exceeding the lamina), lamina broadly
inated as a riverbank pioneer of Eastern North lanceolate to rhomboid or ovate elliptic, ovate or
America, with earlier range expanding through- oblong, with gradually narrowed to blunt or acute
out milder and moister regions to Mexico, Central to subacute mucronulate tip, attenuate or shortly
America and Northern South America. The earli- cuneate base, dimension 19–30 × 3–10 cm.
est record of the species in Europe dates back to Inflorescences are moderately developed in com-
approximately 300 years. The spread of A. hybri- parison with three grain species. Flowers are
dus in Europe took place primarily in the arranged in yellowish, green, reddish or purple
Mediterranean region. The spread of A. hybridus axillary and terminal spikes formed of cymose
was quite slow in comparison with other clusters, which are increasingly closely packed
Amaranthus weeds, especially A. retroflexus. The upward. The terminal inflorescence varies from a
Fig. 3.20 Worldwide distribution of Amaranthus hybridus L.
single spike to a broad, much-branched, panicle A. retroflexus, like A. hybridus and many other
up to 45 × 25 cm; male and female flowers are amaranths, is a riverbank pioneer. Its earliest dis-
intermingled throughout the spikes. Bracts are tribution expanded from the Central Eastern USA
deltoid-ovate to deltoid-lanceolate, pale- to adjacent Canada and Mexico. Linnaeus is
membranous, acuminate with a long, pale to blamed for introducing the weed in Europe,
reddish-tipped, erect arista formed by the stout, where the species quickly spread (Sauer 1967).
excurrent, yellow or greenish midrib, twice as By the early 1800s, the species became a com-
long as tepals, exceeding style branches. Tepals mon weed in the temperate regions of the Old
are five in number, lanceolate or oblong, acute- World. Today, the species is existing as an intro-
aristate or the inner ones sometimes blunt in the duced or naturalised weed worldwide, ranking
female flowers, shorter than utricle and slightly among the most widely distributed weeds of the
recurved, inner tepals are shorter than outer world (Holm et al. 1997). Initial distribution of A.
tepals. Number of stamens 5 and stigmas 2–3, powellii was recorded in canyons, desert washes
erect, flexuose or recurved, 0.75–1.25 mm in and other open habitats west of the Cordilleran
length. Fruits are subglobose to ovoid or ovoid- system of America, with wide gaps in wetter
urceolate, circumscissile utricle. 2–3 mm in regions of Central America. The earliest record of
length, tapering into towers at apex, with a mod- this species in Europe is found in German her-
erately distinct to obscure ‘neck’, smooth lid, barium specimens from the late 1800s and later
longitudinally sulcate, or sometimes rugulose introduced in Southern India and South Africa
below the neck. Seeds are black and shiny or which are evidenced by the samples of Southern
pale, compressed, 0.75–1.25 mm in diameter, India and South Africa. Migration of A. powellii
with spermoderm smooth centrally, faintly retic- to Eastern North America took place only during
ulate in the margins (Fig. 3.21). the last century.
Fig. 3.21 General plant morphology and inflorescence of Amaranthus hybridus L.
Amaranthus tuberculatus is an annual erect mentioned that actual hybridisation among these
herb that bears flowers during the summer or species may be underestimated due to the nature
fall. It has a long terminal inflorescences in the of morphological determinations based on char-
forms of linear spikes to panicles. Female flow- acter intermediacy, which is often diluted after a
ers generally don’t have any tepals, although few generations of backcrossing with the pre-
rarely one or two rudimentary tepals may be dominant genotype. More recent work applying
observed. Sauer (1972) separated Amaranthus molecular and morphological markers suggested
rudis (formerly Amaranthus tamariscina) as dis- the abolition of different species status and con-
tinct from A. tuberculatus, giving emphasis on sidered them to be one and the same (Pratt and
the features like dehiscence of utricle and Clark 2001). At present a single polymorphic
absence of female tepals. Sauer’s A. rudis was species, A. tuberculatus, has got recognition
first described in Oklahoma in the 1830s, and (Mosyakin and Robertson 2003). Costea and
since then it has shown continuous northward Tardif (2003a) however recognised the differ-
and eastward migration into midwestern states, ence between A. tuberculatus and A. rudis at the
overlapping with A. tuberculatus, distributed in variety level and introduced to varieties: A.
the sandy and muddy streambanks, lakeshores tuberculatus var. rudis with more weedy tenden-
and pond margins, along the Missouri, cies than A. tuberculatus var. tuberculatus. Over
Mississippi and Ohio River systems (Sauer the last 20 years, A. tuberculatus has appeared as
1957, 1972), where both A. tuberculatus and A. the most significant weed problem in the mid-
rudis coexisted. The former was recorded on western USA (Steckel 2007), one of the world’s
average 40 years prior to that of the later. Many premier agricultural regions. The success
of the samples collected from these areas were achieved by A. tuberculatus to become a notori-
classified as putative hybrids between A. tuber- ous weed can be attributed mainly to its remark-
culatus and A. rudis, with a higher ratio of able ability of herbicide resistance. Studies on
hybrids to non-hybrids in artificial habitats com- the herbicide resistance of A. tuberculatus are
pared to natural settings. According to Sauer’s discussed in detail in a later section. The capac-
evaluation of dioecious amaranths (1957), A. ity of this species to respond to selection and
tuberculatus and A. rudis represent the most adaptive diversity identified may be of potential
abundant hybrid combination. The author also benefit to less orthodox crop-breeding pro-
gramme though from view point of weed man- variation may occur within a species which may
agement A. tuberculatus is of great concern. lead to occasional intraspecific hybridisation
The prime reasons for the noxious nature of (Sellers et al. 2003). Kansas State University
these weedy species lie in their ability to with- extension has published an excellent guide on pig-
stand and adapt efficiently to the modifications in weed identification with illustrations and a key to
agricultural management practices which are discriminate nine different weed amaranths (Horak
specifically designed to control and prevent weed et al. 1994). Monoecious species are primarily
colonisation. For example, numerous populations self-pollinating (Murray 1940; Weaver and
of pigweeds have evolved herbicide resistance McWilliams 1980), while dioecious species are
(Drzewiecki 2001; Rayburn et al. 2005). obligatory allogamous. Dioecism introduces
Correct identification of pigweed species may genetic diversity by encouraging outcrossing,
be controversial, especially during early vegetative while monoecism supports colonisation of new
phase because of their almost similar morphologi- areas by a single plant. Both the monoecious and
cal look (many look very similar). Significant dioecious pigweeds are highly successful invader.
Infrageneric Classification
of Amaranths 4
4.1 General 4.2 Synopsis of Infrageneric

Classifications of Amaranths
Genus Amaranthus represents a cosmopolitan
assemblage of species showing vast morphologi- The classification of Amaranthus is ambiguous
cal variability and wide geographical distribu- due to the absence of specific and qualitative
tion. Proper taxonomic delimitation of taxa is a species-defining characteristics, the extremely
prerequisite for it’s involvement in any breeding wide range of phenotypic variability among spe-
programme. In case of Amaranthus it is much cies as well as the introgression and hybridisation
more applicable due its taxonomic ambiguity and among weedy and crop species (Hauptli and Jain
presence of a large number of synonyms. Several 1978). There is no general agreement on taxon-
attempts have been made from time to time to omy of Amaranthus. At present the best option to
classify the genus infragenerically. The genus has be adopted is to understand the taxonomic prob-
been classified traditionally into three subgenera, lem of Amaranthus compiling the available
viz. subgenus Acnida (comprising dioecious informations in the literature and studying the
members), subgenus Amaranthus and subgenus phenotypic variability in a comprehensive man-
Albersia. Grain amaranths are included in subge- ner. This genus is nomenclaturally and taxonomi-
nus Amaranthus and vegetable amaranths are cally problematic both for its morphological
included in subgenus Albersia. Infrageneric clas- flexibility and frequent hybridisation. Sauer
sification in subgen. Amaranthus is to some (1967, 1976, 1993) made a taxonomic and geo-
extent satisfactory but, in case Albersia is far graphic survey on the grain amaranths and their
from conclusive, needs further studies. The works putative wild relatives employing morphological
done so far mostly employed morphological data. features and designated two subgenera – Acnida
But with the advent of molecular and biochemi- (which included the dioecious species) and
cal evidences, existing taxonomic delimitations Amaranthus (which included the monoecious
should be evaluated keeping in view phyloge- species). Later Mosyakin and Robertson (1996)
netic relationship. recognised three subgenera – subgen. Acnida

50 4 Infrageneric Classification of Amaranths
(Linnaeus 1753: 1027) Aellen ex K. R. Robertson 4.2.2 Amaranthus Subgen.

(Robertson 1981: 283), subgen. Amaranthus and Amaranthus
subgen. Albersia (Kunth 1838: 144) Grenier and
Godron (1856: 3) on the basis of the inflores- The subgenus Amaranthus consists of 20 species
cence and floral features. Traditionally, the sub- of annual monoecious herbs (Mosyakin and
gen. Amaranthus has been divided into two Robertson 2003). The species are mostly native
sections (Thellung 1919; Aellen 1959; Robertson to the Americas (Mosyakin and Robertson 2003).
1981) sect. Amaranthus [= sect. Amaranthotypus As the female and male flowers are arranged in
Dumortier (1827: 19)] and sect. Blitopsis Dumort. close proximity (Murray 1940), the monoecious
s.l. (1827: 19). Carretero (1985) further splitted amaranths are generally self-pollinating. Stems
the sect. Blitopsis into two groups: sect. Blitopsis are generally erect, and both the axillary and ter-
s.s., including species having indehiscent fruits minal inflorescences are arranged in cylindrical
and n = × = 17, and sect. Pyxidium Moq. in De spikes or panicles (Mosyakin and Robertson
Candolle (1849: 262) comprising species with 2003). Attempts to discriminate or differentiate
dehiscent fruits and n = × = 16. Recently Mosyakin taxa within this group faced much difficulty
and Robertson (1996) recognised four sections because during taxonomic delimitation, empha-
under the subgen. Albersia, of which three sec- sis was given on pigmentation and growth forms
tions [sect. Blitopsis, sect. Pentamorion (Beck which are extremely variable within amaranths
1909: 24) Mosyakin & K. R. Robertson (1996: (Sauer 1967). However, examination of floral
280) and sect. Goerziella (Urban 1924: 301) parts can provide a constant set of characters
Mosyakin & K. R. Robertson (1996: 280)] from which discontinuities can be explored to
include species with indehiscent fruit and remain- define taxa. In this regard, tepal number and mor-
ing one (sect. Pyxidium) includes species having phology are significant in preparing taxonomic
dehiscent, circumscissile utricle. keys. Amaranthus hybridus is a fundamental and
basal species in this crop subgenus and conforms
an interbreeding complex involving two other
4.2.1 Amaranthus Subgen. Acnida Amaranthus weeds, A. retroflexus and A. powel-
Aellen ex K.R. Robertson lii. Partially fertile hybrid swarms between these
species can be found in the USA, in areas where
The subgenus Acnida includes nine dioecious their distributions overlap, and in Europe, where
species which are native to North America and all three species are recent immigrants.
don’t have any immediate phylogenetic affinity Amaranthus hypochondriacus, one of the three
with the amaranth crop. However, recent studies grain amaranths, is cultivated as an alternative
(Trucco et al. 2005a, b) have indicated that gene crop in North America and Asia. Initially the spe-
exchange might have occurred between cies was thought to have an Asian origin, but now
Amaranthus tuberculatus, an infamous member it is believed that distribution in Asia is secondary
of Acnida, and A. hybridus, a probable progenitor in nature, and the species is supposed to have
of domesticated grin amaranths. In fact, studies derived from A. powellii through domestication
on gene exchange between A. tuberculatus and A. in North America. Hybridisation has played a
hybridus provide interesting clues regarding significant role in the evolution of A. hypochon-
applicability of the genetic diversity and novel driacus, with several hybrid races cultivated by
gene pool of the dioecious taxon for crop American aborigines. Sauer (1967) identified
improvement. Moreover, as A. tuberculatus is stable hybrid cultivars derived from crosses pre-
increasingly appearing as a model organism for sumably between A. hypochondriacus and local
the study of weeds (Trucco et al. 2009), a wealth admixtures of A. cruentus – an A. hybridus
of genomic resources are being generated that domesticated form originating in Southern
may be used for programmes designed for the Mexico or Guatemala – and its progenitor. For
crops. instance, in the region of Los Reyes (Michoacan),
4.3 Infrageneric Classification After the Modification by Mosyakin and Robertson… 51
a cultivar is grown to make special ‘dark’ tamales indehiscence of the fruit is not an ultimate con-
which was a putative hybrid between A. hypo- clusive feature for segregation of infrageneric
chondriacus and A. hybridus. Likewise, a Warihio taxa.
Indian crop from Rancho Trigo (Chihuahua) was
identified as a hybrid between A. hybridus and A.
powellii. Another putative hybrid between A. 4.3 Infrageneric Classification
cruentus and A. hypochondriacus is cultivated in After the Modification
the Oaxaca region of Southern Mexico, and the by Mosyakin and Robertson
same crop is grown in small gardens in Madras, (1996)
India. Another important grain species A. cauda-
tus, the grain amaranth of South America, is pre- Recently Mosyakin and Robertson (1996) intro-
sumed to have originated through the duced four sections under the subgen. Albersia,
domestication of A. quitensis in the Andean of which three include species having indehis-
region (Sauer 1967). A. quitensis originally dis- cent fruits [sect. Blitopsis, sect. Pentamorion
tributed as riverbank pioneer of South America, (Beck 1909: 24) Mosyakin & K. R. Robertson
in the mountains of the North-West and at the (1996: 280) and sect. Goerziella (Urban 1924:
lower altitude of temperate south is a weedy 301) Mosyakin & K. R. Robertson (1996: 280)]
member of hybridus complex. Cultivation of A. and one (sect. Pyxidium) includes species with
quitensis type of plant with incipient domestica- dehiscent fruits.
tion can be found from Ecuador to Northern
Argentina, mainly for the production of pigments The Infrageneric Classification After the
needed for colouring of chicha and other maize Modification by Mosyakin and Robertson
dishes. The South American amaranths are not (1996)
supposed to hybridise readily with the North Genus Amaranthus L.
American members of this complex Subgen. Acnida (L.) Aellen ex K. R. Robertson
Sect. Acnida (L.) Mosyakin & K. R.
Robertson
4.2.3 Amaranthus Subgen. Albersia Sect. Saueranthus Mosyakin & K. R.
Grenier & Godron Robertson
Sect. Acanthochiton (Torrey) Mosyakin &
The subgenus Albersia includes all the species K. R. Robertson
traditionally included by many earlier authors in Subgenus Amaranthus
Amaranthus sect. Blitopsis sensu lato. It still Sect. Amaranthus
represents a rather polymorphic group. All the Subsect. Amaranthus
vegetable and weed Amaranthus except A. hybri- Subsect. Hybrida Mosyakin & K. R.
dus are included in subgenus Albersia. Carretero Robertson
(1985), following Moquin-Tandon (1849), Nothosect. Dubia Mosyakin & K. R.
favoured the separation of dehiscent-fruited spe- Robertson
cies in a section Pyxidium. But after this separa- Sect. Centrusa Griseb.
tion, the resulting infrageneric taxa of this Subgenus Albersia (Kunth) Gren. & Godr.
subgenus still appear to be too polymorphic and Sect. Blitopsis Dumort
widespread geographically to be considered as a Sect. Pentamorion (G. Beck) Mosyakin &
natural group. Taxonomic delimitation of vege- K. R. Robertson
table amaranths more particularly those included Sect. Goerziella (Urban) Mosyakin &
in sect. Pyxidium of subgen. Albersia (Mosyakin K. R. Robertson
and Robertson 1996) requires more study from a Sect. Pyxidium Mosyakin & K. R.
morphological point of view. Dehiscence or Robertson
Key for the Infrageneric Delimitation of broadly triangular, occasionally

Taxa abruptly narrowed into a protruding
• Plants normally dioecious ............................... arista or bristle formed of midvein
................... Subgen. Acnida ................... sect. Amaranthus sub-
– Leaves linear-lanceolate; margins of both sect. Amaranthus
bracts and leaves are crisped and minutely Tepals gradually narrowed into acute,
denticulate; pistillate flowers have very often spinulose or bristle-like tip
broad, deltate or rhombic-deltate folia- .... sect Amaranthus subsect.
ceous bracts............................................ Hybrida
...... subgen Acnida sect. Acanthochiton
– Both bracts and leaves with entire margin;
bracts normally elliptic to subulate, linear 4.3.1 Amaranthus Subgen. Acnida
not foliaceous: (L.) Aellen ex K. R. Robertson
Pistillate flower without tepal or rarely
with 1–2 reduced linear or linear- Acnida L. Sp. Pl. 1027. 1753. – Type: Acnida
lanceolate tepals less than 2.mm long; cannabina L. (= Amaranthus cannabinus (L.)
utricle generally indehiscent (dehiscent Sauer). The only species originally assigned by
only in A. rudis) ..................................... Linnaeus in the genus Acnida.
......... subgen Acnida sect. Acnida Acnida L. was formerly recognised as a sepa-
Pistillate flower with five (rarely four) rate genus by most of the authors until 1955. But
spathulate or obovate elliptic tepals, Sauer (1955) on the basis of convincing evi-
utricle generally dehiscent (indehiscent dences included it within Amaranthus. Sauer did
only in A. greggii) .................................. not provide any formal nomenclatural combina-
........ subgen Acnida sect. Saueranthus tion for this taxon either at the subgeneric or sec-
• Plants usually monoecious tional level. Aellen (1959) suggested the
– Plants ascending, prostrate or erect; tepals combination Amaranthus subgen. Acnida (L.)
usually 2–3, but in some species 4–5; in Aellen without any citation of basionym. Later
most cases inflorescence axillary, glomer- the combination was validated by Robertson
ate or short spiciform; in some species ter- (1981). This dioecious Amaranth represents a
minal inflorescence is also developed, but natural and morphologically distinct group origi-
in such cases utricles indehiscent or tepals nally endemic to North America.
less than five ............................................... The subgenus includes nine dioecious species,
.......... Amaranthus subgen. Albersia distributed into three sections each having a
– Plants normally erect, always with more or rather distinct combination of characters related
less long terminal, spiciform, cylindrical, to fruit (dehiscent or indehiscent), bract (folia-
inflorescence; tepals usually five, rarely ceous or not) and tepals of pistillate flower.
4–3, fruit transversely dehiscent (erect A.
bouchonii and some forms of A. spinosus) • Amaranthus subgen. Acnida sect. Acnida (L.)
........................................... Amaranthus Mosyakin & K. R. Robertson, comb. nov.
subgen. Amaranthus
Bracts modified into firm spines; fruit Acnida L. sect. Acnida are autonym created by
transversely or irregularly dehiscent publication of Acnida L. sect Montelia Moquin in
.................... Amaranthus sect. Centrusa DC, Prodr.13/2: 277. 1849 – Type: Acnida can-
Bracts not modified into spines; fruit dehisces nabina L. (= Amaranthus cannabinus (L.) Sauer).
transversely in the equatorial part (fruit This section includes dioecious species char-
indehiscent in A. bouchonii)............... acterised with greatly reduced tepals in pistillate
Amaranthus sect. Amaranthus flower and mostly indehiscent fruits. Three spe-
At least inner tepals spathulate, apex cies belong to this section, viz. Amaranthus can-
obtuse, truncate, emarginate or nabinus (L.) Sauer, Amaranthus australis
4.3 Infrageneric Classification After the Modification by Mosyakin and Robertson… 53
(A. Gray) Sauer and Amaranthus tuberculatus Sauer (often incorrectly mentioned as A. acan-
(Moquin) Sauer. The other two species thochiton (Torrey) Sauer. The plant possesses
Amaranthus rudis and Amaranthus floridamus several features uncommon in amaranths like –
show a morphological transition towards this sec- extremely broad, deltate, foliaceous bracts of
tion. Besides dehiscence of fruit, all other charac- female flowers and narrow lanceolate to linear
teristics of both the species like greatly reduced leaf lamina with crisped margin.
non-spathulate tepals, shape of inflorescence and
general habit, provide ample reasons for their
inclusion in section Acnida. 4.3.2 Amaranthus Subgen.
Sauer (1972) separated A. rudis (formerly A. Amaranthus
tamariscina) as distinct from A. tuberculatus, pri-
marily on the basis of utricle dehiscence and Type: Amaranthus caudatus L., a lectotype des-
absence of female tepals, though more recent ignated by Britton and Brown (1913) and sup-
work using molecular and morphological mark- ported by Hitchcock and Green (1929).
ers suggested the merger of both the species into The presence of massive terminal inflores-
a single polymorphic species A. tuberculatus cence and dehiscent utricle is not the absolute
(Pratt and Clark 2001). identifying feature for subgen. Amaranthus.
These features in combination with other features
• Amaranthus subgen. Acnida sect. Saueranthus should be considered. Mosyakin and Robertson
Mosyakin & K. R. Robertson, sect. nov. (1996) divided this subgenus into several sec-
tions like sect. Amaranthus and sect. Centrusa
Type: Amaranthus palmeri S. Watson Griseb. and one additional sect. nothosect. Dubia
This section is named after J. D. Sauer with Mosyakin & K. R. Robertson. Subsequent segre-
pioneering contribution on grain amaranths. gation of the sect. Amaranthus was recom-
Plants included in this section are dioecious. mended, two new subsects. were proposed, i.e.
Flowers have five (rarely four) tepals spathulate subsect. Amaranthus and subsect. Hybrida
in shape (occasionally only inner spathulate Mosyakin and Robertson subsect. nov. Proper
tepals), with rounded truncate or acuminate apex subsectional delimitation is yet to be achieved.
and usually dehiscent utricle. The section Placement of few species like A. brandegei
includes four species, viz. Amaranthus palmeri Standley, A. bigelowii Uline & Brey, A. viscidu-
S. Watson, Amaranthus watsoni Standley, lus Green, A. scariosus Bentham (Sauer 1967)
Amaranthus arenicola I. H. Johnson and and A. asplundii Thell. (Hunziker 1965) is ques-
Amaranthus greggii S. Watson. The species tionable and requires further study. Sauer (1967)
included in this section are characterised with included the Australian species A. pallidiflorus F.
five well-developed spathulate tepals in pistillate von Muell. closely resembling A. clementii
flowers and dehiscent fruit (except A. greggi hav- Domin in the section Amaranthus, but on the
ing indehiscent utricles). basis of floral morphology, it appears to be
closely related to another Australian species A.
• Amaranthus subgen. Acnida sect. mitchellii Bentham having axillary inflorescence
Acanthochiton (Torrey) Mosyakin & K. R. typical of subgen. Albersia (Kunth) Gren. &
Robertson, comb. nov. Godr. Overlapping of morphological features and
new findings through molecular approach regard-
Type: Acanthochiton wrightii Torrey (= ing phylogenetic affinity have emphasised the
Amaranthus acanthochiton Sauer) need of evaluation of this section Amaranthus
This is a monotypic section comprising the and supported the introduction of more new sub-
only one species Amaranthus acanthochiton sects. under sect. Amaranthus.
• Amaranthus subgen. Amaranthus sect. widespread in the tropics and subtropics of both
Amaranthus the hemisphere. It shows some degree of mor-
phological progress towards developing dioe-
Amaranthus sect. Amaranthus Dumort. cious habit.
(Florula Belgica : 19. 1827.)
Amaranthus sect. Euamaranthus Moquin in
DC, Prodr. 13. 2: 255. 1849 (excluding A. gange- 4.3.3 Amaranthus Subgen. Albersia
ticus L. and A. mangostanus L.)
This section is divided into two subsections: Type: Albersia blitum (L.) Kunth (= Amaranthus
blitum L.)
(1) Amaranthus subsect. Amaranthus compris- This subgenus represents a polymorphic
ing species like A. quitensis Kunth, A. cauda- group comprising all species conventionally
tus L., A. retroflexus L., A. celosioides included in Amaranthus sect. Blitopsis sensu lato
Kunth? and A. asplundii Thell. by many earlier authors. Relative closeness
(2) Amaranthus subsect. Hybrida Mosyakin & among the species included in this subgenus is
K. R. Robertson subsect. nov. less in comparison with subgenus Amaranthus
and subgenus Acnida. Further infrageneric
Type: Amaranthus hybridus L. delimitation in subgen. Albersia is very inconsis-
This subsection comprises species like A. tent and inconclusive as well. Following Moquin-
powellii S. Watson, A. hybridus L., A. hypochon- Tandon (1849), Carretero (1985) supported the
driacus L., A. cruentus L. and A. bouchonii Thell. segregation of section Pyxidium having dehiscent
fruit, but still this group represents a group too
• Amaranthus subgen. Amaranthus nothosect. polymorphic and geographically widespread to
Dubia Mosyakin & K. R. Robertson notho- be considered as natural. The feature like dehis-
sect. nov. (Amaranthus sect. Amaranthus x cence or indehiscence of fruit was not found to be
Amaranthus sect. Centrusa) a conclusive feature for delimiting infrageneric
taxa. Justifying the situation Mosyakin and
The allopolyploid species Amaranthus dubius Robertson (1996) correctly segregated the sub-
Mart. is very closely related with A. spinosus L. gen. into several narrow sections considering
(of sect. Centrusa) and members of sect. overall morphological characters:
Amaranthus. This stabilised allopolyploid spe-
cies are supposed to have evolved as a result of 1. Amaranthus subgen. Albersia sect. Blitopsis
natural hybridisation between A. spinosus and Dumort.
either A. hybridus or A. quitensis of sect. The section includes species having inde-
Amaranthus. hiscent utricles and trimerous flower such as
A. blitum, A. viridis, A. emarginatus, etc.
• Amaranthus subgen. Amaranthus sect. Study on seed coat sculpturing and anatomy
Centrusa Griseb. may demand further segregation of this sec-
tion into narrower subsections (Kowal 1954).
Type: Amaranthus spinosus L. (Fl. British 2. Amaranthus subgenus Albersia sect.
West Indian Islands 68; 1859) Pentamorion (G. Beck) Mosyakin and K. R.
Amaranthus sect. Acanthophora G. Beck in Robertson, comb. nov.
Reichenb Icon. Fl. Germanicae et helveticae 24: The species included in this section have
177. 1909. indehiscent utricle and generally five (occa-
Amaranthus spinosus is the only species sionally four) spathulate or at least distinctly
included in this section. It is probably a native to obovate tepals. Most of the species are native
South America. This polymorphic species is to South America and Australia.
4.4 Provisional Key to Some Edible Species of Amaranthus 55
3. Amaranthus subgen. Albersia sect. Goerziella be developed for proper taxonomic delimitation
(Urban) Mosyakin and K. R. Robertson, (Joshi 1981a, b). Infrageneric classification and
comb. nov. inclusion of various species in it should reflect
This monotypic section includes not only the species delimitation but also phylo-
Amaranthus minimus, the Cuban endemic genetic affinity between them. Keeping in view
species with peculiar floral morphology that the available information achieved through vari-
differs from all other taxa of Amaranthus. ous studies employing morphological and molec-
4. Amaranthus subgen. Albersia sect. Pyxidium ular parameters, infrageneric classification of all
Moquin in DC. the subgenera of Amaranthus especially subgen.
This section includes species with dehiscent utri- Albersia is to be evaluated.
cles like A. tricolor aggregates, A. capensis, A.
thunbergii, etc. It appears to be most ambigu-
ous grouping of taxa in the whole genus. It
traditionally includes monoecious species, but 4.4 Provisional Key to Some
recently Das (2014) described and identified a Edible Species of Amaranthus
gynomonoecious member named A. parga-
nensis from the Lower Gangetic Plain of West Feine-Dudley (1980) compiled the informations
Bengal, India, that closely resembles A. tri- from floras and monographs on Amaranthus rep-
color. This new species demand further subdi- resented as follows:
vision of this section into two subsections at
least comprising monoecious and gynomon- A. Flower unisexual
oecious members, respectively. The proposed B. Three tepals
classification does not appear to be conclusive C. Tepals equal to longer than utricle; utricle cir-
and ultimate, new taxa at section, or varietal cumscissile ........... 1. A. tricolor
level can be incorporated as per taxonomic C. Tepals shorter than utricle; utricle indehiscent
demand (Mosyakin and Robertson 1996). D. Utricle smooth ............. 2. A. blitum
The inclusion of grain amaranths in D. Utricle rugose ............... 3. A. viridis
Amaranthus subgen. Amaranthus along with B. Five tepals
their wild progenitors appears to be beyond any E. Tepals approximately equal in length and
confusion. But subcategorisation of cultivated incurved against utricle
grain and weed amaranths is a difficult job for the F. Plants armed; inflorescence with upper cymes
taxonomists that has been addressed by many staminate and lower cymes pistillate ..............
workers (Sauer 1967; Mosyakin and Robertson .............................. 4. A. spinosus
1996; Hanelt 1968). A comprehensive delimita- F. Plants unarmed; cymes with initial flower sta-
tion at subsection level is yet to be achieved. The minate reminder pistillate ...............................
genus as a whole is known to be taxonomically ..... 5. A. dubius
difficult. Moreover naturally occurring aneu- E. Inner tepals shorter than outer tepals, straight
ploids in the plant populations of the farmer’s or recurved from utricle
field (Joshi1981a, b) and frequent synonymy G. Bract exceeding style branches; inflorescence
have aggravated the taxonomic and nomencla- either short and thick or moderately devel-
tural ambiguity specially in the Himalayan oped; always dark seeded
region. Attempts towards classification and H. Tepals longer than utricle; inner tepals with
nomenclature have been made by Thellung apex obtuse or emarginate; utricle not forming
(1914), Standley (1917), Schinz (1934), Kowal tower at apex; inflorescence short and thick
(1954) and Sauer (1950, 1955, 1957). Keeping in .......................... 6. A. retroflexus
view the intermixing and segregate range among H. Tepals shorter than utricle; inner tepals with
all the three grain species, suitable criteria are to acute apex; utricle narrowing into tower at
apex; inflorescence moderately developed .... and new findings through molecular and bio-
7. A. hybridus chemical approach regarding phylogenetic affin-
G.Bract not exceeding style branches; inflores- ity have emphasised the need of evaluation of the
cence fully developed enormous (domesti- section Amaranthus and supported the introduc-
cated species) seeds usually light, sometimes tion of more new subsects under sect. Amaranthus.
dark Sauer’s key (Sauer 1967) and derived keys (Covas
I.Bracts equalling style branches; inflorescence 1992; William and Brenner 1995; Hendrickson
stiff; style branches forming sharp cleft at 1993; Sanchez-Del Pino et al. 1999) for separat-
base; tepals with acuminate apex ..... 8. A. ing species of grain amaranths could be very
hypochondriacus handy.
I.Branches shorter than style branches; inflores- The infrageneric classification of the genus
cence lax Amaranthus proposed by Mosyakin and
J.Utricle narrowing into tower at apex; style Robertson (1996) is well substantiated by mor-
branches erect; tepals with acute apex ............ phological and biomolecular data but challenged
....................... 9. A. cruentus by AFLP-based UPGMA analysis. AFLP-based
J.Utricle not forming tower; style branches UPGMA analysis (Wassom and Tranel 2005)
spreading meeting in saddle at base; tepals indicated that Sandhill amaranths (A. arenicola)
broad, often overlapping; inner tepals with was genetically more similar to waterhemp (A.
obtuse apex ..... 10. A. caudatus tuberculatus) than to Palmer amaranths (A. palm-
eri) but differing from morphology-based taxon-
Much of the problems in taxonomic delimita- omy proposed by Mosyakin and Robertson
tion of Amaranthus species have been generated (1996). They grouped all the dioecious species in
from the attempts of recognising taxa on pigmen- the subgenus Acnida. This indicates the polyphy-
tation or growth forms, which are extremely vari- letic nature of the subgenus Acnida and the three-
able within amaranths. Delimitation of part classification of the genus Amaranthus. All
infrageneric taxa and placement of various grain the three grain amaranths and their wild progeni-
amaranth species along with their weedy rela- tor though included in the subgen. Amaranthus
tives as envisaged by Mosyakin and Robertson but their distribution in different sections are yet
(1996) were pioneering. Overlapping features to be ratified by biomolecular data.
Taxonomy and Phylogeny of Grain
Amaranths 5
5.1 General grain species are closely related, but A. hypo-

chondriacus and A. caudatus are more closely
Species interrelationship and phylogenetic link- related with each other than to A. cruentus. Size
age among different species especially grain of the chromosome has made the karyological
amaranths have been evaluated by many authors study in the genus very difficult. Three gametic
applying morphological, biochemical, molecular numbers have been reported in the genus (n = 14,
and cytogenetical parameters. Morphological 16 and 17). This tribasic nature of the genus has
evidences have played a significant role in solv- supposed to have originated from dysploidy or
ing taxonomic ambiguities in amaranths which aneuploidy. Information on distribution and vari-
are rich in morphological diversity. Besides con- ability of constitutive heterochromatin, the num-
ventional morphological features like inflores- ber of active ribosomal organiser region and
cence pattern and floral features, several other karyotype analysis are very significant to increase
features like morphology and anatomy of bracte- our knowledge about genetic variability and phy-
ole, seed surface architecture, phyllotaxy, course logenetic linkage among the species. Structural
of vascular supply and pollen morphology have alterations of chromosomes have played an
played a key role in differentiating taxa in many important role in the evolution of species. Wild
cases. Biochemical and molecular evidences species have comparatively more symmetrical
were used to validate the interpretations based on karyotype than the cultivated species. The differ-
morphological data. In most of the cases those entiation was caused by chromosomal repattern-
were concomitant with morphological data. ing, recombination and selection at subspecific
Electrophoresis of seed proteins; isozyme poly- level. These certainly have played a decisive role
morphism; different molecular markers like to their genetic evolutionary process. The grain
RAPD, AFLP, ITS, ISSR, etc.; leaf phenolic and vegetable amaranths are two distinct groups
chromatogram; single nucleotide polymorphisms that have evolved from their respective weed pro-
(SNPs); and tubulin-based polymorphism were genitor through unique domestication event in
applied to have a comprehensive idea about inter- different regions of the world. Two hypotheses
relationship of different species of grain, vegeta- have been proposed regarding the origin of grain
ble and weed amaranths. Most of the molecular amaranths from their wild weed progenitor –
techniques used yielded a common inference that monophyletic and polyphyletic. The monophy-
all the grain amaranths have evolved from weed letic hypothesis based on plant and seed
progenitor A. hybridus. Grain species showed morphology suggests that all three grain ama-
close proximity with weed progenitor A. hybri- ranths have originated from a single progenitor,
dus than with other weed species. The cultivated A. hybridus. The polyphyletic theory based on

58 5 Taxonomy and Phylogeny of Grain Amaranths
phytogeography suggests that all the three grain within a species is very crucial to plant breeders
amaranths have evolved independently. A third because it is an important consideration espe-
hypothesis suggests that all the three grain ama- cially when germplasm is to be selected for
ranths have originated from genetically differen- involvement in a breeding programme.
tiated population of A. hybridus through Morphology contributed a significant part of
independent domestication event. The validity of characters used in taxonomic delimitation in
these hypotheses has been challenged due to lack genus Amaranthus. Grain amaranths are charac-
of adequate sampling of all grain amaranths and terised with few salient features like apical, large
putative weedy progenitors, and modified ver- to moderately large complex inflorescence com-
sions have been proposed. prising aggregates of cymes, unisexual flowers
with five tepals, five stamens, circumscissile utri-
cle, seeds with variable seed coat colour (pale
5.2 Morphological Approach ivory, pinkish, brownish other than black and
in Solving Taxonomic brownish black) and well-defined flange. Due to
Disputes variability in morphological features, accurate
identification of amaranth genetic resources is
Genetic diversity is defined as the variation of not always possible (Transue et al. 1994). For
individual genotypes within and among species. preliminary identification of Amaranthus spe-
It is an important trait for long-term survival of cies, the number, thickness, orientation and den-
species and enables a population to adapt to new sity of branches in inflorescences could be the
conditions brought by environmental change useful tool. The flowers are arranged in small,
(Hamrick and Godf 1996). Genetic variabilities very contracted cymes, and the cymes are
are estimated from experimental data derived agglomerated, axillary and also additionally
from morphological, cytological, biochemical arranged in racemose or spiciform terminal, large
and molecular traits. Application of morphologi- and complex synflorescences. In spite of extreme
cal and cytological trait to estimate genetic vari- variability in inflorescence, there is usually a ten-
ability has the demerits of being influenced by dency towards a morphological ‘type’ (Costea
both environmental and genetic factors. Therefore et al. 2001a). The inflorescence structure of
they may not reflect accurate result (Basu et al. Amaranthus spp. is very complicated and has
2004). The genetic profile of the whole popula- been described by Murray (1940), Weaver and
tions typically varies from place to place across a McWilliams (1980) and Costea et al. (2001a).
species range. These differences may arise due to Flowers are small, green and unisexual and
chance occurrences, such as the genetic composi- develop in numerous dense clusters (dichasial
tion of dispersing individuals that create a new cymes). Each cyme has a determinate main axis
population (founder effect), or changes in allele with a terminal male flower followed by pairs of
frequencies that result from chance matings in opposite, or occasionally single, lateral branches
very small populations (genetic drift). Differences of female flowers (Fig. 5.1a). Each flower is sub-
among the populations may also arise systemati- tended by 1–2 spinescent bracteoles, which are
cally, especially when the individuals are exposed responsible for the overall increases in the den-
in the environments of various places offering sity of the inflorescence (Costea and Tardif
different optima for survival and reproduction 2003c). The cymes are further arranged in numer-
(fitness). For these and other reasons, populations ous spikes (spiciform branches) which grow
often show divergence from each other in their acropetally by the addition of new cymes.
genetic composition. Such divergence is espe- Towards the end of the inflorescence branches,
cially strong and rapid when gene flow among there are several newly developed male flowers
populations is limited (e.g. limited dispersal of that can pollinate female flowers of the lower
seeds or pollen or limited movement of animals cymes. The formation of new male flowers
across physiographic barriers). Knowledge of the decreases and ceases towards the end of the
amount and distribution of genetic variability growing season. Wide morphological variability
5.2 Morphological Approach in Solving Taxonomic Disputes 59
Fig. 5.1a Inflorescence pattern in amaranths. (a) Half glomerule of A. caudatus, (b) half glomerule of A. hypochon-
driacus and (c) full glomerule of A. tricolor (After Mohinder Pal 1972)
though with little taxonomic implications is while spinulose bracteoles are considered as syn-
shown by many Amaranthus species in response apomorphic. The structure of bracteoles in the
to several environmental factors like nutritional grain amaranths was found to be a reliable char-
elements, water availability, light conditions, acter that separates them from their weed rela-
injurious factors, etc. (Costea et al. 2001a, b). tives (Costea and Tardiff 2003b).
The ratio of male/female flower in inflorescence Morphotypes of A. caudatus are generally
is very important in reproductive behaviour of the characterised by a distinctive apical, soft, dense,
species. As per estimation, the ratio of male/ much branched thyrsoid long pendulous
female flowers in the inflorescence of A. powellii spike, like elephant trunk, mostly pinkish seed.
was 7.6 % (43 male to 524 female) and 9.7 % for Morphotypes of A. cruentus are characterised by
A. hybridus (26 male to 241 female). The female an apical massive, lax, complex much branched,
contribution to total reproductive effort, in terms erect reddish, orange, pale-yellow inflorescence
of floral biomass, was estimated to be 98.4 % for and brown or greyish-white seeds. The weed
A. powellii and 95.6 % for A. hybridus (Lemen amaranths generally have small black seeds with
1980). All monoecious species of Amaranthus undifferentiated flange. Wu et al. (2000) in a
are self-compatible and probably self-pollinating study on 229 genotypes from 20 Amaranthus
(Brenner et al. 2000; Costea et al. 2001a, b). The species observed wide variability which was use-
morphology and anatomy of bracteoles were sur- ful for improvement of cultivar in agronomic
veyed in 20 Amaranthus taxa in order to deter- traits, such as plant height, seed, stem and leaf
mine their taxonomic significance. Three types of colour among genotypes within the same species
bracteoles were distinguished: spinulose, folia- and among different Amaranthus species. Similar
ceous and membranous. Foliaceous and membra- results were also observed (Xiao et al. 2000;
nous bracteoles are considered symplesiomorphic, Varalakshmi 2004) in the evaluation of different
accessions of vegetable Amaranthus. Qualitative physiological and genetic integrity of nineteen

characters have been used for plant description. amaranth species during storage. It was found that
They are also useful in separating varieties espe- amaranth seeds start to lose genetic integrity when
cially when the range of quantitative characters is germination capacity is below 40 %. For best per-
limited (Ghafoor and Ahmadt 2003). formance of amaranth seeds in field storage under
The seeds of amaranths were described mor- ambient condition should not exceed 3 months.
phologically by many workers (Kowal 1954; Phyllotaxy and leaf vasculature may be useful
Klopper and Robel 1989; Costea 1997). The seed for species delimitation in amaranths. The phyl-
coat colour is more constant in vegetable and lotaxy and nature of vascular supply were previ-
weed amaranths ranging from black to blackish ously studied in A. caudatus (Gravis and
brown. But the seed coat colours in the grain spe- Constantinesco 1907), A. graecizans L. and A.
cies are more variable, may be dark or light hybridus L. (Wilson 1924). Costea and DeMason
coloured. Micromorphological features of seed (2001) extended this type of study to other com-
surface are proved useful in identifying species mon species. The results proved that a new set of
even accessions of Amaranthus. The seeds of the characters including morphology, epidermal
grain amaranths are quite different from the other characters, phyllotaxy, complexity of leaf vascu-
Amaranthus species in having well-differentiated lar supply and relative amount of secondary
folded flange, ill-defined or well-defined reticula- growth may be of taxonomic value that supported
tion of ridges over spermoderm and presence of the segregation within the ‘hybridus complex’,
verucate processes. Vegetable and weed ama- i.e. separation of cultivated grain amaranths (A.
ranths showed some commonality in seed surface caudatus, A. cruentus and A. hypochondriacus)
features. Both have spermoderm with reticulation from their presumed wild ancestors.
of elevated polygonal areas and are lacking well- Pollen grains of Amaranthus are tectate panto-
defined ridges (Fig. 5.1b, 5.1c, and 5.1d). A large porate, generally with more than 18 sunken pores
number of accessions of Amaranthus cruentus and supratectal granules and spinules (Eliasson
were studied that showed prominent variability in 1988; Costea 1998a, b). This Amaranthus type of
leaf shape, pattern of inflorescence, pigmentation pollen grains (Erdtman 1952, 1966) is also found
and most significantly micromorphology of seed to be present in the other members of
surface that demands for an evaluation and modi- Amaranthaceae as well as in several other
fication of existing infraspecific classification. Centrospermous families (Nowicke 1993).
Amaranthus cruentus have greyish-white or Koracev (1969) reported that pollen diameter in
brown-coloured seeds. All the greyish-white- the ten species of Amaranthus varies from 19.8 to
seeded accessions have rugulate spermoderm 31.4 μ, and the number of pores in a pollen grain
with irregular muriform arrangement of ‘rugae’. varies from 30 to 51, pore size from 2.2 to 5.5 μ
But brown-seeded accessions of A. cruentus have and the distance between the pores 5.5 to 9.6 μ.
reticulate spermoderm with reticulation of prom- Relationship between monoecious and dioecious
inent ridges forming hexagonal or polygonal amaranths as well as between members belong-
cavities. These seed surface features favoured the ing to different ploidy level and interspecific
division of A. cruentus population into two vari- hybrids can be estimated applying pollen grain
eties – Amaranthus cruentus (L.) var. albus features. Pollen grains of dioecious species have
S. Das, var. nov. (with rugulate spermoderm) and a greater number of apertures on the visible sur-
Amaranthus cruentus (L.) var. cruentus (with face (Franssen et al. 2001a). The pollen grains of
reticulate spermoderm showing reticulation of species belonging to lower ploidy level have
honeycomb-like cavities) (Das 2012b) (Figs. comparatively narrow-sized pollen, and the size
5.2a and 5.2b). ranges increase with increase in the ploidy level.
Longevity of seeds in storage condition is a The surface ornamentation appeared more prom-
good parameter of seed quality and vigour in inent in polyploids than in the diploid species
many crops. Kehinde et al. (2013) investigated (Chaturvedi et al. 1997).
Fig. 5.1b Micromorphology of entire seed surface of amaranth seeds by SEM (a) A. tricolor (b) A. blitum (c) A. cau-
datus (d) A. hypochondriacus (e) A. hybridus (f) A. retroflexus
Fig. 5.1c Micromorphology of micropylar region of amaranth seeds by SEM (a) A. tricolor (b) A. blitum (c) A. cau-
datus (d) A. hypochondriacus (e) A. hybridus (f) A. retroflexus
Fig. 5.1d Micromorphology of general seed surface of amaranth seeds by SEM (a) A. tricolor (b) A. blitum (c)
A. caudatus (d) A. hypochondriacus (e) A. hybridus (f) A. retroflexus
Fig. 5.2a Seed surface features delimited two varieties in A. cruentus L. (a). A. cruentus L. var. cruentus with reticulate
spermoderm, (b) A.cruentus L. var. albus S. Das with rugulate spermoderm
Fig. 5.2b Amaranthus

cruentus var. albus
S. Das (a) habit, (b)
portion of inflorescence,
(c) bract (d) bracteole,
(g) female flower
5.3 Biochemical and Molecular Approach in Understanding Systematics and Taxonomy 65
5.3 Biochemical and Molecular Zheleznov et al. (1997) also studied the variation
Approach in Understanding pattern in electrophoretic profile of seed proteins
Systematics and Taxonomy of wild and cultivated Amaranthus and identified
seven biotype groups which revealed the close
Several attempts have been made to explore the relationships between A. cruentus and A. hybri-
putative relationship of the grain amaranths with dus, similarity between A. caudatus and A. cau-
their probable ancestor/ancestors employing bio- datus var. edulis and distinctiveness of A.
chemical and molecular parameters like seed caudatus and A. cruentus. They reported that (1)
storage protein, isozyme polymorphism, pheno- the range of variation in seed protein content
lic chromatograms and molecular markers, viz. among both the wild and cultivated Amaranthus
RAPD, AFLP, ITS, SSR, ISSR, etc. Each and is rather wide; (2) Amaranthus seed proteins are
every parameters yielded data that were more or highly nutritive and, on the whole, consist of eas-
less concomitant not deviating from each other ily digestable albumins and globulins (more than
too much. Any observed variation in protein sys- 50 % of total protein), alkali-soluble proteins-
tems is considered as a mirror for genetic varia- glutelins, which are close to albumins and globu-
tions specifically seed proteins; they reflect the lins by their nutritive value (20.8 % of total
genetic history of the species without being protein), and alkali-soluble proteins-prolamines
affected by the environmental fluctuations. that are deficient in essential amino acids (12 %
Electrophoretic analysis of native or denatured of total protein); (3) the seed protein of
seed storage proteins was used to provide infor- Amaranthus species was heterogeneous in nature
mation concerning the genetic variability, which consisting of 38 bands as appeared through SDS-
represent a source of information for assessing PAGE with buffer pH 3.2. By decreasing electro-
genetic and taxonomic relationships at the spe- phoretic mobility, these bands were
cies level and below (Sammour et al. 2007; conventionally assigned to four zones; and (4)
El-Esawi 2008; Drzewiecki 2001). Stebbins the study of electrophoretic patterns of seed pro-
(1971) reported that polyacrylamide gel tech- teins is very promising to establish phylogenetic
niques provide clues to (1) explore interspecific relationship among the species of genus
or intraspecific variation, (2) screen a large num- Amaranthus. Electrophoretic characterisation of
ber of cultivars or accessions for genetic purity, seed proteins is rapid, relatively cheap method,
(3) verify whether or not two or more morpho- largely unaffected by the growth environment
logically identical accession in the germplasm (Juan et al. 2007; Jugran et al. 2010), and has
collection is also electrophoretically identical been applied by many workers to study the sys-
and (4) exploit the important traits of landraces tematics of Amaranthus spp. Drzewiecki (2001)
and wild relatives to facilitate increase in crop used SDS-PAGE of urea-soluble proteins of ama-
production and stabilise crop yield. The seed ranth seeds to differentiate both – species and
storage proteins of 44 taxa of Amaranthus spp. their cultivars. On the basis of similarity in pro-
were extracted in buffer solution and analysed on tein profile, samples of seven species were classi-
SDS-PAGE under reducing conditions. The result fied into three groups. By protein patterns A.
showed clear division in the studied amaranth tricolor (leafy type of Amaranthus) clearly dif-
taxa into two groups, one group containing basic fers from other species. The study suggested a
chromosome number x = 17 and the other group closer similarity between A. caudatus and A.
containing basic chromosome number x = 16 cruentus species than between the pairs of spe-
(Sammour 1991). This information clearly indi- cies A. hypochondriacus/A. caudatus and A.
cated the relation between the chromosome num- hypochondriacus/A. cruentus. Only slight differ-
ber and the electrophoretic pattern. The data on ences were seen among cultivars, especially of
seed storage protein also confirmed the separa- grain amaranths. According to Drzewiecki (2001)
tion of A. cruentus from A. hybridus and A. syl- crossing rate can be evaluated based on electro-
vestris and A. sylvestris from A. graecizans. pherogram of urea-soluble protein extracted from
single seed. On the basis of solubility, Osborne phoresis proved useful for genetic determination
(1907) divided proteins into four classes: albu- of A. retroflexus populations and identification of
mins soluble in water, globulins soluble in high- biotypes with a typical morphology.
salt concentration, prolamines soluble in aqueous Isozyme polymorphism analysis has been used
alcohol and glutelins soluble in acid or alkaline for over 60 years as useful parameter in biological
solutions (Segura-Nieto et al. 1994). This classi- research to establish phylogenetic relationships,
fication into four protein fractions has enabled to estimate genetic variability, identify cultivars and
differentiate various seed samples more clearly. genes and study population genetics and develop-
Gorinstein et al. (1991) and Drzewiecki et al. mental biology (Sammour et al. 2007; El-Esawi
(2003) generally characterised the protein frac- 2008; Rahman 2001). It was also utilised in plant
tion spectra of amaranth species. Dzunkova et al. genetic resource management, taxonomic delimi-
(2011) established the methodology for clear tation and plant breeding. Furthermore, isozyme
identification of the amaranth species using glu- analysis was used in control of breeding, estima-
telin protein fraction. The washing of water-, tion of outcrossing, testing purity and in species
salt- and alcohol-soluble proteins in protein frac- delimitation and conservation (Ar’us et. al. 1985;
tion separation process makes polymorphic peaks Becker et al. 1992; Chamberlain 1998). Finally
of amaranth glutelins to be distinguished very isozyme technique may be used by plant breeders
easily. SDS-PAGE analysis has been traditionally to generate, evaluate and select desired genotypes
used to analyse glutelin subunit composition of in early stage of the breeding programme, which
wheat, but the procedure is slow, laborious and saves time, money and efforts of the breeders
non-quantitative. The chip microfluidic technol- (Tanksley and Orton 1983). Thirty-four popula-
ogy based on capillary electrophoresis represents tion of New World amaranth along with 21 weedy
new approaches for the analysis of wheat HMW- New World populations were screened for allo-
GSs (high molecular weight glutenin subunits). zyme variability at electrophoretic loci using nine
This procedure is rapid and simple to apply and enzymatic system (Hauptli and Jain 1984). Eleven
enables automatic and immediate quantitative population of cultivated amaranth from Indian
interpretation. Other advantages over traditional state of UP and six population from Nepal were
gel electrophoresis method include lesser require- also considered in survey for comparison. In the
ment of sample and reagents and a reduced expo- New World populations, heterozygosity was low,
sure to hazardous chemicals (Bradova and and polymorphic loci ranged from 0 to 44 %.
Matejova 2008). The study of Janovská et al. Diversity index was partitioned into the intra- and
(2008) on the seed protein profiles of 15 interpopulation as well as the interspecific com-
Amaranthus accessions from the Czech Gene ponents of variability. The crop versus weed
Bank using both SDS-PAGE and chip electro- genetic distances was the largest, whereas the
phoretic profiles exhibited that (1) chip electro- intra- and interpopulation components of
phoretic technique is highly sensitive and Diversity index were found to be equal. Genetic
produces wider range of bands and (2) the structure of all the three grain species of the New
obtained data confirmed the classification of World amaranths together was described as a col-
Amaranthus species studied. The analysis of the lection of distinct populations, each represented
total seed proteins was very useful to assess the more or less a heterogeneous collection of highly
genetic differences between two grain popula- homozygous individuals. The North Indian popu-
tions of Amaranthus retroflexus collected from lations showed relatively less allozyme variability
field of the Maize Research Institute Zemun with the most common alleles same as those of
Polje, Serbia (Snezana et al. 2012). Two popula- Mexican landraces. Alleles at several loci proved
tions showed difference in protein profile; out of to be diagnostic of the crop and weed groups and
18 protein fractions obtained in protein analysis, of the three individual crop species. Genetic dis-
populations differed in four protein fractions of tances based on pooled gene frequencies showed
different molecular weight. Seed protein electro- three crop species to be generally more closely
related than they were to their putative weedy DNA product is present in one parent but absent
progenitor species, respectively (with the excep- in the other. For repositories with large collec-
tion of the weed-crop pair A. quitensis and A. cau- tions, this technique represents an important
datus). This consolidated the concept of single advancement towards detailed characterisation
domestication event where A. hybridus performed and screening of individual accessions at the
as the common ancestor, rather than three sepa- molecular level (Waycott and Fort 1994). RAPD
rate domestication events. Close similarity analysis is a powerful tool in determining inter-
between A. caudatus and A. quitensis might have as well as intraspecies genetic relationships
resulted from transdomestication based on a (Hardys et al. 1992). Such studies have been done
weedy or semi-domesticated species having involving wild and cultivated species (Ranade
migrated from Meso-America to South America. and Sane 1995), among self- and cross-pollinated
Genetic variability and species relationships of a species (Sharma et al. 1995) and even within
total of 23 species and 60 populations of culti- germplasms of a single-species population
vated and wild amaranths were studied using iso- (Pammi et al. 1994). Earlier, germplasms of grain
zyme marker (Chan 1996). High degree of amaranth accessions were analysed by RAPD
interspecific and intraspecific variations was (Virk et al. 1995). However, this study did not
observed between the investigated species and reflect interspecific relationship.
populations; 132 alleles were detected for 15 The RAPD technique has been successfully
enzyme systems. Total gene diversity for grain used for evaluating variation within plant acces-
amaranths and wild species was 0.39 and 0.72, sions and for establishing differences among
respectively. The polymorphism analysis demon- lines of apparently closely related populations in
strated the relationships of grain amaranths (A. germplasm collections, for example, American
caudatus, A. cruentus, A. hypochondriacus) with chestnut (Huang et al. 1998), barley (Russel
their putative ancestors (A. hybridus, A. powellii et al. 1997, Hong et al. 2001), Pinus longaeva
and A. quitensis), and the results also consolidated (Lee et al. 2002), strawberry (Zebrowska and
the idea about monophyletic origin of the grain Tyrka 2003), Trigonella (Dangi et al. 2004),
amaranths. In addition, the genetic diversity and Morus (Awasthi et al. 2004), Orobanche (Román
relationships of other species of amaranths were et al. 2007), Lactuca (Yoo and Jang 2003),
determined. Genetic diversity and relationships of Curcuma species (Syamkumar and Sasikumar
23 cultivated and wild Amaranthus species were 2007) and Amaranthus (Yoshimi et al. 2007).
examined using isozyme marker. A total of 30 loci Genetic diversity and relative closeness
encoding15 enzymes were resolved, and all were among six Amaranthus species collected from
polymorphic at the interspecific level. High levels eight phytogeographic regions were analysed
of inter-accessional genetic diversity were found using a random amplified polymorphic DNA
within species, but genetic uniformity was (RAPD) marker. RAPD primers yielded a total of
observed within most accessions (Chan and Sun 262 amplicons, ranging from 250 to 3000 bp of
1997). Iudina et al. (2005) examined the electro- which 254 amplicons (96.94 %) were polymor-
phoretic patterns of five isozyme systems in total phic. The genetic similarity coefficient among all
52 populations and two varieties (Cherginskii and the Amaranthus species ranged from 0.16 to 0.97
Valentina). Allozyme variation of these materials with a mean value coefficient of 0.56, which indi-
was low. Irrespective of species affiliation, 26 cated that variation exists in the genetic diversity
populations and two varieties were monomorphic of different populations (Yoshimi et al. 2007).
for five enzymes. RAPD marker polymorphism tend to support a
Genetic variability analysis with RAPD mark- closer genetic relationship between A. caudatus
ers is a fast, technically less complex, less expen- and A. hypochondriacus species (Avise and
sive methodology and involves no radioactivity Hamrick 1996), clustering of hybrids between A.
and hybridisation. RAPD markers are usually edulis and A. caudatus with A. caudatus and clus-
scored as dominant alleles, since the amplified tering of hybrids between A. hybridus and A.
hypochondriacus with A. hypochondriacus ment was conducted using eight decamer primers.
(Khoshoo and Pal 1972). The low genetic dis- After evaluation of analytical data on interspecific
tance values between these hybrids and other hybridisation and hybrid fertility, it was also con-
accessions of A. caudatus and A. hypochondria- cluded that these two (i.e. A. hypochondriacus
cus, respectively, indicated a weak genetic differ- and A. caudatus) are the most closely related pair
entiation between them. Chan and Sun (1997) in the grain Amaranthus species group (Grant
generated molecular progenies of cultivated and 1959b). From the similarity/dissimilarity percent
wild amaranth employing both isozyme and and subsequent clustering in dendrogram com-
RAPD marker. They evaluated 23 different culti- puted on RAPD data, it was reasonable to appre-
vated and wild Amaranthus species comprising hend that at least A. hypochondriacus and A.
the three cultivated grain species as well as acces- caudatus must have a common ancestor.
sions of all the species included in the A. hybridus Transue et al. (1994) applying RAPD tech-
aggregates. Fifteen isozyme systems encoded 30 niques also supported monophyletic origin of the
loci that showed polymorphism at the interspe- grain amaranths from A. hybridus as suggested by
cific level. Most accessions were monomorphic, Sauer. They classified 29 accessions of A. cauda-
but there was considerable diversity among the tus, A. cruentus and A. hypochondriacus into dis-
accessions as well. Out of 600 RAPD amplicons tinct groups and also classified and assigned 79
generated from 27 arbitrary ten-base RAPD prim- other accessions not previously assigned to spe-
ers, polymorphism percentage as revealed by cies (all assigned to A. cruentus or A. hypochon-
RAPD amplicons within cultivated grain species, driacus). The study showed that these three species
within progenitor species, within vegetable spe- could be classified unambiguously despite over-
cies and within other wild species were 39.9 %, lapping variations for morphological traits.
42.8 %, 51.0 % and 69.5 %, respectively. The evo- Analysis of 282 polymorphic RAPD markers
lutionary relationships between grain Amaranthus showed that A. hypochondriacus and A. caudatus
and their probable ancestors were investigated are more closely related to each other than their
utilising both the RAPD and isozyme data sets individual closeness with A. cruentus. RAPD anal-
that not only supported a monophyletic origin of ysis was also successful in the investigation to
grain amaranths and projected A. hybridus as the establish relationships between four A. hypochon-
common ancestor but also indicated a closer rela- driacus varieties (Barba de la Rosa et al. 2009).
tionship between A. hypochondriacus and A. cau- AFLP, ITS and ISSR markers were proved to be
datus. Dendrogram computed from RAPD and very efficient molecular parameters to study
isozyme date showed difference. Information genetic diversity and phylogenetic relationships
from both isozymes and RAPD when applied in a (Nolan et al. 2010), to assess the putative wild pro-
cumulative fashion appeared complementing genitor of grain Amaranthus (Costea et al. 2006;
each other and generated more accurate estimate Xu and Sun 2001) and to resolve taxonomic confu-
of genetic diversity and relative closeness within sion between the taxa of Amaranthus (Costea et al.
and among crop species and their wild relatives, 2006). The ‘Morelos’ accessions of Amaranthus
than data set when applied separately (Avise and from Mexico show taxonomic ambiguity at the
Hamrick 1996). This interpretation was ques- basic morphologic level. Although basic morpho-
tioned by the facts that the number of accession logical criteria are helpful for quick taxonomic
considered was limited and the wild species anal- delimitation and identification of specimens or
ysed were not from the area of their origin in germplasm collections, interpretation derived from
many cases. So, they were not the proper repre- morphological data alone often can be misleading.
sentatives of diversity. Mandal and Das (2002) To determine the taxonomic identity of the
demonstrated a high degree of genetic similarity ‘Morelos’ accessions and their speculative species
between A. hypochondriacus and A. caudatus affiliation to either Amaranthus caudatus or
based on RAPD analysis which supports earlier Amaranthus cruentus, Costea et al. (2006) con-
observations of Chan and Sun (1997). The experi- ducted a comparative analysis of evolutionary link-
age among these taxa/accessions applying basis of micromorphology and AFLP analysis
amplified fragment length polymorphism (AFLP) (Costea et al. 2006). Phylogenetic relationships of
and micromorphology methods. AFLP data helped grain amaranths with their wild progenitor and taxo-
to assign all the controversial ‘Morelos’ accessions, nomic disputes that exist among three cultivated
consistently and unambiguously into a single A. grain amaranths and their putative wild progenitors,
cruentus clade. This A. cruentus clade containing A. hybridus, A. quitensis and A. powellii, were eval-
‘Morelos’ accessions was clearly separated from A. uated using ITS, AFLP and ISSR (Xu and Sun
caudatus clade. This result clearly indicated that 2001). Phylogeny was poorly resolved due to low
the ‘Morelos’ accessions are affiliated to A. cruen- ITS divergence exhibited by gain amaranths and
tus. The AFLP-based phylogenetic relationship of their wild progenitor, though extensive polymor-
‘Morelos’ accessions and sharp delimitation of A. phism exists within and among the species studied
cruentus and A. caudatus are further consolidated both at AFLP and ISSR loci. In the cladogram com-
by micromorphology, proving the fact that the puted on either AFLP or ISSR or the combined data
combination of these techniques can provide more sets, nearly all intraspecific accessions could be clus-
reliable information for germplasm identification tered in their corresponding species clades, indicat-
than each method used separately. Qiang and Jin ing these taxa as well separated species. The AFLP
(2013) conducted a comprehensive investigation trees share many similarities with the ISSR trees,
applying 14 simple sequence repeat (SSR) to assess showing a close relationship between A. caudatus
the genetic variability and population structure of and A. quitensis, placing A. hybridus in the same
70 amaranth accessions collected from South and clade where all grain amaranths are included also
Southeast Asia. In total, 67 alleles were detected, indicating A. powellii as the most divergent taxon in
with an average of 4.79 per locus. A large portion the A. hybridus species complex. This study has also
of detected alleles (46.3 %) were raw alleles, while demonstrated that both AFLP and double-primer
29 unique alleles associated with rice accessions fluorescent ISSR have a great potential for develop-
were also detected. The mean major allele fre- ing a large number of informative characters to
quency (MAF), genetic diversity (GD) and poly- explore phylogeny of closely related species espe-
morphic information content (PIC) of the 14 SSR cially when ITS polymorphism is insufficient. Most
loci were 0.77, 0.36 and 0.34, respectively. A of the molecular techniques used to study the species
model-based structural analysis showed the pres- relationship in amaranths yielded a common infer-
ence of three subpopulations among the accessions ence regarding the origin of grain amaranths. The
studied. The genetic relationships configured by grain amaranths are more closely related with wild
the neighbour-joining tree method were fairly con- progenitor A. hybridus rather than with other wild
comitant with the structure-based membership species. The cultivated grain species are closely
assignments for most of the accessions. All 70 related, but A. hypochondriacus and A. caudatus are
accessions showed a clear relationship to each more closely related than to A. cruentus.
cluster without any admixtures. Qiang and Jin Though molecular studies do not always sup-
(2013) observed a relatively low extent of genetic port unanimous conclusion regarding the evolu-
exchange within or among amaranth species from tionary origin and proximity of crop-wild allies,
South and Southeast Asia. This result of genetic conventional studies considering hybrid fertility
diversity analysis could be very useful to identify and chromosome number tend to support the
amaranth germplasms and also to facilitate their hypothesis of independent domestication of grain
use for crop improvement. amaranths and close relationship between A.
Lanoue et al. (1996) studied restriction site varia- hypochondriacus and A. caudatus. Gupta and
tions in chloroplast DNA and nuclear ITS1 and ITS2 Gudu (1990) identified closer affinity between A.
region of 28 Amaranthus species. The data revealed caudatus and A. hypochondriacus applying
closer affinity between A. caudatus and A. cruentus hybridisation technique. Analysis of phenolic
than to their respective progenitor. Amaranthus chromatograms has been successfully used in
cruentus could be separated from A. caudatus on the solving taxonomic disputes (Bate-Smith and
Lerner 1954; Baker and Ollis 1961; McClure and polymorphism of acid phosphatase were applied
Alston 1966; Das and Mukherjee 1995) also as to get a comprehensive idea about relative close-
genetic marker to assess species affinity (Gornall ness among the species and morphotypes/acces-
and Bohm 1980). Das (2012a) successfully sions of both the vegetable, grain and weed
applied leaf phenolic compounds to assess the amaranths. Twenty-two plant specimens affiliat-
species relationship and taxonomic delimitation ing to ten species of Amaranthus were included in
in amaranths. Methanolic extracts of leaf pheno- the study (Das 2012a). The compiled chromato-
lics of different amaranth species were analysed gram (Fig. 5.4) on phenolic spots was generated
by thin-layer chromatography. Phenolic spot pro- incorporating all phenolic spots detected in indi-
files were viewed under UV light after spraying vidual chromatogram of each species, and such
flavone reagent (diphenyl boric acid ethanol- compiled chromatogram showed 21 spots. Few
amine, Sigma). Variability in the distribution of spots were exclusive for vegetable amaranths and
coloured spots proved to be very useful in evalu- few for grain amaranths. The zymogram of acid
ating species relationship (Figs. 5.3a and 5.3b). phosphatase showed 29 polymorphic bands. Not
Along with general morphological features, leaf only species, the morphotypes and varieties also
phenolics (secondary metabolites) and isozyme showed apparent variability in band profile. Both
Fig. 5.3a Leaf phenolic chromatography differentiating varieties and morphotypes of A. tricolor L.
Fig. 5.3b Leaf phenolic chromatography differentiating grain amaranths
Fig. 5.4 Compiled

phenolic chromatogram
showing total number of
phenolic spots found in
all the species studied
Similarity index
1.25 1.00 0.75 0.50 0.25 0.00
Amaranthus tricolor (Morphotype)

Amaranthus tricolor var. tricolor
Amaranthus tricolor var. tristis
Amaranthus tricolor var. acutus
Amaranthus tricolor (Morphotype) Group A
Amaranthus virisid
Amaranthus spinosus
Amaranthus dubius
Amaranthus graecizans
Amaranthus retroflexus
Amaranthus hybridus (Morphotype)
Amaranthus hypochondriscus (Morphotype)
Amaranthus hypochondriscus (Morphotype) Group B
Amaranthus caudatus (Morphotype)
Amaranthus cruentus (Morphotype)
Fig. 5.5 Dendrogram computed from morphological data
the vegetable and grain amaranths are supposed to and grain amaranths with their putative weed rela-
have originated from their respective weed pro- tives and also consolidated the concept of hybrid
genitor through occasional outcrossing and grad- origin of A. dubius from A. spinosus and A. hybri-
ual domestication. The study also strongly dus and allopolyploid nature of A. dubius as well.
vouched the previous conclusion regarding inter- Chan (1996) did not consider A. spinosus as one
relationships between grain amaranths, and their of the progenitors of A. dubius, according to him
probable derivation from their putative progenitor A. hybridus and A. hypochondriacus supposed to
also showed clear separate clustering of vegetable have undergone hybridisation and following dou-
and grain amaranths. A sharp congruence in clus- bling of chromosome gave rise to A. dubius. A.
ter pattern between morphological dendrogram dubius appears to be an autotetraploid of A. hybri-
(Fig. 5.5) and cumulative dendrogram computed dus and A. hypochondriacus. Stefu nova et al.
on morphological and biochemical data set (Fig. (2014) utilised ISSR primers to study the intra-
5.6) was apparent. Dendrogram on morphology and interspecific variability of a large number of
as well as consensus dendrogram revealed clear accessions of A. caudatus, A. cruentus and A.
separate clustering of accessions belonging to hypochondriacus. Highest intraspecific variabil-
three grain amaranths. Accessions of A. cruentus ity was shown by A. hypochondriacus in compari-
formed a distinct cluster, while accessions of A. son with A. caudatus and A. cruentus. It proved
hypochondriacus and A. caudatus revealed closer that ISSR markers were significant enough to
relationship keeping parity with previous obser- generate sufficient level of informative characters
vations. The dendrograms computed from data for intra- and interspecific molecular analysis of
also showed separate categorisation of vegetable genus Amaranthus.
Similarity index
1.25 1.00 0.75 0.50 0.25 0.00
Amaranthus tricolor var. tricolor
Amaranthus tricolor var. acutus
Amaranthus tricolor (Morphotype) Group A

Amaranthus viridis
Amaranthus spinosus
Amaranthus graecizns
Amaranthus retroflexus
Amaranthus dubius
Amaranthus hypochondriacus (Morphotype)
Amaranthus hypochondriacus (Morphotype) Group B

Fig. 5.6 Dendrogram computed from compiled date from all parameters
Many of the earlier molecular studies involv- enriched for the microsatellite motifs AAC, AAT
ing RAPD marker polymorphism yielded incon- and AC. Of these, 353 (24 %) contained unique
sistent results. Current studies are applying more microsatellites. An additional 29 microsatellite
reliable genetic marker like microsatellite loci were also identified among 728 BAC-end
marker and genome sequencing techniques to sequences of a newly developed amaranth BAC
address evolution-related ambiguities. Mallory library. Flanking primers were designed for 319
et al. (2008) applied microsatellite markers and of the microsatellite loci, and all were screened
genome sequencing to address evolutionary on a panel of diverse amaranths, including grain
linkage among grain amaranths and identified and weedy Amaranthus species. A total of 179
and characterised 179 microsatellite markers for (56 %) microsatellites were found polymorphic
amaranths. The objective of the study was to across accessions screened from the three grain
generate and characterise a set of highly infor- amaranths. Among these polymorphic microsat-
mative, reproducible microsatellite markers for ellite loci, a total of 731 alleles were identified
the three grain amaranths. A total of 1457 clones with average of four alleles per locus.
were sequenced from three genomic libraries Heterozygosity values ranged from 0.14 to 0.83
with a mean value of 0.62. Thirty-seven (21 %) called next-generation sequencing technique to
of the markers found were polymorphic between identify a huge number (27,658) of single nucle-
the parents of a segregating population and otide polymorphisms (SNPs) among four diverse
appeared to be inherited in a normal Mendelian amaranth accessions. Amaranthus hypochondri-
fashion proving the applicability of these mark- acus showed the highest total number of poly-
ers for linkage mapping of the amaranth genome. morphic SNP marker, while A. cruentus showed
Phylogenetic tree computed from microsatellite the lowest genetic diversity which may be indic-
marker data showed the placement of A. hybri- ative of the specialised domestication process,
dus accessions in two of the three major grain limited and uniform cultivation range (Mallory
amaranth clades, suggesting the polyphyletic et al. 2008). On the other hand A. hybridus, the
origin of the three cultivated species from differ- putative wild progenitor of grain amaranths,
ent A. hybridus ancestors. The microsatellite showed the highest genetic diversity of all the
markers have been proved to be very promising species studied which is explainable for its spec-
for further evaluation of the relative closeness ulated progenitor status. Significant SNP poly-
among the grain amaranths and their relatives morphism further consolidated the ancestral
also for use in marker-assisted breeding pro- relationship between the grain amaranths and A.
grammes, germplasm analysis and varietal iden- hybridus. One recently developed genotyping
tification. These markers may be useful to other method is tubulin-based polymorphism based on
species within the genus Amaranthus, including intron-length polymorphism available in plant
economically important weeds, vegetable ama- β-tubulin gene family (cTBP). The genes code
ranths and ornamentals due to the transferability for protein relevant for growth. The genomic
of these markers to A. hybridus, A. powellii and structure of these genes is such that a multiple
A. retroflexus. Microsatellite loci generated for approach can be adopted to study genetic
Amaranthus hypochondriacus were character- variability.
ised, and their cross amplification in wild species Bardini et al. (2004) developed the new tech-
was studied by Lee et al. (2008). Twelve poly- nique called tubulin-based polymorphism (TBP)
morphic microsatellite markers for Amaranthus based on the length of the first intron present in
hypochondriacus were isolated and character- the coding region of plant β-tubulin genes.
ised in the study. A total of 92 alleles were Introns of plant tubulin genes function as impor-
detected from the 20 accessions, with an average tant regulatory elements to support gene expres-
of 7.7 alleles per locus. The observed heterozy- sion. Introns can either increase the level of
gosity (HO) and expected heterozygosity (HE) expression through intron-mediated enhance-
values ranged from 0 to 0.95 and from 0.49 to ment, or they can change the actual site of gene
0.92, respectively. At significance threshold expression. Introns of tubulin genes are also use-
(P < 0.05), nine loci deviated from Hardy- ful for genotyping plant species and varieties, for
Weinberg equilibrium (HWE), whereas signifi- assessment of parental status and also for assist-
cant linkage disequilibria (LD) were observed ing breeding programmes (Braglia et al. 2010;
between five pairs of loci. Twelve microsatellite Breviario et al. 2007, 2008). Molecular markers
loci were successfully amplified in 18 other are now the most widely used parameter to
amaranth species representing cultivated grain assess genetic diversity (Karp et al. 1998;
and vegetable categories, their probable progeni- Koebner et al. 2001). TBP method was success-
tors and wild members. These results demon- fully applied on a wide range of species like oil-
strated great utility of these markers to study seed rape, coffee, Lotus (Bardini et al. 2004),
intra- and interspecific genetic variability as well Brassica, Eleusine and Arachis (Breviario et al.
as evolutionary relationships among cultivated 2007). Popa et al. (2010) used TBP method on
and wild amaranths. A significant progress was Amaranthus species for highlighting a possible
achieved when Maughan et al. (2010) utilised a polymorphism in the gene for β-tubulin. The
novel genomic reduction strategy linked with so- results revealed an intraspecific polymorphism
5.4 Cytogenetical Approach in Understanding Species Relationship 75
in A. cruentus (Alegria) V1-R1, while no poly- Amaranthus hypochondriacus is a native of

morphism was detected among the other studied North America where its closest wild relative A.
species. However, this low level of polymor- powellii is also existing. Sauer (1993) suggested
phism in these species may be indicative of high that it may be of hybrid origin from A. powellii
level of inbreeding in these Amaranthus species and A. cruentus. This relationship was also con-
or the fact that the primers which they have used solidated by some molecular information
were not successful enough in amplifying bands (Transue et al. 1994; Kirkpatrick 1995; Chan
in Amaranthus species under study (Bardini and Sun 1997). Thellung (1926) described a new
et al. 2004). Clear differentiation between two species A. bouchonii in Europe. Costea et al.
grain amaranths targeting a gene was achieved (2001a) considered it as a subspecies of A. pow-
by Park and Nishikawa (2012). They delimited elli, i.e. A. powellii subsp. bouchonii (Thell.)
A. caudatus and A. hypochondriacus from each Costea & Carretero. Though Sauer (1967) con-
other at species level by sequencing and PCR- sidered European taxon A. bouchonii to be con-
RFLP simple method of targeting two amaranth specific to A. powellii, a study on isozyme
starch synthase genes. polymorphism (Wilkins 1992) failed to establish
The versatile polymorphism and genetic vari- separate identity of A. bouchonii, but cytological
ability in amaranths especially in ‘hybridus com- evidences supported their distinctiveness. To
plex’ created many misapplications of names avoid confusion it would be better to consider
and created disputes between synonymy and both at subspecific level, i.e. Amaranthus powel-
species delimitation. Besides these major taxo- lii S. Watson subsp. powellii and Amaranthus
nomic problems involving grain amaranths, powellii subsp. bouchonii (Thell.) Costea &
there are few more diverse taxonomic disputes Carretero.
which are yet to be solved clearly like hybrid ori-
gin of A. dubius Mart., synonymy versus species
delimitation of A. quitensis Kunth and A. hybri- 5.4 Cytogenetical Approach
dus, conspecific nature of A. powellii S. Watson in Understanding Species
and A. bouchonii Thell., taxonomy of water- Relationship
hemp, etc. The widespread Northern races of A.
hybridus can be distinguished from other Karyotypical studies in Amaranthus are limited,
Amaranthus species. But tropical races are mor- probably due to the small size of the chromo-
phologically intermediate between A. hypochon- somes, which makes morphological analysis of
driacus, A. cruentus and other species (Sauer chromosome very difficult (Grant 1959a, b, c).
1950). Sauer (1950) misapplied the binomial Updated data have indicated that there are two
‘Amaranthus quitensis H.B.K.’ which should be basic chromosome numbers in Amaranthus,
the synonym of A. hybridus (Coons 1978). x = 16 and x = 17, and it is diploid except for a
According to Coons (1975) A. quitensis Kunth solitary tetraploid (2n = 64, x = 16) species, A.
as mentioned by Sauer (1950, 1967) is not a dubius Mart ex. Thell., and in some cases, both
valid biological species but should be classified numbers were mentioned for the same species
as Amaranthus hybridus var. sangorache which (Grant 1959a; Pal and Khoshoo 1972; Pal 1972;
unfortunately has not been published validly. Pal and Khoosho 1973a, b; Poggio 1988). Pal
However the validity of A. quitensis as a biologi- et al. (1982) suggested that the gametic number
cal species in Sauer’s sense is supported by n = 17 has originated from n = 16 through primary
molecular data (Chan and Sun 1997) and cytoge- trisomy. Greizerstein and Poggio (1992) sup-
netical observations (Greizerstein and Poggio ported this hypothesis analysing meiotic behav-
1992). Hybrids between these two species (A. iour of species and interspecific hybrids. The
hybridus and A. quitensis) yielded pollen grains existence of reproductive barrier during the
with 60 % viability. The data showed closeness crossing between Amaranthus species was sug-
at the same time divergence between these two. gested to be the contributory factor for variability
and suggested the possibilities of genic or chro- Table 5.1 Genome formulas for grain amaranths and
some other related wild species
mosomal differences causing sterility in hybrids.
Studies carried out on chromosome morphol- Genome
ogy of some species of Amaranthus have indi- Amaranthus species formulas N
cated variation in number of chromosome pairs Amaranthus caudatus A1A1B1B1 16
Amaranthus cruentus A2A2B2B2 17
with satellites. Palomino and Rubí (1991)
Amaranthus hypochondriacus A4A4B4B4 16
reported the karyotypic formula in some cultivars
Amaranthus mantegazzianus A3A3b3B3 16
of Amaranthus hypochondriacus and Amaranthus
Amaranthus quitensis AABB 16
cruentus, suggesting the existence of 6–10 pairs
Amaranthus hybridus A5A5B5B5 16
of chromosomes with satellites in different culti-
Amaranthus spinosus A6A6CC 17
vars. Greizerstein and Poggio (1994) proposed
Minor differences are expressed in the subscript for A and
karyotypic formulae of various accessions of cul- B genome. The genomes are x = 8 except for B2 and C,
tivated species (Amaranthus cruentus, which are x = 9 (Greizerstein and Poggio 1995)
Amaranthus hypochondriacus, Amaranthus man-
tegazzianus and Amaranthus caudatus). In all the
species studied, only one pair of chromosome hypochondriacus with 2n = 32 and A. mantegaz-
was found with a satellite (Greizerstein and zianus with 2n = 32 showed slight differences in
Poggio 1994). Kolano et al. (2001) reported the karyotype formulae and asymmetry index (Table
presence of one and two pairs of chromosomes 5.1). The variation in karyotype of different spe-
with ribosomal hybridisation signals for two cul- cies could be brought about by dysploidy, pericen-
tivars of Amaranthus caudatus applying FISH tric inversions, deletions, unequal translocation,
technique with 45 s ribosomal probes. etc. It suggests that the chromosome size is not
Basic chromosome number in Amaranthaceae constant, and it can vary according to selection
is x = 8 or 9 (Turner 1954). Three gametic numbers factor or adaptive norms. Restricting their studies
have been reported in the genus (n = 14, 16 & 17), on four species of Amaranthus, viz. A. viridis, A.
i.e. the genus is tribasic. In few cases n = 16 and 17 spinosus, A. blitum and A. tricolor, Srivastava and
occur in the same species. Amaranthus dubius is Roy (2012) concluded that by virtue of having
an exception, which is an allopolyploid with more symmetrical karyotype than other species, A.
2n = 64. The species might have originated as a spinosus is considered to be more primitive, and in
result of interspecific hybridisation between A. having asymmetrical karyotype, A.blitum is con-
spinosus L. and most probably A. hybridus. sidered more advanced.
According to Pal and Khoshoo (1982), the basic The genus Amaranthus is characterised by
chromosome number n = 17 has evolved from small chromosomes with indistinguishable sec-
n = 16 through primary trisomy. On the basis of ondary constrictions, satellites, etc. even at mitotic
cytogenetic analysis, Greizerstein and Poggio metaphase that act against a lucid and vivid work-
(1994) supported the above idea and proposed that ing out of its karyotype. This fact has so far
the species with somatic chromosome number restricted the cytogenetical studies of this crop to
2n = 32 are actually polyploid (having basic chro- base mainly on its male meiosis. However, to
mosome no. X = 8), and X1 is a derived basic num- explore the genetic system of any organism, a pre-
ber. The basic chromosome no. X2 = 17 might have cise knowledge of its karyotype is essential.
originated by primary trisomy. Genus Amaranthus Srivastava and Roy (2012) recorded the deviation
itself appears to be allotetraploid with chromo- in the karyotypic formulae of Amaranthus from
some count of n = 16 or 17 (Greizerstein and the previous count. Amaranthus spinosus (2n = 34)
Poggio 1994, 1995), and the genus behaves as dip- earlier showed the presence of one subtelocentric
loid during meiosis. They reported the karyotype and ten telocentric pairs of chromosomes
formulae and total DNA content of four species of (Greizerstein et al. 1997), but later study
Amaranthus. According to their observation A. (Srivastava and Roy 2012) showed presence of
caudatus with 2n = 32, A. cruentus with 2n = 34, A. only five subtelocentric pairs of chromosomes. It
is reported that A. viridis has two metacentric, 20 morphological and ecological condition may rep-
submetacentric, eight subtelocentric and two telo- resent most common cyto-evolutionary pattern in
centric chromosomes, and A. tricolor has six angiosperm. Amaranthus represents a genus that
metacentric, six submetacentric, 16 subtelocentric shows a great variability in chromosome number
and six telocentric chromosomes (Madhusoodanan and ploidy level (Table 5.2).
and Nazeer 1983). According to later study The distribution and variability of constitutive
(Srivastava and Roy 2012), A. viridis showed 14 heterochromatin and the number of active ribo-
metacentric, 12 submetacentric and eight subtelo- somal organiser regions were studied by Bonasora
centric chromosomes, and A. tricolor showed 14 et al. (2013) in two different cultivars of the spe-
metacentric, ten submetacentric and ten subtelo- cies Amaranthus cruentus (2n = 34), Amaranthus
centric chromosomes. No satellite was found to mantegazzianus (2n = 32), Amaranthus hypochon-
be present in any species. On the basis of above driacus (2n = 32) and Amaranthus caudatus
observation, it could be presumed that the karyo- (2n = 32). The aim of the study was to increase the
type differences are probably due to divergence knowledge about the genetic variability of genus
and frequent repatterning of chromosome over Amaranthus. The distribution and variability of
long periods of time and support the earlier reports constitutive heterochromatin were studied using
as analysed in different accessions (Greizerstein DAPI-CMA3 banding techniques (Bonasora et al.
et al. 1997; Madhusoodanan and Nazeer 1983). 2013). The position of the nucleolar organiser
Srivastava and Roy (2012) reported the chromo- (NOR) was observed using Ag-NOR banding and
some number in A. blitum (2n = 28), which is sim- fluorescent in situ hybridisation (rDNA-FISH).
ilar to the new basic chromosome number (x = 14), All the cultivated Amaranthus species showed
and this report was supported by observation of two active NOR regions except A. caudatus cv.
Pal et al. (2000). Pal et al. (2000) also reported EEA INTA Anguil. Furthermore the number of
chromosome number 2n = 28 in Amaranthus tenu- DAPI+/CMA3 bands allowed the characterisa-
ifolius. This consolidated the possibility of new tion and identification of heterochromatin in culti-
basic chromosome number (x = 14) for the genus vars and species.
Amaranthus, and genus could therefore be triba- Madhusoodanan and Nazeer (1983) con-
sic (x = 14, 16 and 17). ducted a comprehensive morphological and
There are several groups of angiosperm having karyotypical analysis of five species of sect.
variable chromosome numbers as a result of dys- Blitopsis, viz. A. tricolor, A. lividus, A. graeci-
ploidy or aneuploidy (Stebbins 1971). Amaranthus zans, A. viridis and A. albus, of which all the spe-
is one such group of plants. The term dysploidy cies except A. albus were collected from India.
indicates the process by which the euchromatin of The species of sect. Blitopsis are taxonomically
a genome is rearranged by translocation on to a well defined. Forms of A. tricolor are rather tall
greater or lesser number of centromeres, which is compared to other pot-herb species and their
evidenced in morphology of chromosome. It is wild relatives. In general for all the phenotypic
evidenced in the study of Srivastava and Roy characters studied, namely, height of the plant,
(2014) that there is whole chromosome loss at number of primary branches, leaf size, stomatal
diploid level in high number. It is assumed that size, flower size, etc., variation pattern is similar
dysploid reduction from relatively high chromo- for all the aspects indicating gigantic nature,
some number (2n = 34) has taken place in thereby better yielding ability of A. tricolor
Amaranthus due to repeated chromosome manip- compared to that of the other species. This may
ulation during hybridisation in between landraces. possibly be due to controlled evolution in A. tri-
This type of changes in chromosome numbers color resulting from recombination and selec-
represents a case of dysploidy rather than aneu- tion in which all the desirable qualities for
ploidy. Extensive dysploid reduction from high commercial growing are combined. In the mem-
primitive chromosome number to lower chromo- bers of other species, the morphological discon-
some number species associated with derived tinuity is more pronounced which can be
Table 5.2 The number of chromosomes and ploidy levels in some amaranths
Species No. of chromosomes (2n) Ploidy level
Amaranthus albus 32 or 34 Diploid
Amaranthus blitum 28 or 34 Diploid
Amaranthus tricolor 34 or 35a Diploid
Amaranthus viridis 34 Diploid
Amaranthus lividus 34 Diploid
Amaranthus aureus 34 Diploid
Amaranthus gangeticus 34 Diploid
Amaranthus hypochondriacus 32 or 34 Diploid
Amaranthus caudatus 30 or 32 Diploid
Amaranthus cruentus 32 or 34 Diploid
Amaranthus hybridus 32 Diploid
Amaranthus leucocarpus 32 Diploid
Amaranthus mangostanus 32 Diploid
Amaranthus mantegazzianus 32 Diploid
Amaranthus caturtus 64 Tetraploid
Amaranthus dubius 64 Tetraploid
Amaranthus edulis 32 Diploid
Amaranthus chlorostachys 32 Diploid
Amaranthus graecizans ssp. graecizans 34 Diploid
Amaranthus graecizans ssp. sylvestris 32 Diploid
Amaranthus giganteus 34 or 64 Diploid and tetraploid
Amaranthus retroflexus 32 Diploid
Amaranthus powellii 32 or 34 Diploid
Amaranthus quitensis 32 Diploid
Amaranthus paniculatus 32 or 34 Diploid
Amaranthus polygamous 34 or 68 Diploid and tetraploid
Amaranthus salicifolius 34 Diploid
Amaranthus spinosus 34 Diploid
Amaranthus sylvestris 32 Diploid
Amaranthus tenuifolius 28 Diploid
Amaranthus blitoides 32 Diploid
Amaranthus tuberculatus 32 Diploid
Amaranthus palmeri 32 or 34 Diploid
Amaranthus roxburghianus 34 Diploid
a
Representing an aberrant form
Reference: 1. Behera and Patnaik (1974), 2. Greizerstein and Poggio (1994), 3. Kulakow and Jain (1990a, b), 4.
Madhusoodanan and Nazeer (1983), 5. Mohideen and Irulappan (1993), 6. Sammour et al. (1993,) 7. Sauer (1976), 8.
Sreelathakumary and Peter (1993), 9. Pal et al. (2000), 10. Srivastava and Roy (2014), 11. Rayburn et al. (2005).
attributed to their limited usage as greens. The karyotypes of the species analysed are
Moreover, unlike the cultivated forms of A. tri- highly asymmetric because all the chromosomes
color, most of them are usually collected from of the basic set vary in size as well as in form as a
their wild habitat for use. They were found to be good number of them used to be subterminal and
quite distinct, differing in height, mode of terminal, while the others are submetacentric and
branching, shape as well as size of the leaf, inflo- metacentric. However, there was no correlation
rescence structure, nature of bract, utricle and between these two features in any of the species. It
dehiscence of utricle. is evident from the karyotype formulae that none
of the five species have identical chromosome evolutionary relationship (Ranade et al. 1997).
sets. As a whole, symmetrical karyotypes are usu- On the basis of karyotype analysis, Sharma and
ally considered as primitive and most asymmetri- Banik (1965) discussed species interrelation-
cal karyotypes as derived. Hence, the presence of ship in the family Amaranthaceae. Several
a large number of metacentric chromosomes and authors have demonstrated the existence of
its reasonably symmetric nature in A. albus is compatibility barrier to the interspecific cross
indicative of its primitiveness, while an increase in among Amaranthus species due to genic or
the number of telocentrics as observed in A. livi- chromosomal differences (Greizerstein and
dus shows that the genome has undergone consid- Poggio 1992; Pal and Khoshoo 1973a, b). It was
erable structural modifications. The karyotypic observed that wild species have more symmetri-
variation detected in different species may be due cal karyotype than the cultivated species. The
to pericentric inversions, deletions, unequal trans- karyotypic differentiation is caused by repat-
location, etc. In Amaranthus both chromosome terning of chromosomes, recombination and
size and gross morphology are not constant, and it selection at the subspecific level, and these are
varies being influenced by the selection factors important contributing factors to their genetic
and/or adaptive norms. The results obtained by evolutionary process (Srivastava and Roy 2012).
Madhusoodanan and Nazeer (1983) support the Interspecific hybridisation would be a confir-
earlier finding (Madhusoodanan 1978) that in the matory parameter for identification of
members of sect. Blitopsis, neither hybridisation Amaranthus specimens. Murray (1940) using
nor polyploidy seems to have played any role in controlled pollinations demonstrated that
their evolution, and species differentiation has interspecific hybridisation in Amaranthus is pos-
been brought about by chromosomal repatterning, sible. Interspecific hybridisation and transfer of
recombination and selection at the subspecific herbicide resistance in Amaranthus have been
level which are regarded as the most important demonstrated between two dioecious species, A.
factors contributing to their genetic evolution. palmeri and A. tuberculatus, and between A.
Sharma and Banik (1965) are also of the opinion tuberculatus and a monoecious species A. hybri-
that structural alterations of the chromosomes dus (Tranel et al. 2002; Wetzel et al. 1999b).
have played a very significant role in the evolution Interspecific hybridisation has also been reported
of the species. In the morphological and karyo- in wild Amaranthus populations. Grant (1959b)
typical analyses of five species of Amaranthus described the karyotypes of putative interspecies
sect. Blitopsis (Madhusoodanan and Nazeer 1983) amaranth hybrids. Hauptli and Jain (1984)
the species were found to be quite distinct, differ- described hybrids derived from interspecific
ing in height, mode of branching and shape as well crossing between cultivated grain amaranth (A.
as size of the leaf, structure of infloresence, nature caudatus) and redroot pigweed growing wild
of bract as well as utricle and mode of dehiscence near cultivated amaranth that showed intermedi-
of ripe utricle. Compared to the wild species, the ate morphology, but isozyme profile was more
cultivated ones showed advanced characters due similar to cultivated grain amaranth with a low
to selection. The karyotypes of all the species are frequency of redroot pigweed isozymes.
very much asymmetric with variation in size as Cultivated amaranths have relatively lower chi-
well as in form of chromosome sets. No two spe- asma frequency in comparison with wild species.
cies are karyotypically alike. Evidently, the spe- This phenomenon has attributed to repeated cycles
cies differentiation in members of the section is of hybridisation and selection in cultivation which
caused by chromosome repatterning, and in evolu- produced an inherent heterozygosity in the culti-
tion of the species, structural alterations of chro- vated amaranths (Madhusoodanan and Pal 1981).
mosomes are playing a key role. Cultivated species are characterised by more chi-
The trend in species relationship among the asma per bivalent than semiwild species like A.
amaranths as revealed by RAPD analysis is con- spinosus also with bigger chromosomes and pol-
sistent and supportive to their cytogenetic and len grains (Sreelathakumary and Peter 1993).
5.5 Taxonomic Delimitation 2013). The morphometric study recognised three

in Vegetable Amaranths varieties (i.e. var. tricolor, var. tristis and var.
acutus var.nov.) of which one (Amaranthus tri-
This group of plants are gradually feeding out of color var. acutus) was new for the science.
global crop directory, but it is true that at least 50 Amaranthus tricolor var. acutus differs from
tropical countries grow vegetable amaranth and other varieties in having ovate-oblong leaves
in quantities that are far from negligible. Nearly with emarginate apex, ovate-spathulate tepals
all the vegetable amaranths are included in the with acute apex and tepals as long as or shorter
subgenus Albersia. Recently four sections have than fruit (Fig. 5.7). Morphological characters
been recognised under subgen. Albersia – three delimiting three varieties of A. tricolor are men-
(sect. Blitopsis, sect. Pentamorion and sect. tioned in Table 5.3.
Goerziella) comprising members having indehis-
cent fruit and one (sect. Pyxidium) including
members having dehiscent fruit. Taxonomic 5.6 Key to the Varieties
delimitation and application of nomenclature in of Amaranthus tricolor L.
vegetable amaranths are still very tentative for its
morphological variability and frequent 1. Leaf ovate-oblong, apex emarginate, tepal
hybridisation. apex acute, as long as or shorter than fruit ......
Few species of Amaranthus are known as veg- ............................ A. tricolor var. acutus
etable amaranths like A. tricolor, A. blitum, A. 1. Leaf usually ovate-lanceolate or deltoid-ovate,
dubius, A. cruentus, etc., of which the most sig- tepal apex awned, tepal much longer than
nificant are A. tricolor and A. blitum. Amaranthus fruit(2)
tricolor is included in sect. Pyxidium and is a 2. Branching from base of the stem ....................
native species of tropical Asia. A. tricolor proba- .................. A. tricolor var. tristis
bly originated from weed progenitor in tropical 2. Branching from middle of the stem ................
Asia through outcrossing and domestication pro- ...................A. tricolor var. tricolor
cess. As a result several new taxa were described
at subspecies, variety and form ranks. A. tricolor Pan et al. (1992) studied 45 indigenous and
is the lectotype of the section Pyxidium and is a exotic genotypes of A. tricolor taking into con-
taxon of renewed taxonomic interest from the sideration ten quantitative traits like days to first
view point of misapplication of names, morpho- clipping, number of clippings, diameter of stem,
logical variability, as well as due to the presence length of internode, leaf-stem ratio, length of
of a large number of synonyms. Several authors lamina, width of lamina, days to flowering, dura-
(Mathai 1978; Mosyakin and Robertson 1996; tion of harvest and total yield. Variance analysis
Das 2013) recognised A. tricolor as aggregate or showed differences among the genotypes for all
complex comprising various synonyms like A. ten characters. In clustering pattern, genotypes
gangeticus L., A. mangostanus L., A. polygamus were distributed in ten clusters, and the geno-
L., A. melancholicus L. and A. tristis L. These types in these clusters showed significant diver-
synonyms are creating confusion in delimitation gence from each other. Clustering pattern did not
of taxa. Infraspecific morphological variability of reflect any parity with geographic distribution of
A. tricolor can be addressed by introducing taxa the indegenous and exotic genotypes of A. tri-
at variety level ignoring the synonyms. color studied. Devdas et al. (1992) investigated a
Lakshminarasimhan and Godbole (2001) recog- total of 25 accessions belonging to Amaranthus
nised two varieties under A. tricolor – A. tricolor tricolor, A. dubius, A. spinosus and A. viridis for
var. tricolor and A. tricolor var. tristis (L.) 13 biometric characters. The accessions were
Thellung. During comprehensive field survey in classified into seven clusters. The study of inter-
West Bengal, India, a large number of plants and intra-cluster differences revealed greatest
were collected and studied morphologically (Das variability in varieties of A. tricolor.
5.6 Key to the Varieties of Amaranthus tricolor L. 81
Fig. 5.7 A. tricolor var.

acutus S. Das (a) habit,
(b) bract, (c) bracteole,
(d) tepal, (e) male
flower, (f) female flower,
(g) utricle
Table 5.3 Morphological characters delimiting three A. tricolor var. A. tricolor var. A. tricolor var.
varieties of A. tricolor tricolor tristis acutus
A. tricolor var. A. tricolor var. A. tricolor var. Tepal apex Tepal apex Tepal apex acute
tricolor tristis acutus awned or awned or
Small erect Robust erect Robust erect or setaceous setaceous
herb herb procumbent herb Tepal length Tepal length Tepal length
Branching from Branching from Branching from 4.5–5.0 mm 4.5–5.0 mm 2.25–2.5 mm
middle of the base of the stem middle of the Fruit shorter Fruit shorter Fruit equal to
stem stem than the tepals than the tepals longer than the
Leaf Leaf Leaf tepals
deltoid-ovate ovate-lanceolate ovate-oblong Ratio tepal Ratio tepal Ratio tepal
Leaf apex acute Leaf apex acute Leaf apex length/fruit length/fruit length/fruit
notched or length length length
emarginate 0.9–1.0 mm 1.6–2.25 mm 2.0–2.25 mm
Petiole shorter Petiole shorter Petiole equal or
than the leaf than the leaf longer than the Amaranthus blitum L. included in sect.
blade blade leaf blade Blitopsis is a trailing ascending glabrous herb
Bracteole Bracteole Bracteole with light-green ovate-rhomboid emarginate,
ovate-lanceolate ovate-lanceolate ovate-spathulate cuneate leaves widespread in warm tropical cli-
Bracteole apex Bracteole apex Bracteole apex
mate. It might have originated in the Mediterranean
long awned long awned acute
region. A. blitum known as livid amaranth (Britton
Bracte broad Bract broad Bract
ovate ovate ovate-spathulate and Brown 1896) is quite widespread in Indian
Bract apex Bract apex Bract apex acute tropics as well as other Southeast Asian countries
awned awned with a large number of morphotypes. Gradual
Tepal Tepal Tepal domestication process has resulted in the appear-
ovate-lanceolate ovate-lanceolate ovate-spathulate ance of its cultivated form. The wild A. blitum has
(continued) 1.5–2.5 cm long ovate-emarginate leaves, while
A. lividus L. is an erect cultivated form of A. bli- and form ranks) were published, and the situation
tum with large leaves up to 7.0–9.0 cm known as is quite complicated.
separate species. Both have long been known as Moquin-Tandon (1849: 265) accepted five
synonyms (Hooker 1885; Thellung 1914). varieties (α − ε) under A. blitum, three of which
A. blitum is a species with remarkable variabil- (α-sylvestris, δ-graecizans, ε-angustifolius)
ity, and several subspecies have been described; referred to A. graecizans Linnaeus (1753: 990).
some of them have been given species rank (Hugin On the other hand he recognised the genus
1987; Costea et al. 2001b). Keeping in view the Euxolus Rafinesque (1836: 42) and proposed the
morphological variabilities, it would be logical to combinations E. lividus (L.) Moq. (≡ A. lividus),
treat A. blitum as a complex or aggregate compris- E. oleraceus (L.) Moq. (≡ A. oleraceus) and E.
ing several taxa just like A. tricolor aggregate. viridis (Linnaeus 1763: 1405) Moq. (≡ A. viri-
The Amaranthus blitum aggregate belonging dis), the latter of which has four varieties:
to the subgenus Albersia includes four taxa: A. β-ascendens (Loiseleur 1810: 141) Moq. (≡ A.
blitum Linnaeus (1753: 990) sensu stricto, A. bli- ascendens), γ-purpurascens Moq. (new variety),
tum var. oleraceus (Linnaeus 1763: 1403) Costea δ-rubens Moq. (new variety) and ε-polygonoides
et al. (2001b: 984), A. emarginatus (Moquin- Moq. (new variety). The name A. emarginatus
Tandon ex Uline & Bray 1984: 319) sensu stricto Salzmann ex Moquin-Tandon (1849: 274) was
and A. emarginatus var. pseudogracilis (Thellung published as synonym of Euxolus viridis L. var.
1914: 321) Iamonico (2014c). (‘ε’) polygonoides Moq., so it is to be considered
As far as the revision of the genus Amaranthus illegitimate under art. 36.1c of the ICBN (McNeill
is concerned, Iamonico (2010a, b, 2012, 2013, et al. 2012).
2014a, b, c) contributed a lot in several projects, Roxburgh (1832: 605–606) recognised the
and Das (2012a, b, 2013) investigated morpho- taxa A. lividus and A. oleraceus as separate spe-
logical variability of the genus in India. They cies; A. blitum was cited as similar, but not a
jointly contributed to the knowledge of the genus well-defined species, under A. polygamus Willd.
Amaranthus in India emphasising A. blitum (Roxoburgh 1832: 603).
aggregate (Das and Iamonico 2014). Hooker (1885: 721) accepted the name A. bli-
Amaranthus blitum and A. lividus were first tum and recognised three varieties under A. bli-
described by Linnaeus in the first edition of tum: var. blitum, var. oleraceus and var. sylvestris.
Species Plantarum (Linnaeus, 1753: 990). These The latter is distinguishable from the other variet-
two names have generated an interesting nomen- ies in having dehiscent fruit, while the other two
clatural problem. Moquin-Tandon (1849) and varieties have indehiscent utricles. The var. syl-
Thellung (1914) partially included A. graecizans vestris is not part of the A. blitum aggregate and
in the synonymy of A. blitum, leading Brenan is currently accepted as A. graecizans Linnaeus
(1961) to state A. blitum as nomen confusum. subsp. sylvestris (Villars 1807: 111) Brenan
Brenan and Townsend (1980) agree with Brenan (1961: 273) (see, e.g. Costea et al. 2001; Iamonico
(1961) and proposed to list the name A. blitum as 2014d). The var. oleraceus was mentioned as cul-
nomen rejiciendum. Few years later the Committee tivated in India and elsewhere.
for Spermatophyta (Brumitt 1984) rejected this Thellung (1914) accepted the name A. lividus
proposal, reporting ‘…since the last century, and and proposed the following classifications:
in the present century (A. blitum) was used in the
correct sense of A. lividus or at not be used at – lividus proles polygonoides (Moq.) Thell.
all…’. Filias et al. (1980) pointed out that the [now Amaranthus emarginatus]: wild plants
choice between the two names (A. blitum and A. with prostrate or ascending stems, small
lividus) was made by Hooker (1885) favouring A. leaves and fruit about 1.5 mm long. Two new
blitum as the currently accepted name. forms were described: f. pseudogracilis Thell.
Concerning the infraspecific classification of (‘2’) and F. axillaris Thell. (‘3’); the first one
A. blitum, several names (at subspecific, variety differs from the other in having the inflores-
cence in terminal spike (the flowers are in diameter. Also the size of leaves can be used to
arranged in axillary cluster in the F. axillaris) separate these two groups (see the diagnostic key
– lividus proles ascendens (Loisel.) Thell. (now below). The plants with larger seeds can be
A. blitum sensu stricto): wild plants with pros- referred to A. blitum s.l., with two varieties (var.
trate or ascending stems, small leaves and blitum and var. oleraceus, both occurring in
fruit 2.0 − 2.5 mm long India) that distinguish each other on the basis of
– lividus proles týpicus (L.) Thell. (now A. bli- the seed dimensions and surface. The population
tum s. s.): cultivated forms with vigorous, from West Bengal showed a clear distinctiveness
erect or ascending stems and large leaves, having minute seeds, leaves with obtuse or
red-coloured obscurely emarginate apex, bract as long as tepals
– lividus proles oleraceus (L.) Thell. (now A. (ratio bracts/tepals about 1) and acute tepals. The
blitum var. oleraceus): cultivated plants, very populations from Europe are distinct from those
much like proles týpicus but whitish-coloured of West Bengal having leaves with clearly emar-
ginate to bilobed apex and bract always shorter
Hügin (1987) renamed A. emarginatus Moq. than tepals (ratio bracts/tepals 0.5 − 0.6) and
ex Uline & Bray and the combination A. emar- acute tepals. This European population should be
ginatus subsp. pseudogracilis (Thell.) Hügin was assigned to a separate species A. emarginatus s.l.
proposed. [containing two varieties – var. emarginatus and
Based on morphological and ecological obser- var. pseudogracilis – distinguished on the basis
vations, Costea et al. (2001b) recognised the of the habitus and the synflorescence structure].
taxon emarginatus at subspecies rank of A. bli- The Bengal population shows features that can-
tum, also including two varieties: var. emar- not be ascribed to any known Amaranthus spe-
ginatus and var. pseudogracilis (Thell.) Costea cies but certainly belong to the A. blitum
et al. (2001: 981). aggregate but cannot be assigned to A. emar-
Walter and Dobes (2004) carried out a mor- ginatus, given a species status, named as
phometric study confirming the recognition of the Amaranthus bengalense Das & Iamonico sp.nov.
classification proposed by Costea et al. (2001b). In India specially in the lower gangetic plain of
More recently Iamonico (2014c) was in favour Bengal, the ‘Blitum complex’ is represented by
to the separation of the taxa blitum and emar- three varieties of Amaranthus blitum – A. blitum
ginatus at species rank due to their different ori- var blitum, A. blitum var. oleraceus (L.) Hook
gins (Mediterranean Basin, Europe, North Africa and A. bengalense Saubhik Das & Iamonico (Das
and Tropical America, respectively). The taxon and Iamonico 2014).
oleraceus was accepted at variety rank of A. bli-
tum, the taxon pseudogracilis at variety rank of Amaranthus blitum var. blitum A monoecious,
A. emarginatus. annual, erect to prostrate herb attaining a height of
Morphometric investigation on Amaranthus 15–80 cm. Leaves are ovate (2.5–4.5 × 1.5–3.5
blitum complex in India and elsewhere showed cm), entire, long petioled (petiole usually as long
that there exist two varieties of A. blitum in India as the blade) apex emarginate to bilobed, often
(A. blitum var. blitum and A. blitum var. oleraceus mucronate, base cuneate and green.
(L.) Hooker filius (1885: 721). As pointed out in Synflorescences are arranged in terminal spike-
previous works (see, e.g. Costea et al. 2001b; like fashion, erect or reflexed. Bracts are ovate to
Walter and Dobes 2004; Iamonico and Iberite oblong shorter than the perianth. Staminate flow-
2012), the diameter of the seeds is an important ers have three tepals and three stamens; tepals are
character to distinguish the taxa belonging to the usually ovate. Pistillate flowers have three equal
A. blitum aggregate. The analysis reveals two or subequal tepals and three stigmas; tepals are
main groups: the first one including plant with oblong-lanceolate or elliptic (1.2–2.0 mm long),
seeds of 1.1–1.8 mm in diameter and the second membranous with acute apex and thick abaxial
group including plants with seeds of 0.7–1.1 mm midvein. Fruits are reddish brown, compressed,
Fig. 5.8 A. blitum var.

oleraceus (L.) Hooker f.
(a) habit, (b) small
portion of inflorescence,
(g) female flower
subglobose to ellipsoidal utricle, 1.5–2.5 mm and three stamens shorter than the tepals.
long, as long as or longer than the perianth, Pistillate flowers have three equal or subequal
smooth to finely rugose and indehiscent; seeds are tepals and three stigmas; tepals are oblong-
lenticular, black to dark reddish, with minutely lanceolate to spathulate (1.5–2.5 mm long),
punctiform surface 1.1–1.2 mm in diameter. membranous with acute apex. Fruits are reddish
brown, compressed, subglobose utricle (1.8–
2.8 mm long), longer than the perianth, usually
Amaranthus blitum var. oleraceus A monoe- smooth and indehiscent. Seeds are broadly len-
cious, annual, erect, cultivated herb attaining a ticular, black with smooth surface (1.2–)1.4–
height of 25–90 cm. Leaves are green ovate- 1.7(1.9) mm in diameter (Fig. 5.8).
rhomboid (8.0–10.0 × 6.0–7.0 cm), mucronate,
petiolate (petiole as long as or shorter than the A. bengalense Saubhik Das & Iamonico Annual,
blade) with entire margin and cuneate base. monoecious, ascending, cultivated herb attaining
Synflorescences are arranged in terminal spike- a height of 30–40 cm. Leaves are green deltoid-
like pattern and in axillary glomerules. Bracts are ovate to ovate-lanceolate (4.0–4.5 × 3.0–3.5 cm),
ovate, as long as or shorter than the perianth, petiolate (petiole shorter than the blade) with
membranous, with acute to acuminate tip. obtuse to obscurely emarginate apex, entire mar-
Staminate flowers have three tepals usually ovate gin and cuneate base. Synflorescences are
Fig. 5.9 Amaranthus

bengalense Das &
Iamonico (a) habit, (b)
synflorescence part, (c)
bract, (d) bracteole, (e)
tepal of male flower, (f)
tepal of pistillate flower,
(g) male flower, (h)
female flower
arranged in terminal spike-like fashion and in – Seed diameter 0.7–1.1 mm, length of the fruit
axillary glomerules. Bracts and bracteoles are (1.2–)1.4 =−1.8 mm, leave blade size 1−3.5
ovate-lanceolate (4.0–4.5 mm long), as long as (−4.5) × (0.5−) 0.8−3.5 cm ........................ (3)
the perianth, with green adaxial midvein and acu- 2. Seeds with minutely punctiform surface and
minate apex. Staminate flowers have three tepals diameter 1.1−.2 mm .......................................
and three stamens; tepals are lanceolate (4.0–4.5 A. blitum var. blitum
× 1.0–1.8 mm) and acuminate; stamens are shorter – Seeds with smooth surface and diameter (1.2−)
than tepals. Pistillate flowers have three equal or 1.4 − 1.7(−1.9) mm.................................. A.
subequal, oblanceolate to spathulate (2.5–3.0 × blitum var. oleraceus
0.8–1.0 mm) tepals and three stigmas. Fruits are 3. Bracts as long as tepals (ratio bract/tepal about
greenish brown, compressed, subglobose (1.5– one); tepals tip acuminate; leaves with apex
1.8 mm long) utricle, slightly shorter than the obtuse or slightly emarginate .........................
perianth, smooth and indehiscent. Seeds are ..................................................A. bengalense
broadly lenticular, black with smooth surface, – Bracts shorter than the tepals (ratio bract/tepal
0.7–1.0 mm in diameter (Fig. 5.9). 0.5−0.6); tepals acute; leaves with apex clearly
emarginate to bilobed ................................ (4)
4. Stem ascending, synflorescence in axillary
Key to the Varieties of Amaranthus bli- glomerules or short thickened terminal spike-
tum L. like (up to 2 cm long) ................................ A.
A diagnostic key of all the taxa of the A. blitum emarginatus var. emarginatus
aggregate is as follows: – Stem prostrate, synflorescence in terminal long
1. Seed diameter 1.2−1.8 mm, length of the fruit and slender spike-like (up to 10 cm long) often
1.9–3.5 mm, leave blade size (3.0−) thin and flexuous (1.5–7.5 mm long) ................
3.5−9.0 × 1.5−6.2 cm ................................. (2) A. emarginatus var. pseudogracilis
Das (2015) introduced a new vegetable ama- only dioecious species, and the other two, sub-
ranth, Amaranthus parganensis Saubhik Das sp. gen. Albersia and subgen. Amaranthus, comprise
nov. from Lower Gangetic Plain of West Bengal, only monoecious species. There are no records of
India. In general appearance and morphology, any other amaranths having gynomonoecy or
Amaranthus parganensis closely resembles incipient gynomonoecy.
Amaranthus tricolor L. Both the species are Amaranthus parganensis is a robust, erect,
adapted to tropical climate, sympatric in distribu- branched, 100–120 cm tall herb, appearing
tion having similar ecology, distinguished by gynomonoecious but not functionally. Leaves
common features such as the erect habit, leaf are exstipulate, ovate to lanceolate, 13–17 cm ×
size, bracts smaller than the tepal and the fruit as 5–9 cm, with acute or minutely notched apex
a dehiscent, circumscissile utricle with a smooth having a small apicula. Flowers are arranged in
surface and blackish-brown seeds. It is further axillary glomerules and a terminal compound
assigned to sect. Pyxidium under the subgen. inflorescence of cymes, terminal inflorescence
Albersia. The new species shows a structural massive but less branched. Flowers are either
gynomonoecy with rudimentary gynoecia in bisexual or pistillate, unequal in size, intermixed;
bisexual flowers (Fig. 5.10). There is no func- tepals of the bisexual flowers are larger than
tional gynomonoecy, and no seeds are found those of pistillate flowers. Bract, bracteoles and
within the rudimentary carpels. Among the three tepals have apical awns. Bract and bracteoles are
subgenera as delimited by Mosyakin and broadly ovate to lanceolate, 1.5–2.0 mm long.
Robertson (1996), subgen. Acnida comprises Perianth consists of three ovate to lanceolate or
Fig. 5.10 Amaranthus

parganensis Saubhik
Das (a) fertile habit, (b)
aristate bract, (c) aristate
bracteole, (d) aristate
tepal, (e) bisexual
flower, (f) bisexual
flower with stamens and
rudimentary carpel, (g)
pistillate flower, (h)
gynoecium
oblong, membranous tepals with acuminate uated using protein and RAPD markers. Though
apex, 3.0–3.5 mm long in pistillate flowers. high level of genetic diversity was common within
Bisexual flowers have three stamens, 4.5– species, genetic uniformity was observed within
5.0 mm long. Style is short and thickened, ovary most accessions. Srivastava and Roy (2012) stud-
ovoid, three stigmas, filiform and papillose. ied four species (A. viridis, A. spinosus, A. tricolor
Gynoecium in the bisexual flower is rudimentary and A. blitum) and their accessions. The study
and has a short thickened gynophores, but revealed that the accessions of each species were
gynoecium in the pistillate flowers lacks a gyno- very close to each other, but variation was observed
phore. Fruits are circumscissile utricle, produced at the species level. A. spinosus was shown to be
only by pistillate flowers, ovoid and 1.5–2.0 mm the next most closely related to A. tricolor after A.
long. Seeds are blackish brown, shiny, com- blitum, though Zheleznov et al. (1997) reported no
pressed and 1.2–1.5 mm in diameter. phylogenetic relationship between A. tricolor and
Morphological features separating the prime A. spinosus; Chan and Sun (1997) also reported
vegetable species of Amaranthus are mentioned dissimilarity among A. viridis, A. tricolor and A.
in Table 5.4. spinosus. Samour (1991) observed a definite pro-
Genetic diversity and relationship among the tein profile, intermediate between A. viridis and A.
cultivated and wild Amaranthus species were eval- hybridus in a population of A. viridis indicating a
Table 5.4 Comparative morphology of major vegetable amaranths

Morphological
characteristics A. parganensis A. tricolor A. blitum
Breeding system Incipiently gynomonoecious Monoecious Monoecious
Stem Erect, 100–120 cm Erect, 90–100 cm Erect or ascending,
35–45 cm
Leaf shape Ovate-lanceolate or Deltoid-ovate or ovate to Ovate to lanceolate or
lanceolate, not rhombic lanceolate or oblong rhomboid-ovate
Leaf sizes 13.0–17.0 × 5.0–9.0 cm 9.0–7.0 × 5.0–9.0 cm 2.5–8.0 × 1.5–6.0 cm
Ratio lamina/petiole 2.1–2.6 0.89–1.66 0.5–1.5
length
Leaf apices Acute or minutely notched Acute or slightly Emarginate
emarginate
Inflorescence Terminal inflorescence Terminal inflorescence Terminal inflorescence
massive but less branched massive and much short
branched
Flowers Both bisexual and unisexual Unisexual, staminate and Unisexual, staminate and
(female flower), bisexual and pistillate flowers are pistillate flowers are
female flowers unequal equal equal
Bract vs. tepal Bract < tepal Bract < tepal Bract ≤ tepal
Bract length 1.5–2.0 mm 1.5–2.0 mm 0.5–2.5 mm
Tepal shape Ovate to lanceolate or oblong Ovate to lanceolate or Lanceolate to spathulate
spathulate
Tepal tip Acuminate Acute or awned Acute or awned
Tepal length 3.0–5.0 mm 2.25–2.5 (−5) mm 1.2–2.5 mm
Fruit vs. tepal Fruit not exceeding tepals Fruit may or may not Fruit marginally
exceed tepal exceeding tepals
Fruit surface Smooth Smooth Faintly rugose
Dehiscence Circumscissile utricle, Circumscissile utricle, Indehiscent
dehiscent dehiscent
Seed diameter 1.2–1.5 mm 1.2–1.5 mm 1.1–1.8 mm
Seed colour Blackish brown Blackish brown Black
probable hybridisation between the two taxa. be merged and considered as a single species or
These results confirmed the suggestions of their separate identity would be maintained.
Drzewiecki (2001) that the grain and leafy types of Wassom and Tranel (2005) used AFLP-based
cultivated amaranths (from India) could be easily marker to highlight the phylogeny of both dioe-
distinguishable as two genetic groups on their cious and monoecious amaranth. Eight weed spe-
fixed allozyme alleles. The wild amaranths as a cies represented by 141 individuals from 98
rule showed high RAPD polymorphism than the different accessions were taken into consider-
cultivars (Srivastava and Roy 2012). Highest poly- ation. Interestingly, the dioecious weed A. rudis
morphism was shown by A. viridis, while A. bli- and A. tuberculatus did not group together, per-
tum showed the lowest level of polymorphism. haps indicating independent evolutionary ways
Dendrogram computed on RAPD and protein and separate genetic entities as well. Correct iden-
marker showed that cultivated amaranths are most tification of Amaranthus species is often difficult
closely related to each other than the wild mem- and frequent misidentification is common (Horak
bers. A high degree of genetic variability shown by et al. 1994; Wax 1995; Sauer 1953; Arhens et al.
the wild species is expected and explainable 1981). For example, Ahrens et al. (1981) found
because they are not subjected to any selection that 13 of 14 accessions of weed amaranths that
pressure of domestication; as such A. viridis weed scientists earlier identified as redroot pig-
showed highest polymorphism and A. blitum the weed were actually smooth pigweed (Amaranthus
least polymorphism. Observation of Chan and Sun hybridus L.) or Powell amaranth (Amaranthus
(1997) was also concomitant to this interpretation powellii S. Wats.). As another example, Wetzel
as they found 39.9 % polymorphism in crop spe- et al. (1999a) applying molecular marker analysis
cies, 51 % polymorphism in vegetable species and of ribosomal internal transcribed spacer (ITS)
69.5 % polymorphism in wild species of regions found that 12 of 92 Amaranthus acces-
Amaranthus. sions that had been collected and identified by
weed scientists earlier were misidentified. In spite
of accuracy it is not possible to apply molecular
5.7 Taxonomic Delimitation marker-based identification for routine use. It
in Weed Amaranths could be applied to verify the identity of trouble-
some biotypes with ambiguous or atypical mor-
Most of the people try to identify weeds based on phology. Response to herbicides can be a very
‘how the plant looks’, i.e. to recognise taxa on the promising tool to delimit taxa. As for examples,
pigmentation or growth forms which are Coetzer et al. (2002) reported significantly differ-
extremely variable (Sauer 1967); more accurate ent responses to glufosinate herbicide among
identification needs vivid morphological study of waterhemp (Amaranthus tuberculatus (Moq.)
the floral parts. Historically, taxonomic delimita- Sauer and Amaranthus rudis Sauer), redroot pig-
tion in Amaranthus species has been done mostly weed (Amaranthus retroflexus L.) and Palmer
based on differences in floral characteristics, but amaranth (Amaranthus palmeri S. Wats.). Mayo
new methods using molecular biological tech- et al. (1995) reported that Palmer amaranth was
niques are also gaining importance. Correct iden- more difficult to control with various herbicides
tification of Amaranthus weed species is necessary than were other Amaranthus species.
for efficient weed control (Horak et al. 1994;
Sweat et al. 1998). Some molecular studies have
been applied to weed species. Wetzel et al. (1999a) 5.8 Different Phylogenetic
developed ribosomal ITS restriction site-based Concepts on Grain
PCR-generated marker to identify common ama- Amaranths
ranth weeds which were otherwise difficult to be
identified morphologically. Pratt and Clark (2001) It is generally considered that the vegetable and
used isozyme polymorphism to address the ques- the grain amaranths are two distinct groups that
tion whether A. rudis and A. tuberculatus would have originated from their respective weed pro-
5.8 Different Phylogenetic Concepts on Grain Amaranths 89
genitor. Both the groups have different centres of domestication of A. caudatus by crossing between
origin and unique processes of domestication. A. cruentus and A. quitensis (Fig. 5.11b). Since
Weedy and grain Amaranthus spp. are inseparably 1940 Amaranthus powellii has become a wide-
linked phylogenetically and historically. The grain spread and troublesome weed in Eastern North
amaranths are supposed to have derived from their America. In Europe it was misidentified as A.
respective weed progenitors through frequent out- chlorostachys Willd., a synonym of A. hybridus.
crossing and gradual unique domestication pro- The species is exceptional in producing flowers
cess. Two hypotheses have been proposed by with tepal and stamen numbers varying from 3 to
Sauer (1967, 1976). The first hypothesis is based 5, even in a single plant. Individual plants with
on geographical distribution, which suggests that indehiscent utricle also occur sporadically in
all the three grain amaranths have evolved inde- Native American populations and have formed the
pendently, i.e. origin is polyphyletic. A. hypochon- entire local population in Europe (Brenan 1961;
driacus was domesticated in Mexico from A. Aellen 1961). The hybrids between other ama-
powellii, A. cruentus from A. hybridus in Central ranth species may show such aberrant forms, but
America and A. caudatus from A. quitensis in aberrations in A. powellii evidently arisen without
South America (Fig. 5.11a). The second hypothe- hybridisation. Amaranthus quitensis is a riverbank
sis based on plant and seed morphology suggested pioneer of South America. The plant is semi-culti-
monophyletic origin of the grain amaranths. The vated or tolerated as a source of food additives.
three cultivated grain species have originated from Three grain amaranths along with A. hybridus are
a single weed progenitor A. hybridus, following supposed to have formed a complex or aggregate
subsequent introgressive hybridisation with two (‘hybridus species complex’) in which taxonomic
other wild species in different regions. According disputes and phylogeny are yet to be resolved with
to this hypothesis, the first domesticated species extreme clarity, especially because of apparent
was A. cruentus, originated from A. hybridus in common hybridisation and misapplication of
Central America, followed by the domestication of nomenclature.
A. hypochondriacus by repeated crossing between More recently a third hypothesis was pro-
A. cruentus and A. powellii in Mexico and the posed by Mallory et al. (2008). He suggested
Fig. 5.11a Polyphyletic

theory regarding origin of
grain amaranths as proposed USA
by Sauer (1967)
Mexico
Central America
A. powelli
A. cruentus
A. hybridus
A. hypochondriacus
A. caudatus A. quitensis
South America
Fig. 5.11b Monophyletic

theory regarding origin of
grain amaranths as proposed
by Sauer (1967)
Amaranthus hypochondriacus Amaranthus caudatus Amaranthus cruentus
Independent domestication Independent domestication

event event
Genetically differentiated Genetically differentiated

Genetically differentiated
population population
population
Amaranthus hybridus
Fig. 5.12 Derivation of grain amaranths from ancestral A. hybridus through separate genetically differentiated popula-
tions as proposed by Mallory et al. (2008)
that A. hybridus is the progenitor species for all were monophyletic, while A. hybridus was
the three domesticated grain species, but each polyphyletic as it appeared from the presence
was derived through independent domestica- of A. hybridus accessions in each of the three
tion events from genetically differentiated pop- grain amaranth monophyletic clades. They also
ulations of A. hybridus (Fig. 5.12). The study observed that A. powellii (a previously recog-
designated A. hybridus as the only progenitor nised progenitor of grain amaranths) has
species of all the three grain amaranth species. formed a monophyletic group distinct from any
According to neighbour-joining analysis, A. of the grain amaranths. It is obvious that a
caudatus, A. cruentus and A. hypochondriacus larger and more inclusive and comprehensive
investigation, involving substantially more domestication event has occurred involving A.

diversified accessions of both the grain ama- hybridus as the common ancestor rather than
ranth and progenitor species (A. hybridus, A. separate domestication event (Kulakow et al.
powellii and A. quitensis), is still needed to 1985; Kulakow and Jain 1990a, b). But relative
have a clear idea about origin and phylogeny of closeness between A. caudatus and A. quitensis
the grain amaranths. However, one can pre- shows some sort of deviation from Saure’s per-
sume that taxonomic identification of these ception. This suggests a separate domestication
Amaranthus species may be ambiguous due to event of A. caudatus from A. quitensis. It is also
reciprocal gene flow that resulted from out- presumed that A. quitensis may have also derived
crossing in regions where the species are from A. caudatus through introgressive hybridi-
sympatric. sation with A. hybridus, or it is semi-domesticate
Progenitor status of A. hybridus is supported which arose from the escape of A. caudatus from
by the observation that it forms hybrid with all cultivation. Amaranthus quitensis is found to be
the other species in the ‘hybridus complex’ (Pal ecologically a semi-domesticate in Ecuador
and Khoshoo 1972, 1973a, b, 1974). Study occurring only in cultivated field (Coons 1982;
involving neighbour-joining method based on Hauptli and Jain 1984).
combined ITS, ISSR and AFLP data (Xu and Sun Electrophoretic profile of total seed protein
2001) also supported the same monophyletic ori- revealed that A. cruentus and A. hypochondria-
gin of the grain amaranths and close relationship cus are closely related to A. hybridus (Sammour
between A. caudatus and A. hypochondriacus. et al. 1993). Sammour concluded that A. cruen-
Kirkpatrick (1995) utilised morphological fea- tus and A. hypochondriacus are subspecies of A.
tures, isozyme polymorphism and nuclear ribo- hybridus. Amaranthus cruentus and A. hypo-
somal DNA analytical data to study interspecific chondriacus have more or less the same amount
relationship. Morphological features of A. hybri- of DNA content, whereas A. caudatus differs
dus were overlapping with that of cultivated grain significantly from other two species in DNA
amaranths, while A. retroflexus L., A. spinosus, A. content (Greizerstein and Poggio 1994). On the
palmeri S. Watson and A. powellii were morpho- other hand, studies involving interspecific fertil-
logically distinct. Amaranthus hybridus was also ity, electrophoretic data or RAPD data favoured
grouped with grain amaranths, while the other much closer genetic affinity between A. cauda-
weed species were distinct. All these evidences tus and A. hypochondriacus than either one with
favoured the concept of monophyletic origin of A. cruentus (Gupta and Gudu 1991; Transue
grain amaranths from A. hybridus. Studies on et al. 1994). Vegetable amaranths were studied
allozyme variation in amaranths also proved for their morphology and karyotype
helpful in the investigation of phylogenetic rela- (Madhusoodanan and Nazeer 1983). The study
tionship among weedy and crop species (Hauptli revealed that both, the chromosome size and
and Jain 1984, 1985). Molecular marker analysis gross morphology, vary according to the selec-
has contributed a lot to explore origin and evolution factor and/or adaptive protocols.
tion of cultivated amaranth and allied wild spe- Chromosome repatterning and structural modifi-
cies, and Hauptli and Jain (1984) were among the cation played a key role in differentiation and
pioneering workers who firstly used molecular evolution of Amaranthus species.
marker to address evolutionary relationship Investigation on morphological and genetic
among grain amaranths. In their study, they con- variability in Indian amaranth showed a high
sidered nine enzyme systems, and the study degree of morphological polymorphism but least
revealed that the grain species are related more allozymic variability, i.e. maximum monomor-
closely to each other than they are to their respec- phism of allozyme loci. The lack of positive cor-
tive wild progenitors. The study supported the relation between morphological polymorphism
monophyletic origin of the grain amaranths as and genetic variability may be due to relatively
proposed by Sauer and indicated that a single recent introduction and rapid spread of grain
amaranths in India. Putative progenitor-derived cious counterparts. The investigation by Chan

species pair often shows high genetic identities, (1996) involving RAPD assay and isozyme poly-
isolated by reproductive barriers such as chromo- morphism supported Sauer’s monophyletic
some repatterning or deviation in breeding sys- hypothesis and projected A. hybridus as the pro-
tem. Speciation can still proceed with little or no genitor of all the three grain amaranths but did
divergence at isozyme loci (Crawford 1983). The not support the single origin hypothesis of the
phylogenetic affinity of 28 species of Amaranthus dioecious amaranths like Lanoue et al. (1996)
were studied by Lanoue et al. (1996), applying (Fig. 5.13).
restriction site analysis of PCR-generated chloro- Previous phylogenetic studies investigating
plast and nuclear DNA. Genetic variation found the origin of grain amaranth involved inadequate
by the restriction site analysis was used to con- sampling of all the grain amaranths as well as the
struct a cladogram of 28 species. The two putative weedy progenitors A. hybridus and
tree reflected A. dubius and/or A. hybridus as A. quitensis across their native species range as
the possible progenitors of A. cruentus and well as inadequately investigated the genetic
A. quitensis as the probable progenitor of A. cau- diversity component of both putative weedy
datus. Contradicting the previous observations species. Sampling errors like under-representa-
(Hauptli and Jain 1984; Transue et al. 1994), tion of weedy species compared to the cultivated
Lanoue found much closer affinity between A. species or over-representation of A. hybridus
caudatus and A. cruentus than each of them with with exclusion of A. quitensis or involvement of
A. hypochondriacus. The study does not support weedy representatives collected from outside the
the single origin hypothesis of the dioecious species native range accounted for discordant
amaranths like A. australis, A. cannabinus, A. interpretation between previous investigations
floridanus, A. palmeri and A. rudis as proposed (Kietlinski et al. 2013). Both the Sauer’s hypoth-
by Sauer (1955). Lanoue suggested that A. aus- eses, i.e. single progenitor theory and indepen-
tralis, A. cannabinus and A. rudis are more dent domestication hypothesis, are based on
closely related to each other, whereas A. florida- morphology that is to be evaluated. The specula-
nus and A. palmeri are more similar to other tions made by Sauer based on morphological
monoecious amaranths rather than other dioe- evidences validated by several workers in a
A. dubius A. retrofexus
(Allotetraploidization)
A. powellii A. quitensis
(Intogression)
A. hypochondriacus A. caudatus
(Intogression)
A. hybridus A. cruentus
(Early domesticate)
Fig. 5.13 Concept of Chan (1997) regarding origin of grain amaranths

A. caudatus A. hypochondriacs A. cruentus
A. hypochondriacus
A. caudatus
A. quitensis
A. cruentus
A. hybridus
Early
domesticate
A. hybridus
Fig. 5.14 Studies on evolutionary aspects of grain amaranths (a) phylogenetic relationship exclusively between three
grain types and (b) phylogenetic relationship among all members of hybridus complex
fragmentary fashion not in a conclusive manner. regarding the domestication of A. hypochondria-

All the studies were channelised in either of the cus and A. caudatus:
two directions, i.e. (1) studies addressing the
phylogenetic relationships exclusively between 1. Single domestication hypothesis – in which
the three grain amaranths (Fig. 5.14a) and (2) one domestication event occurring in Meso-
studies addressing the phylogenetic relation- America or the Andes in A. hybridus, followed
ships among all members of the hybridus com- by geographical separation giving rise to A.
plex (Fig. 5.14b). Kietlinski et al. (2013) hypochondriacus and A. caudatus.
considered 258 individuals from 56 taxa repre- 2. A dual-lineage domestication hypothesis – in
senting the three grain amaranths species and which a single A. hybridus lineage spanning
two putative weedy progenitor samples from Meso-America to the Andes was indepen-
their native species range and subjected to mic- dently domesticated at least twice giving rise
rosatellite analysis. Data analysed by PCA, clus- to A. hypochondriacus and A. caudatus.
tering algorithms and genetic distance Amaranthus cruentus appears to have origi-
neighbour-joining tree dendrogram highlighted nated separately from second widespread A.
few focal points, viz. (1) the grain amaranths and hybridus lineage spanning across Guatemala
Amaranthus hybridus were found to group and Central Mexico. This species appeared to
together and all the grain amaranths have evolved be more diverse relative to its cultivated coun-
from the progenitor A. hybridus via at least two terparts as evidenced by high allelic diversity
domestication event. (2) A. quitensis is its own and numerous private allele (Kietlinski et al.
species, a distinct group not a variant of A. hybri- 2013). The ‘hybridus species complex’ of
dus. (3) Amaranthus caudatus L. and Amaranthus Amaranthus comprises a group of cultivated
hypochondriacus L. appear to be closely related grain species and weedy species from the
though both the species are geographically sepa- New World where taxonomic delimitation and
rated. A strong inbreeding coefficient was found systematic are poorly understood. The ‘hybri-
in all the studied taxa. Amaranthus hybridus may dus species complex’ comprises widespread
consist of two widespread morphologically agricultural weed A. hybridus, South American
cryptic species or may consist of a single highly endemic A. quitensis and three grain
variant species from which all grain amaranths amaranths. Adhikary and Pratt (2015) estab-
arose. The study favoured two hypotheses lished an interrelationship between the com-
ponent species of this complex studying the gene flow, though genome size and seed size did
floral variation pattern within the species com- not appear to be significantly different between
plex. Twenty-one pistillate and 12 staminate wild and domesticated amaranth. It was con-
floral characters from 41 specimens affiliating cluded that A. caudatus is an incompletely
to five species were analysed morphologically. domesticated species, which may be due to lack
Results indicated that the hybridus complex to of strong selection process or due to high level of
be divided vertically into two morphologically gene flow from its sympatric wild relatives pre-
distinct larger groups by A. hybridus each venting the fixation of key domestication traits in
comprising different cultivated taxa. One A. caudatus.
group includes A. cruentus, A. caudatus, A. In grain crops, grain size and seed coat colour
quitensis and one morphotype of A. hybridus are important traits for selection and supposed to
and another group comprises A. hypochondri- play a key role in domestication of various plants
acus and a different morphotype of A. hybri- (Hake and Ross-Ibarra 2015). Key domestication
dus. The result though supported the concept traits such as shape of inflorescence, seed shatter-
of considering A. hybridus as a progenitor spe- ing and seed size were rather similar between
cies, but closer affinity of A. hybridus with wild and cultivated amaranth. Domesticated
both A. cruentus and A. caudatus was left con- amaranth shows morphological differences from
troversial. In the study staminate morphologi- wild amaranth having large and more compact
cal variation appeared to be more taxonomically inflorescence (Sauer 1967). Trait like white seed,
informative than assumed before. surely a domestication-related trait, is predomi-
nant in cultivated amaranth though other seed
Stetter et al. (2015) investigated the domesti- coat colour may have been preferred. Genes for
cation events of A. caudatus and its relationship white seed coat colour were not fixed. Lack of
with the two wild relatives A. hybridus and A. knowledge about genetic basis of domestication
quitensis applying genotyping by sequencing trait, lack of strong domestication syndrome and
(GBS) to genotype on 119 amaranth accessions fixation of putative domestication trait in spite of
from the Andean region. The population genetic long cultivation period may be due to genetic
analysis based on 9485 SNPs revealed very little constraints and ongoing gene flow between wild
genetic differentiation between the two wild spe- and domesticated amaranth. The genotyping of
cies, negating their separate identity, but a strong wild and cultivated amaranth accessions revealed
differentiation between wild and domesticated a strong genetic differentiation between wild and
amaranth A. caudatus. Earlier separate identity cultivated amaranths and a high level of genetic
of A. quitensis and A. hybridus was ignored by differentiation within domesticated A. caudatus
Coons (1978). The taxonomic differentiation due to ongoing gene flow making it incompletely
between these two wild amaranths is resting on a domesticated. The history of cultivated ama-
very fragile morphological trait namely the shape ranths represents a multiregional, multiple and
of the tepals which is very prone to misidentifica- incomplete domestication process with frequent
tion (Sauer 1967). The genome size and pheno- and ongoing gene flow from sympatric wild rela-
typic variation in two domestication-related traits tives showing similarity to the history of rice,
like seed size and seed coat colour were deter- apple or barley (Londo et al. 2006, Cornille et al.
mined and compared between A. caudatus and its 2012, Poets et al. 2015). Experimental data were
two wild relatives by Stetter et al. (2015). consistent with the concept of Kietlinski et al.
Amaranthus caudatus has a higher genetic diver- (2013) who proposed a single domestication of
sity than its wild relatives, and about 10 % of A. caudatus and A. hypochondriacus in Central
accessions showed a strong admixture between America followed by migration of A. caudatus to
the wild and cultivated species suggesting recent South America.
Weed and Herbicide Resistance
6
6.1 General 6.2 Resistance to Different Types

of Herbicides
Weed management in agriculture has become
very problematic and costly due to the increase Herbicide resistance in crops is generated through
in the development of resistance to common a few mechanisms like enhanced metabolism,
herbicides by the obnoxious weeds. More than sequestration, target site resistance, reduced
180 different weed species showed resistance uptake and overproduction of herbicide target
and amaranths amount more than 5 % of the site. Resistance against the mechanism of ten
total biotypes. Eleven weedy species of ama- herbicides has been confirmed. The most wide-
ranths showed resistance. There are different spread resistances developed are against photo-
groups of herbicides and amaranths showed system II (PS II) inhibitors, photosystem I (PS I)
variable response to those. Among the 60 pho- inhibitors, acetolactate synthase (ALS) inhibi-
tosystem II inhibitor resistant angiosperms tors, EPSP synthase inhibitors (e.g. glyphosate),
recorded, amaranths contribute nine species. protoporphyrinogen oxidase (PPP or PROTOX)
More than 90 weeds developed resistance inhibitor, acetyl-coa carboxylase (ACCase)
against ALS inhibitor; pigweeds are repre- inhibitors, mitotic inhibitor and auxinic inhibitor.
sented by eight resistant species. Amaranthus In each case resistance is induced by multiple
tuberculatus is one of the three species that amino acid changes within herbicide-binding
have developed resistance against protoporphy- domain.
rinogen oxidase (PPO) inhibitor. It is a trouble- A worldwide catalogue of herbicide-resistant
some weed that shows multiple resistances to weeds (Heap 2008) includes more than 300 resis-
different herbicides. It has evolved resistance to tant biotypes. A biotype is a group of plants
five different classes of herbicides. In midwest- within a species with a distinct genetic variation.
ern agricultural field of the USA, it shows two A herbiside- resistant biotype is a particular weed
distinct groups of populations – agricultural species with resistance to a particular herbiside
and nonagricultural – with varying degrees of or a group of herbisides with same site of action.
resistance against ALS. It is possible to transfer Herbiside-resistance is an inherited ability of a
resistance trait from weed to crop through weed or crop species to survive a herbiside appli-
weed-crop hybridisation. cation. This catalogue of herbicide-resistant

96 6 Weed and Herbicide Resistance
weed includes more than 180 different weed spe- mechanism was the first ever characterisation of
cies, and amaranth is considered as the leading evolved herbicide resistance at DNA level
member in the list comprising more than 5 % of (Hirschberg and McIntosh 1983).
the total resistant biotypes. Among weedy ama-
ranths, herbicide resistance has been detected in
11 species (Heap 2010). The first report of herbi- 6.2.2 Resistance to Acetolactate
cide resistance shown by Amaranthus species Synthase (ALS) Inhibitors
came up from North America; Amaranthus hybri-
dus showed resistance against the triazine herbi- More than 90 weed species are known to have
cide atrazine in 1970 (Ryan 1970). The biotypes evolved resistance to ALS inhibitor. Eight species
of Amaranthus tuberculatus have been reported of pigweeds showed confirmed resistance against
to show multiple resistances against three herbi- ALS inhibitor. Target site resistance is the most
cide actions (ALS, PPO and PS II) (Patzoldt et al. common mechanism of resistance against ALS
2005). Amaranthus palmeri and Amaranthus inhibitors, and it is indicative of why ALS target
rudis that have shown resistance against herbi- site mutation occurs so frequently. In a survey of
cide glyphosate and ALS inhibitor, respectively, herbicide resistance among Illinois waterhemp
are creating problems in the weed management population, several were observed with varied
system in cultivation field of soybean, maize and response to imazethapyr and thifensulfuron, two
cotton in the USA. Amaranthus species provide acetolactate synthase (ALS)-inhibiting herbi-
ample scope to study herbicide resistance mecha- cides. Multiple ALS mutations were a determin-
nism and may be treated as models to examine ing factor for herbicide resistance in waterhemp
the evolution of herbicide resistance across broad (Patzoldt and Tranel 2007). Due to the impor-
geographical region. tance of ALS-inhibiting herbicide and the fre-
quency at which weed population have developed
resistance in them against ALS inhibitor, a wealth
6.2.1 Resistance to Photosystem II of data has been generated (Guttieri et al. 1996;
Inhibitors Duggleby and Pang 2000; Tranel and Wright
2002).
Resistance to photosystem (PS) II inhibitor
(e.g. triazine herbicide) in Senecio vulgaris is
often regarded as the first major report of evolved 6.2.3 Resistance
herbicide resistance in a weed (Ryan 1970). More to Protoporphyrinogen
than 60 weed species have been identified to Oxidase (PPO) Inhibitors
resist PS II inhibitor, of which nine species
belong to genus Amaranthus. Early research on Resistance to herbicide that target the enzyme
triazine resistance in A. hybridus contributed protoporphyrinogen oxidase has been documented
much to the understanding of the fitness penalty only in three species (Heap 2008). The first
associated with this resistance (Arhens and Stoller species which showed this type of resistance
1983; Ort et al. 1983) and also to the exploration was A. tuberculatus (Shoup et al. 2003). The
of mechanism of resistance (Steinbank et al. resistance to Protox-inhibiting herbicide in an A.
1981; Hirschberg and McIntosh 1983). Triazine tuberculatus biotype from Illinois was due to
resistance in A. hybridus occurred due to glycine DNA mutation caused by codon deletion in the
for serine codon change in the psbA gene of gene encoding the mitochondrial Protox isomer
triazine resistance. The identification of this (PPX2) (Patzoldt et al. 2006).
6.2 Resistance to Different Types of Herbicides 97
The general characteristic that contributed to (a) Whether the Amaranthus species possess
the efficacy of Amaranthus species as weed in ‘innate genetic properties’ like transposable
modern agriculture is their well-exemplified abil- element that preconditions the rapid evolu-
ity to evolve herbicide resistance. Weed amaranths tion of herbicide resistance
contribute over 5 % of herbicide-resistant biotypes (b) Whether hybridisation among Amaranthus
distributed worldwide and has evolved resistance species is responsible for its innate ability to
to diverse herbicide like that which inhibit photo- evolve resistance
system II (PS II), acetolactate synthase (ALS), (c) Whether insertion/deletion (which is com-
protoporphyrinogen oxidase (PPO) and 5-enolpyr- mon in Amaranthus species) are contributing
uvyl-shikimate-3-phosphate synthase (EPSPS) factors to the evolution of herbicide
(Heap 2010; Tranel and Trucco 2009). In some resistance
cases resistance is evolved against more than one
of these herbicides within a single population or These are some of the prime questions that
even a single plant, making the control of weed require investigation. Pigweeds have already
like A. tuberculatus a real problem (Patzoldt et al. offered a stiff challenge towards herbicidal con-
2005). Frequent occurrence of herbicide resistance trol on weed through resistance evolution. A
in weedy amaranths indicates the possibility of large number of herbicide-resistant varieties of
occurrence of the same trait in cultivated amaranth weed amaranths have evolved in different parts of
and the feasibility of possible transfer of this resis- the world (Table 6.1).
tant trait from weed to crop via interspecific The evolution of invasiveness of weeds has
hybridisation. Nevertheless, the only Amaranthus been extensively studied in natural ecosystems.
species that has developed resistance against PPO In spite of its great economic impact on crop pro-
inhibitors is A. tuberculatus and against EPSPS ductivity, little is known about the evolution of
inhibitors are A. tuberculatus and A. palmeri. agricultural invasiveness of weed. Evaluation of
Dioecious nature of A. tuberculatus undoubtedly the population structure of recently arisen weeds
contributed to its ability to develop multiple resis- can be very informative about evolutionary trends
tances. Gene flow via pollen movement from one to the invasion of agroecosystems. Waselkov and
resistant biotype to another effectively combines Olsen (2014) addressed several questions about
different resistance traits, although two self-polli- the origin of the Native North American agricul-
nating monoecious species A. powellii and A. ret- tural weed waterhemp (Amaranthus tubercula-
roflexus have also evolved resistance against tus), which invaded corn and soy fields in the
multiple herbicides (Heap 2008). Tranel et al. midwestern USA in the twentieth century.
(2002) examined introgression of herbicide resis- Agricultural invasion by native, wild plant spe-
tance allele of ALS from A. tuberculatus to A. cies can follow different evolutionary directions
hybridus, and they also observed that A. tubercula- from weeds related to domesticated plants, which
tus allele could not be introduced into A. hybridus has great implications in evolutionary biology
with monoecious background. Considering mon- and weed control. Amaranthus tuberculatus has
oecism and dioecism as the taxonomic distin- evolved resistance to five different classes of her-
guishing character for both the taxa, it appears that bicides: (1) photosystem II inhibitors, (2) aceto-
gene flow between these species is unidirectional. lactate synthase (ALS) inhibitors, (3)
Available informations have raised several queries protoporphyrinogen oxidase (PPO) inhibitors,
regarding efficient multiple herbicide resistance of (4) p-hydroxyphenylpyruvate dioxygenase
Amaranthus species: (HPPD) inhibitors and (5) EPSP synthase inhibi-
98 6 Weed and Herbicide Resistance
Table 6.1 Herbicide-resistant Amaranthus species

Herbicide Species Year of documentation Country where resistance documented
PS II inhibitors A. hybridus 1972 USA, South Africa, France, Italy, Switzerland,
Spain, Israel
A. powellii 1977 USA, Canada, France, Switzerland, Czech
Republic
A. lividus 1978 Switzerland, France
A. retroflexus 1980 USA, Canada, France, Germany, Switzerland,
Bulgaria, Czech Republic, Spain, Poland,
Greece, Italy, Serbia, China
A. blitoides 1983 Israel, Spain
A. albus 1987 Spain
A. cruentus 1989 Spain
A. palmeri 1993 USA
A. tuberculatus 1994 USA, Canada
ALS inhibitor A. blitoides 1991 Israel
A. palmeri 1991 USA
A. retroflexus 1991 Israel, USA, Canada, Italy
A. hybridus 1992 USA
A. lividus 1993 USA
A. tuberculatus 1993 USA, Canada
A. powellii 1996 USA, Canada
A. quitensis 1996 Argentina, Bolivia
Glyphosate A. palmeri 2005 USA
A. tuberculatus 2005 USA
Protoporphyrinogen A. tuberculatus 2001 USA
oxidase inhibitor
Bipyridyliums A. lividus 1990 Malaysia
Dinitroanilines A. palmeri 1989 USA
Heap (2008)
tors, glyphosate. The species has extremely large gests that gene flow is responsible for its presence
agricultural populations. Its constantly evolving in natural habitat. It may be possible that allele
resistance makes it one of the weeds difficult to arose independently in natural population and
control in midwestern agricultural field of the persisted at low frequencies. On average, agri-
USA. The waterhemp has two distinct groups of cultural waterhemp populations showed higher
populations, agricultural and nonagricultural, ALS inhibitor resistance than did natural popula-
with varying resistance potentials against ALS tion. It is possible that the mutation enhanced the
inhibitor. Eleven Ohio populations of Amaranthus performance of the plants in the fluctuating natu-
tuberculatus from both agricultural and natural ral environment of riverbank habitat. The varia-
habitat in the riverbanks were screened to evalu- tion in resistance level among the agricultural
ate the concept that agriculturally adaptive ALS populations may be due to herbicide rotation
inhibitor resistance alleles are found in natural practices that vary from year to year and differ-
habitat. The presence of the same alleles in natu- ence in rotation schedule as well (Waselkov and
ral and agricultural waterhemp population sug- Olsen 2014).
Distribution and Maintenance
of Amaranth Germplasm 7
Worldwide
7.1 General 7.2 Germplasm Collection

Centres in the World
Amaranths show a wide geographical distribution,
evolution of landraces and domestication in The amaranths are characterised with wide
widely spaced areas. To carry out research on geographical distribution that has resulted in the
genetic improvement of the crop, proper collec- evolution of many landraces in widely separated
tion, maintenance and periodical evaluation of areas. The wide gene pool of amaranths is to be
germplasm are a prerequisite. There are quite a assessed and characterised for future development
few amaranth research centres and germplasm of the crop. This needs systematic collection and
collections all over the world which are maintain- maintenance of germplasm worldwide. Amaranth
ing a working germplasm collection. The germ- germplasm is catalogued and stored in germplasm
plasm collections are maintained in at least 61 banks in 11 countries (Toll and van Slotten 1982).
collection centres. Prominent among them are A descriptive taxonomic key for all the cultivated
Rodale Research Center, Pennsylvania; USDA species of the genus Amaranthus was developed
Plant Introduction Center at Ames, Iowa; by Feine-Dudley (Grubben and van Slotten
National Bureau of Plant Genetic Resources 1981). Since 1977, Rodale Research Center
(NBPGR), New Delhi; etc. Amaranth genome (RRC) Pennsylvania, USA, is maintaining a
may serve as a model system to study weediness. working germplasm collection. The 1400 acces-
A bacterial artificial chromosome (BAC) library sions in the amaranth germplasm collection
was constructed from A. hypochondriacus con- include representatives from 12 amaranth spe-
taining about 37,000 clones and an anticipated cies. The collection includes germplasm from
genome coverage of more than tenfold. banks in other countries, material from germ-
Microsatellite markers developed from A. hypo- plasm collection trips and materials which have
chondriacus and A. tuberculatus would be valu- been donated by collaborating researchers. The
able for more detailed study on phylogeny and germplasm is catalogued according to ‘grain
breeding efforts. These markers were shown type’. The grain type categories may actually be
transferrable to other cultivated as well as weedy a regrouping of the germplasm according to land-
amaranth species. Besides, shotgun sequence race. The grain type categories have proven to be
data from A. tuberculatus and collection of useful to manage the huge amount of variability
recombinant lines (RIL) derived from initial that exists within each species. Descriptions of
weed-crop cross between A. hypochondriacus all species and the grain types can be found in the
and A. hybridus can also be very useful for the RRC germplasm catalogue (Kauffman and
development of amaranth genetic map. Reider 1986). To meet the problem of genetic

100 7 Distribution and Maintenance of Amaranth Germplasm Worldwide
erosion, Amaranthus germplasm has been col- 1993), but later three new lines have been devel-
lected for ex situ conservation (Grubben and Van oped in China (Wu 1998). Three cultivar lines,
Sloten 1981). Field studies have been conducted namely, ‘Vietmeyer’, ‘Oscar Blanco’ and ‘Alan
by Kauffman during 1978 and 1979. In 1978 Garcia’, were released through selection in Peru
yields were tested on 23 of the newly selected (Summar et al. 1992). Covas (1991) developed
lines. Most of the lines were selected from the five cultivars in Argentina. Bansal (1996)
Mexican lines. The yield levels ranged from 5.8 described several cultivars in India, where
to 18.8 q/ha. In California Davis Jain et al. (1984) ‘Plainsman’ replaced ‘Annapurna’ as the top
reported grain yield of some selection as high as yielding line.
36 q/ha (National Research Council 1984). Jorge The germplasm collections are maintained in
Mario and Bressani (1987) while testing eight at least 61 collections (IPGRI 1999). The collec-
grain amaranth selections reported a grain-yield tions are gradually becoming more and more
range from 20.3 to 38.q/ha in Guatemala. The important because of their accessibility and doc-
highest grain yields were recorded in 20-USA umentation through the Internet. Most collec-
(Rodale Selection of A. cruentus) followed by tions have less than 100 accessions except six
28-USA (Rodale Selection). In China and collections which are maintaining quite a large
Mongolia, maximum grain yields have been number of accessions (Table 7.1). Amaranthus is
reported to be 5500 kg/ha and 3400 kg/ha, respec- well adjusted to ex situ conservation due to their
tively. Four amaranth cultivars have been regis- small and long-lived seeds (Kigel 1994). Brenner
tered in the USA – ‘Montana 3’ (MT-3), ‘Montata and Widrlechinar (1998) described an efficient
5’ (MT– 5), ‘Amont’ and ‘Plainsman’ (Schulz- protocol for regenerating seeds of Amaranthus
Schaffer et al. 1991; Baltensperger et al. 1992). germplasm and maintenance of its genetic integ-
Several lines have been developed by the RRC, rity in ex situ situation.
Nu-World Amaranth and American Amaranth A majority of the accessions in the RRC germ-
that have been widely distributed and evaluated plasm collection were collected as mass selec-
but never registered. All registered cultivars were tions or single plant selections of cultivated
developed by the RRC; ‘MT-3’ was a selection landraces of those species which are commonly
from RRC 1041, ‘MT-5’ was selected from RRC grown for their light-coloured seed. Many selec-
425 and ‘Amont’ was a selection from ‘MT-3’. tions have been made from the segregating acces-
‘Plainsman’ was a selection from the cross RRC sions in an effort to create uniform lines. Seed
1024 x RRC 1004 and widely distributed and from the amaranth accessions has been distrib-
treated as ‘K 343’ prior to release. Plainsman has uted to thousands of researchers and farmers
become the most widely grown amaranth cultivar around the world. The germplasm collection is
in the USA due to its relatively high yield poten- the backbone of the varietal improvement pro-
tial, lodging resistance, limited seed shattering, gramme which is aimed at developing agronomi-
seed colour and maturity range. Yield variation cally acceptable lines using classical plant
was high with Plainsman ranging from 2500 kg/ breeding and selection methods.
ha in Colorado in 1991 down to 220 kg/ha in Germplasm characterisation is being con-
Missouri in 1990, typically in the range of ducted at a number of locations around the world.
700 kg/ha to 1700 kg/ha considering all the Extensive, well-documented amaranth germ-
entries over the year and locations tested. Several plasm characterisation has been conducted by
cultivars have been developed through the world organisations in India, Peru and Mexico. Since
including Russian, ‘Pastevny 1’, ‘Turkestan’ and 1982, the staff at RRC has collaborated with
‘Ural’, and South America’s A. cruentus geno- researchers who are conducting observations on
type ‘Anden’ (Kaul et al. 1996). None of these 14 selected grain amaranth accessions (represent-
cultivars have been widely tested in the USA. The ing grain types from the species A. hypochondri-
main cultivars in China are five RRC lines (Corke acus L., A. cruentus L., A. caudatus L. and A.
et al. 1997) especially RRC 1011 (Yue and Sun hybridus L.) to collect information on their
7.2 Germplasm Collection Centres in the World 101
Table 7.1 Six largest ex situ Amaranthus germplasm collections in the world
Institute No. of accessions Year updated References
1. Institute of Crop Germplasm Resources 438 1996 IPGRI (1999)
(CAAS)Beijing, China
2. Universidad Nacional del Altiplano 440 1990 IPGRI (1999)
Puno, Peru
3. Instituto Nacional de Investigaciones 495 1993 IPGRI (1999)
Forestales y Agropecuarias (INIFAP)
4. Univ. Nacional San Antonio Abad del 740 1990 IPGRI (1999)
Cusco (UNSAAC/CICA)
5. National Bureau of Plant Genetic 3000 1995 Joshi (1985)
Resources
6. North Central Regional Plant 3380 1999 USDA, ARS (1999)
Introduction Station (NCRPIS), Ames,
Iowa, USA
performance at distinctly different climates and Introduction, Indian Agricultural Research

latitudes (Bressani et al. 1987; Senthong 1986; Institute, New Delhi (now National Bureau of
Gupta 1986; Duriyaprapan 1986; Espitia 1986). Plant Genetic Resources or NBPGR), which then
Information generated at each location serves as held a small germplasm collection (less than 50)
a starting point to help determine which sources at Shimla Station, assembled from indigenous
of germplasm should be exploited to develop sources as well as from a few foreign countries,
improved lines of grain amaranth for any given viz. Nepal and the USA. It was only in 1978
area. Similar observations have also been made onwards when more efforts were directed towards
to characterise germplasm which was collected germplasm collection and evaluation of the exist-
throughout India (Joshi 1986). Populations of ing genetic resources to enrich these collections
landraces which were collected from Mexico to and to analyse and study the potential of this
Argentina have been observed for their genetic dual-purpose crop (Joshi and Rana 1991). A great
structure (Hauptli and Jain 1984). deal of variability was observed, especially in
Germplasm enhancement is being conducted inflorescence, leaf and stem colour; inflorescence
at RRC since 1977 using recurrent single plant size and shape; inflorescence and stem branching
selection and mass selection with an intention to pattern; axillary inflorescence branching pattern;
develop ideotypes which would meet the needs plant height, spiny/glabrous nature of the bracts;
of modern agriculture. seed colour, size, weight and transparency; termi-
The USDA Plant Introduction Center at Ames, nal inflorescence length and shape; and maturity.
Iowa, has been working to characterise germ- The collected amaranth germplasm includes
plasms and to determine the amount of outcross- nearly 2722 indigenous accessions and 293
ing that has occurred. This Plant Introduction exotic accessions. Now the total amaranth germ-
Center is of great significance in maintaining a plasm collection of Shimla Regional Station is
large number of amaranth germplasm collected 3081 comprising mainly of grain amaranth
from all over the world and their distribution to (Table 7.4). Apart from exchange of germplasm
promote amaranth research (Tables 7.2 and 7.3). within India, exchange of seeds under physiosan-
USDA, Ames, Iowa, USA, also published an ity vigilance between India and other countries
amaranth catalogue of 2783 collections with ori- are being encouraged by NBPGR making
gin and grouped them under respective species of exchange links with about 70 different countries.
the genus Amaranthus (Brenner 1990). It is linked with several international institutes
In India initial efforts to study grain amaranths like IRRI, ICARDA, IITA, CIAT, AVRDC,
were made in the 1960s by the Division of Plant CSIRO, USDA and IBPGR.
Table 7.2 Various accessions of Amaranthus maintained Table 7.2 (continued)

at Plant Introduction Center at Ames, Iowa
Sl. Number of
Sl. Number of no. Amaranthus species accessions
no. Amaranthus species accessions 30 Amaranthus fimbriatus 4
1 Amaranthus spp. NC7-amaranth 41 NC7-amaranth
2 Amaranthus albus NC7-amaranth 7 31 Amaranthus floridanus 1
3 Amaranthus hybridus 154 NC7-amaranth
NC7-amaranth 32 Amaranthus graecizans 14
4 Amaranthus blitum NC7-amaranth 10 NC7-amaranth
5 Amaranthus dubius NC7-amaranth 43 33 Amaranthus retroflexus 24
6 Amaranthus crispus NC7-amaranth 1 NC7-amaranth
7 Amaranthus greggii NC7-amaranth 2 34 Amaranthus tenuifolius 1
NC7-amaranth
8 Amaranthus palmeri NC7-amaranth 15
35 Amaranthus tucsonensis 1
9 Amaranthus pumilus 7 NC7-amaranth
NC7-amaranth
36 Amaranthus californicus 1
10 Amaranthus torreyi NC7-amaranth 1 NC7-amaranth
11 Amaranthus viridis NC7-amaranth 18 37 Amaranthus polygonoides 1
12 Amaranthus caudatus 557 NC7-amaranth
NC7-amaranth 38 Amaranthus standleyanus 2
13 Amaranthus cruentus 365 NC7-amaranth
NC7-amaranth 39 Amaranthus tuberculatus 51
14 Amaranthus deflexus 5 NC7-amaranth
NC7-amaranth 40 Amaranthus acanthochiton 2
15 Amaranthus hybridus 191 NC7-amaranth
NC7-amaranth 41 Amaranthus tamaulipensis 1
16 Amaranthus powellii 18 NC7-amaranth
NC7-amaranth 42 Amaranthus hypochondriacus 1523
17 Amaranthus spinosus 24 NC7-amaranth
NC7-amaranth
18 Amaranthus tricolor NC7-amaranth 182
19 Amaranthus watsonii 1 Table 7.3 Various accessions of Amaranthus available at
NC7-amaranth Plant Introduction Center at Ames, Iowa for distribution
20 Amaranthus wrightii 2 Sl. Number of
NC7-amaranth no. Amaranthus species accessions
21 Amaranthus arenicola 7 1 Amaranthus albus NC7-amaranth 7 7
NC7-amaranth 2 Amaranthus hybridus NC7-amaranth 153
22 Amaranthus asplundii 1 153
NC7-amaranth 3 Amaranthus blitum NC7-amaranth 10 10
4 Amaranthus dubius NC7-amaranth 31 31
23 Amaranthus australis 2 5 Amaranthus crispus NC7-amaranth 1 1
NC7-amaranth 6 Amaranthus greggii NC7-amaranth 2 2
24 Amaranthus blitoides 7 7 Amaranthus palmeri NC7-amaranth 15 15
NC7-amaranth 8 Amaranthus viridis NC7-amaranth 17 17
25 Amaranthus crassipes 2 9 Amaranthus caudatus NC7-amaranth 543
NC7-amaranth 543
26 Amaranthus muricatus 1 10 Amaranthus cruentus NC7-amaranth 335
NC7-amaranth 335
11 Amaranthus deflexus NC7-amaranth 5 5
27 Amaranthus quitensis 51
12 Amaranthus hybridus NC7-amaranth 184
NC7-amaranth
184
28 Amaranthus acutilobus 2 13 Amaranthus powellii NC7-amaranth 16
NC7-amaranth 16
29 Amaranthus cannabinus 3 14 Amaranthus spinosus NC7-amaranth 23
NC7-amaranth 23
(continued) (continued)
7.2 Germplasm Collection Centres in the World 103
Table 7.3 (continued) Table 7.4 Germplasm of amaranths maintained at

Sl. Number of Regional Station Shimla, NBPGR
no. Amaranthus species accessions No. of
15 Amaranthus tricolor NC7-amaranth 181 Sl. no. Species of Amaranthus accessions
181
1. Amaranthus hypochondriacus 2452
16 Amaranthus watsonii NC7-amaranth 1
1 2. A. cruentus 556
17 Amaranthus wrightii NC7-amaranth 2 2 3. A caudatus 27
18 Amaranthus arenicola NC7-amaranth 6 4. A. edulis 08
6 5. A. dubius 10
19 Amaranthus asplundii NC7-amaranth 1
1 6. A. hybridus 09
20 Amaranthus australis NC7-amaranth 2 7. A. viridis 05
2 8. A. retroflexus 07
21 Amaranthus blitoides NC7-amaranth 7 9. A. lividus 03
7
10. A. tricolor 04
22 Amaranthus crassipes NC7-amaranth 2
2
23 Amaranthus muricatus NC7-amaranth 1
1 Apart from NBPGR, Regional Station Shimla,
24 Amaranthus quitensis NC7-amaranth 49 collections have also been made by the
49 Vivekananda Parvatiya Krishi Anusandhan Shala
25 Amaranthus acutilobus NC7- 2 (VPKAS) Almora, GB Pant Agricultural
amaranth 2
26 Amaranthus cannabinus NC7- 2 University, Ranichauri, Tehri Garhwal, North
amaranth 2 Eastern Hill Complex, Shillong, Maharashtra and
27 Amaranthus fimbriatus NC7-amaranth 4 Gujarat State in India. Collection activities have
4 also been reported for grain amaranth germplasm
28 Amaranthus floridanus NC7-amaranth 1
1 including IBPGR (1984)-supported explorations
29 Amaranthus graecizans NC7- 14 in Nepal, Bhutan, Thailand, Indonesia, Ethiopia,
amaranth 14 Zambia, Nigeria, Kenya, Argentina, Bolivia,
30 Amaranthus retroflexus NC7- 24 Ecuador, Mexico, Guatemala and Peru. The
amaranth 24
31 Amaranthus tucsonensis NC7- 1 global germplasm collections have been stored in
amaranth 1 the National Seed Storage Laboratory, Fort
32 Amaranthus californicus NC7- 1 Collins, Colorado, USA, where IBPGR has des-
amaranth 1 ignated NBPGR as a regional base centre for
33 Amaranthus polygonoides NC7- 1
amaranth 1 amaranth germplasm collection in view of pio-
34 Amaranthus standleyanus NC7- 2 neering work done on this crop by this station.
amaranth 2 The evaluation of indigenous and exotic germ-
35 Amaranthus tuberculatus NC7- 50 plasm collections of grain amaranths over several
amaranth 50
36 Amaranthus acanthochiton NC7- 2 years led to the identification of strains/acces-
amaranth 2 sions which offer good opportunities for its utili-
37 Amaranthus tamaulipensis NC7- 1 sation and improvement (Annual Report 1981 to
amaranth 1 1988, NBPGR, Regional Station, Phagli, Shimla).
38 Amaranthus hypochondriacus 1497
NC7-amaranth These have been multiplied and maintained sepa-
rately for distribution to the persons involved in Table 7.5a Amaranth germplasm collection at Tamil
crop improvement and those who are interested Nadu Agricultural University, Coimbatore
to take up inheritance studies. The most promising Sl. No. of accessions
genotypes in indigenous and exotic material of no Species of Amaranthus maintained
interest to breeders include IC-38541, IC-38577, 1 Amaranthus dubius 18
EC-170304 and EC-169626 (dwarf types, 2 Amaranthus tricolor 22
68–81 cm); IC-38137, EC-38323 and EC-151544 3 Amaranthus tristis 6
4 Amaranthus 14
(high number of spikelets, 65–95); IC-38052,
hypochondriacus
IC-38508 and EC-157415 (high inflorescence
5 Amaranthus cruentus 19
length, 78–95 cm); IC-5626, IC-7934 and 6 Amaranthus paniculatus 18
IC-38 l36 (hard threshability); IC-38133, Total 97
EC-157413 andBDJ86–329 (early flowering
40–52 days); IC-38133, IC-38422 and
EC-1574 l7 (early maturing 100–110 days); trials at Shimla and Almora during 1986. The
IC-38l3l, IC-386 l1, IC-38665 and BDJ-86–259 highest grain yield (34.8 q/ha) was recorded for
(bold seeded 1000 grain weight more than 1 ‘Annapurna’ followed by VL-21 (32.5 q/ha) and
gramme); and IC-38269, IC-38280, IC-42258–1, IC-42290–17 (30.9 q/ha). Multilocation trials of
VL-21 and BDJ86–129 (high grain yield per 14 promising varieties were conducted at eight
plant). A very high yielding variety of grain ama- centres during 1988–1989. The varieties that
ranth A. hypochondriacus named ‘Annapurna’ appeared to be significant were S.K. Nagar,
was developed in the National Bureau of Plant ‘Annapurna’, ‘Akola local’, A. edulis (ex Taiwan),
Genetic Resources (NBPGR) Regional Station, IC-5564, etc. with varying yield amounts at dif-
Shimla, through screening germplasms of 2700 ferent locations.
collections and multilocation trials after a con- In South India, Tamil Nadu Agricultural
tinuous research of 14 years (Joshi et al. 1983). It University, Coimbatore, is a prominent centre of
has given an average grain yield of 22.3 q/ha. and amaranth germplasm collections, specially vege-
34 q/ha of seed yield in Shimla condition. The table amaranths (Tables 7.5a and 7.5b), which
variety has few distinguishing features like tall released few improved varieties of vegetable
habit, medium or late maturing, dark-green broad amaranths.
leaves, yellowish-green inflorescence with long
terminal and lateral spikelet, about 74 cm long
and creamish-white seed with 14.5 % protein 7.3 Genome Resource
content. The popping quality of the seeds is Development of Amaranthus
excellent, about six times of the seed size. The
other significant varieties of grain amaranth like Amaranth may serve as one of the best model
R 104, 20 USA, Jumla, VL-21 and S.K. Nagar systems of genome investigation of weediness
have also been developed and employed in culti- (Basu et al. 2004; Chao et al. 2005). Amaranth
vation in different parts of the world. The variety possesses several characteristics which made
‘Annapurna’ has a wide adaptability and can be them a desirable model system:
grown in rain-fed land in the hills as well as in the
drought-prone areas and also under Himalayan 1. Most species are functional diploid having
watershed. It has given 68.9 % higher increase n = 16 or 17 (exception A. dubius is a poly-
in grain yield over VL-21, another promising ploid with 2n = 64). Amaranthus species have
selection from Almora, Uttarakhand. All India detectable genome size 3 to 4 times that of
Coordinated Research Project has recommended model plant Arabidopsis thaliana.
it as a national check (Joshi and Rana 1991). 2. Amaranthus species are generally easy to cul-
Nine promising varieties selected from landraces ture and manipulate under greenhouse or
of grain amaranths were evaluated by repeated other experimental condition.
7.3 Genome Resource Development of Amaranthus 105
Table 7.5b Few new cultivars/varieties of vegetable amaranths were released from vegetable research station, TNAU
No. of accessions at
Sl. no Amaranthus species VRS, Palur Remarks
1. Amaranthus polygonoides 8 Called Sirukeerai in Tamil
A variety named PLR1 was released in 2013
from this university
2. Amaranthus tristis 2 Called Araikeerai in Tamil
A variety named CO3 was released in 2013
from this university
3. Amaranthus tricolor 7 Called Mulaikeerai/Thandukeerai in Tamil
Varieties: CO2 and CO5
3. As a typical weed, Amaranthus spp. exhibit tive markers to study genetic variability. The mic-
plasticity in adaptability in diverse environ- rosatellite markers will be of great value to
mental condition. investigate queries related to gene flow, evolution
4. The monoecious species are readily self- and hybridisation within and among Amaranthus
pollinated but outcross, and F1 hybrids are weeds. The microsatellite markers can also be
easily obtainable if selectable (herbicide resis- used to construct genetic linkage map from map-
tance), detectable or scorable (pigmentation) ping populations and for multilocus genome scan-
markers are present in one of the parents. ning to identify genetic target of selection (Smith
5. Production of a large number of small seeds. and Haigh 1974). Microsatellite markers have
been developed from A. hypochondriacus (Lee
The first genome-based resource derived specifi- et al. 2008; Mallory et al. 2008), and markers
cally from Amaranthus species is now available. A were demonstrated to be transferable to the other
bacterial artificial chromosome (BAC) library was cultivated as well as weedy amaranth species.
constructed from Amaranthus hypochondriacus Nearly 400 unique microsatellite markers were
(Maughan et al. 2008). It can be used as a tool for obtained primarily from microsatellite-enriched
isolation and identification of the full-length gene libraries but also from BAC-end sequence data.
sequence. This library contains about 37,000 clones About 180 of these proved polymorphic among
and a presumed genome coverage greater than ten- three grain amaranths. Many of these markers are
fold. Thus any specific DNA sequence of A. hypo- transferable to the dioecious amaranth species as
chondriacus has greater than 99 % chance of being well. A preliminary phylogenetic analysis using
represented in this library. A full-length genome some of these markers placed A. hybridus within
sequence for desirable target site of the gene ALS multiple grain amaranth clades, suggesting mul-
and PPX2 can be obtained. Due to the sequence tiple domestication events from A. hybridus
similarity among the Amaranthus species, the BAC (Mallory et al. 2008). Additional microsatellite
library from A. hypochondriacus can be utilised for markers were obtained recently from A. tubercu-
genome-based investigation of other weedy species latus (Lee et al. 2009); collectively these micro-
as for example, transferability between A. hypo- satellite markers will be valuable for more detailed
chondriacus and other weed species. The BAC phylogenetic studies and breeding effects.
library can be used to generate a physical map of the The third genome resource obtained recently
Amaranthus genome, which ultimately can enable is shotgun sequence data from A. tuberculatus
map-based cloning of Amaranthus genes of interest (Lee et al. 2009). Random sequencing was done
(Maughan et al. 2008). using massively parallel pyro-sequencing tech-
The second genome resources available are a nology (Margulies et al. 2005). From a single
set of microsatellite marker. The microsatellite pilot sequencing run, approximately 160,000
markers are one of the most robust and informa- sequencing reads were obtained within an aver-
age read length of about 270 nucleotides, yield- The fourth resource is a collection of recombi-
ing a total of about 43 million nucleotides of A. nant inbreed lines (RILs) derived from an initial
tuberculatus sequence data. The data sheet crop-wild cross between A. hypochondriacus and
included nearly a complete sequence of the A. hybridus, selected based on herbicide resis-
chloroplast genome and partial sequence of tance. They can be the useful source to provide
most currently known herbicide target-site ideal populations for development of an ama-
genes. ranth genetic map.
Breeding of Amaranths
8
8.1 General transfer of feature like herbicide resistance from

a weed species to another weed species of ama-
Amaranths are characterised with remarkable ranth. The crossability barrier between grain
germplasm diversity, adaptability to different amaranths and their weedy relatives are very
growing conditions and unique matting behav- fragile showing variable frequency of cultivar-
iour ranging from obligate outcrossing (dioe- weed outcrossing. The use of interspecific ama-
cious species) to greater outcrossing to greater ranth hybrids appears to be promising to increase
self-pollination. The breeding mechanism in biomass productivity. In the case of vegetable
amaranths is variable due to variability and versa- amaranths used as parents, heterosis from inter-
tility of inflorescence, ratio and distribution of specific hybridisation may double to quadruple
male and pistillate flowers in inflorescence. the biomass of the parents. Crosses between
Breeding work in amaranths is to be adopted grain species and their weedy relatives may yield
giving priority to traits like non-shattering seed, sterile hybrids but with increased biomass. Grain
reduced plant height, high yield, increased seed yield heterosis has been estimated in A. hypo-
size, synchronised drydown of plant, taste and chondriacus. Male sterility found only in A.
nutritive value, improved pest resistance/or toler- hypochondriacus could be used to simplify
ance, low content of anti-nutrient factors, etc. hybridisation. High heterozygosity and low heri-
The genetic improvement of grain amaranths has tability in some traits are some of the problematic
been done so far mostly applying conventional areas which breeders face in amaranth breeding
selection methods from local collections or land- that are to be solved. Wide range of genetic vari-
races, and new varieties or improved lines are ability in cultivated landraces of some grain spe-
released. In amaranth breeding, hybridisation has cies offers much scope for its improvement
been proved as the most effective breeding pro- through selection. Molecular markers are very
gramme to produce new improved variant with useful for diversity analysis to distinguish geneti-
useful trait. Naturally occurring hybrids between cally similar or distinct accessions or landraces.
A. caudatus and A. cruentus and A. hybridus Marker-guided selection method can assist to
were identified by intermediate characters. One select individual carrying molecular markers
of the most popular and widely utilised grain associated with the trait of interest. Mutation
amaranth varieties in the USA, A. hypochondria- breeding has a great scope in amaranth to induce
cus var. Plainsman, was obtained through a cross variability using radiation. Drought tolerance in
with A. hybridus accession from Pakistan having A. tricolor and increased yield potential in A.
a feature of earliness. Interspecific hybridisation caudatus were induced through gamma radia-
techniques have been employed to study the tion. Three mutant varieties have been developed

108 8 Breeding of Amaranths
and registered in MVD, viz. centenario in Peru, like reduced plant height, reduced branching and
New Asutake in Japan and Sterk in Russian non-shattering seeds facilitate easy grain harvest-
Federation. Polyploidy was induced in A. cauda- ing. For vegetable amaranths branching is desir-
tus, A. tricolor, A. hypochondriacus and A. edulis able. Vegetable cultivars with more branching are
by colchicine treatment with positive effect. suitable for multiple cutting. This produce more
Tetraploid A. caudatus had 60 % more protein, total biomass yield compared to unicut cultivars
more lysine and threonine than diploid. (Sreelathakumary and Peter 1993).
Polyploidy increased grain size and weight not In most of the species of Amaranthus investi-
decreasing the fertility or nutritive value. gated so far, nearly 40 have a diploid number of
Amphidiploidy was induced in fertile F1 hybrids chromosome equal to 32 or 34, and the basic
between A. caudatus and A. retroflexus which chromosome number is 16 or 17 (Madhusoodanan
were perfectly fertile and phenotypically alike. In and Pal 1981). There are few tetraploid species
plant breeding, genetic engineering involves arti- like A. caudatus and A. dubius having basic
ficial incorporation of foreign genetic materials chromosome number either 32 or 34. Natural
followed by selection. Agrobacterium-mediated polyploids occur in wild amaranths; Amaranthus
transformation protocol in A. tricolor and A. dubius is a naturally occurring tetraploid.
hypochondriacus has been standardised. This Induced polyploid forms of A. dubius and grain
achievement made it possible to reduce the anti- amaranth A. hypochondriacus have been
nutrient oxalate level in vegetable amaranths obtained through colchicine treatment of the
through the insertion of cDNA of oxalate decar- seeds and seedlings.
boxylase gene. The insertion of oxalate decar- Broadly speaking, the genus Amaranthus
boxylase gene in tomato has been possible shows three types of mating system, viz. (a) obli-
through Agrobacterium-mediated transforma- gate outcrossing in dioecious species, (b) rela-
tion. A storage protein gene AmM1 isolated from tively greater outcrossing in monoecious member
A. hypochondriacus can be introduced in any of sect. Amaranthus and (c) relatively greater
other crop through genetic transformation as has self-pollination in members of sect. Blitopsis.
been done in potato and wheat. Plant tissue cul- Dioecious species are confined to a small area in
ture can be very useful though not utilised so North America (Sauer 1955, 1957), though spo-
much in amaranth breeding. Variability in germ- radic appearance has been reported from time to
plasms can be used for screening cell lines in time (Aellen 1961; Brenan 1961; Thakur 1964).
search of plant more tolerant to stress, a potential The narrow geographic range of the dioecious
source of herbicide resistance. Such traits can be species can be attributed to the fact that, for any
introduced into grain and vegetable forms via long-distance dispersal, at least two propagules
protoplast fusion. (male and female) are necessary before a particular
species can successfully establish and colonise
the new area. The second reason may be that few
8.2 Breeding Behaviour plants which supposed to have escaped their
in Amaranths native place are often involved in interspecific
hybridisation with monoecious amaranths and
Amaranths have been subjected to a little domes- lead to the totally sterile hybrids. These perhaps
tication pressure as compared to major crops. It is were the reasons for reduction of their reproduc-
notable for its remarkable germplasm diversity tive potential and ultimate elimination.
and adaptability to different growing conditions. The monoecious species of Amaranthus are
Several characteristics are key attributes for self-compatible. They are nearly cosmopolitan
success in the agronomic use of amaranths. species, often weedy and seem to have achieved a
According to Kauffman (1982), varieties of grain significant balance between fitness and long-
amaranths should have synchronised drydown of range flexibility. The breeding mechanism of
the entire plant at maturity. The characteristics Amaranthus is complex and variable due to
8.2 Breeding Behaviour in Amaranths 109
variability and versatility of inflorescence. The the other hand, the single staminate flower per
basic units of the inflorescence are little dichasial glomerule and big inflorescence should lead to
cymes, usually called glomerules, each ordinarily more cross-pollination in the sect. Amaranthus
consisting of an initial male or staminate flower (the grain amaranths). However, grain amaranths
and an indefinite number of female or pistillate are also basically self-pollinated with 0–34.9 %
flowers. The glomerules are crowded on a leafless cross-pollination which may be helped by insect
axis to form complex inflorescence, technically pollinators (Mohideen and Irulappan 1993). The
thyrses, which are generally called spikes. A main inflorescence in sect. Amaranthus is a dense
completely developed glomerule may have as spike or panicle, which is usually profusely
many as 250 flowers, while completely pollinated branched. The panicles or spikes may be either
flowers are represented in lesser number. In all drooping (A. caudatus), semidrooping (A. spinosus,
the grain species, each flower is subtended by a A. cruentus, etc.) or erect (A. hypochondriacus
sharp-pointed bract. The perianth consists of five and A. edulis). The growth of the terminal
free ‘tepals’; the male flowers characteristically inflorescence and its branches is indeterminate
have five stamens, and the female flowers have a (racemose), and thus, it may reach a length of one
single circumscissile utricle. metre or even more. However, A. edulis is the
The main axis of the inflorescence is usually only exception in which the inflorescence is
branched. The length and number of these determinate due to the presence of a terminal
branches and their angle with the main axis polymerous male flower, and therefore, the elon-
determine the shape of the inflorescence. The gation of the inflorescence in this species is due
clusters of individual flower develop along the to intercalary growth of the axis. Individual
axis in an alternate fashion. The first or primary flower clusters or glomerules develop alternately
branch of inflorescence is terminated by the first along the axis of the main inflorescence. In sect.
flower. From the base of the primary branch, two Amaranthus glomerules develop chiefly in com-
branches arise, which are terminated by the sec- pact fashion on the terminal inflorescences and
ond and the third flowers. At its base the next two are rather feebly developed in the axils of the
flowers arise and the process continues until all branches. In the sect. Blitopsis, due to the absence
the available space is occupied. The monoecious of prominent terminal inflorescence, axillary
species exhibit two types of arrangements of the glomerules are well developed and resemble
staminate and pistillate flowers. These types are glomerules in the terminal inflorescence of sect.
important because of their different breeding Amaranthus. The growth pattern of the individual
behaviour. In the first type, the first flower of each dichasia, as described above, is common to all
flower cluster is a staminate and all the secondary species irrespective of the sex. As expected in
ones are pistillate. There is only one staminate dioecious species, all the glomerules on a plant
flower in each flower cluster of the inflorescence have the flowers of the same sex. The breeding
and that withers soon after shedding pollen. All pattern in grain amaranths varies from high rates
the species, except A. spinosus, belong to this of selfing (over 90 %) to mixed mating with as
group. In the second type, all the pistillate flow- much as 30 % outcrossing (Jain et al. 1982).
ers develop only in the axis of the branches and at Outcrossing rates vary significantly with geno-
the base of the terminal inflorescence, while the type and environmental factors between years. A
clusters of staminate flowers are borne terminally bidirectional selection experiment based on male/
on the main axis and lateral branches. The spe- female flower ratio per plant suggested that
cies A. spinosus belongs to this type. breeding system is under genetic control, and
Species included in the sect. Blitopsis (the thus, it can be modified to suit breeding procedures
vegetable amaranths) have a number of staminate for either inbreeding or outbreeding species.
flowers per glomerule and small axillary inflores- Kauffman (1979) while studying the floral
cence, resulting to predominant self-pollination morphology reported that each head of amaranth
of these species (Pal and Khoshoo 1973a, b). On consists of hundreds of floret arranged in a
panicle. Each floret consists of three to six flowers tion, it is possible to control the outcrossing
with one staminate (male) flower surrounded by rate and to raise true-to-type lines from segre-
several pistillate (female) flowers. Occasionally, gating breeding lines involving only a few gen-
a floret consists of only female flowers. Anthesis erations of selection.
begins when the stigmas of the female flowers The cultivated grain amaranths are monoe-
exert. Available pollen does not come from the cious. The basic emasculation and pollination
male flower in the same floret but from the stami- techniques for breeding amaranths have been
nate flowers of other florets. Due to the fact that described (Murray 1938) and refined (Kauffman
flowers of both the sexes are located on the same 1981a, b). Later concrete programmes are being
inflorescence, the potential for selfing is followed at the University of California-Davis to
increased. So breeding behaviour in amaranth, identify genetic traits to be emphasised for
featured with variable sex ratio within glomer- improving grain amaranths (Jain et al. 1984).
ules and inflorescence as a whole, and variable Efficacy of different breeding strategies has been
outcrossing rate attributed to bidirectional selec- determined and evaluated (Kulakow and Jain
tion in amaranth breeding. Such unique genetic 1987; Ayiecho 1986). Applicability of additional
features in breeding system have great impact on techniques has been evaluated keeping in view
the evolution of amaranth species and domestica- the manipulation of plant height and increase in
tion of landraces. grain yields (Vaidya and Jain 1987).
The outcrossing rate was found to be nega- Research programme carried out at the
tively correlated with the ratio of staminate to University of California-Davis and other loca-
pistillate flowers within an inflorescence under tions related to the development of hybrids has
strong environmental influence (Hauptli and enriched the breeding work by the identification
Jain 1985). Interspecific hybrid is formed as a of gene markers for the useful traits like time of
result of cross-pollination of two different spe- flowering, determinate vs. indeterminate growth
cies. Natural interspecific hybridisation occurs pattern, plant and seed pigment patterns and leaf
frequently in amaranths, eight naturally occur- characteristics (Kulakow et al. 1985; Kulakow
ring interspecific hybrids have been reported in and Jain 1985; Kulakow 1987). An additional
vegetable amaranth species, and they show vary- marker like bract length has been identified in
ing degrees of sterility. Interspecific hybrid of A. breeding trials at RRC. Okuno and Sakaguchi
lividus and A. tricolor showed an even higher (1982) have identified the perisperm and grain
degree of hybridisation defects, with about starch of grain amaranths as potential gene mark-
90 % of pollen sterility, because of the occur- ers. Markers are useful to trace and reject self-
rence of two or more interchanges of several pollinated plants in the F1 generation and also to
chromosomes and subsequent meiotic abnor- allow breeders to avoid the complex process of
malities (Sreelathakumary and Peter 1993). On emasculation in crossing event. The identifica-
the other hand, interspecific hybridisations tion of male-sterile lines in A. hypochondriacus
between grain amaranths and vegetable ama- has further eased the way for getting hybrids as
ranths often produce hybrids with malforma- additional hybridisation techniques (Peters and
tion, high pollen sterility and chromosome Jain 1985; Peters and Jain 1987; Gudu and Gupta
aberrations, suggesting the presence of a great 1988a).
genetic barrier between species included in the In India efforts have been devoted to learn more
sections Amaranthus and Blitopsis (Mohideen about traits which govern yield potential including
and Irulappan 1993). The gene pool of harvest index, weight per 1000 seeds and yield per
Amaranthus provides an excellent source of plant (Pandey 1984b). Investigations have also
materials for geneticists to work with. been directed to identify the inheritance pattern of
Amaranths are predominantly self-pollinating trait that governs nutritional characters of grain.
crop, with varying outcrossing rate (Hauptli The factors regulating starch characteristic of
and Jain 1985). By growing amaranth in isola- grain were explored in Japan (Okuno 1985;
8.2 Breeding Behaviour in Amaranths 111
Konishi et al. 1985). In India also the inheritance • High protein content
pattern of grain protein percentage has been inves- • Functional attributes of the grain
tigated (Pandey and Pal 1985).
A breeding programme was stated in 1978, in Montana Agricultural Experiment Station first
Rodale Research Centre (RRC), USA, to raise released an improved line of grain amaranth
improved plant types for commercial production named ‘MT-3’ (Cramer 1988) selected from seg-
in the USA and Canada. A series of improved regating accessions of A. cruentus L. originally
breeding lines (F6 generation and beyond) was collected in Mexico. Similar programmes were
generated and grown at experimental station and carried out in Mexico, Peru and Kenya. In Peru,
commercial farm in the USA, applying classical the University of Cuzco has released three culti-
breeding methodologies. The lines were included vars of A. caudatus L. named as ‘Oscar Blanco’,
and distributed as a part of ‘K-series’. One of ‘Noel Vietmeyer’ and ‘Alan Garcia’. These three
such lines named ‘K-343’ is a derivation of cross- cultivars are now being grown on commercial
ing between white-seeded A. hypochondriacus scale by the farmers in Peru (Sumar K., L. per-
from Mexico and a black-seeded weed A. hybri- sonal communication, 1988). A cultivar of A.
dus from Pakistan. This single accession was hypochondriacus L., named ‘Annapurna’, has
widely utilised in the Great Plains of the USA in been released by NBPGR in India.
1988 and grown on commercial scale. It is char- The concentration on only one source of
acterised with features like a short unbranched germplasm is not helpful for improvement of the
plant habit at high plant densities and slightly crop. Additional germplasm resources are to be
early maturity, as compared to ‘MT-3’ and other explored and evaluated to initiate and expand
selected Mexican landraces which gained popu- applied breeding programmes for expected
larity among the farmers. increase in grain amaranth production. The seed
Scientists at RRC have identified a number of certification process initiated in 1988 will be
easy to isolate traits in order to proceed with the helpful to address the need for sources of
genetic improvement programme, which include improved quality seed of grain amaranth.
the following: The future improvement of amaranth is pri-
marily dependent on the appropriate selection of
• Reduced stature of plant (1.0–1.5 m) genotypes from the existing germplasm collec-
• Flowering above the leaf canopy tions. We have to explore unusual and minor
• Lack of branching sources of germplasm also to find traits such as
• White or gold seed-coat colour early maturity and shorter plant stature. A dark-
• Maturity between 100 and 120 days seeded accession (‘African’ grain type) of A.
• Higher yield due to traits like a large flower cruentus from West Africa, having a feature of
head with a high ratio of pistillate flowers that greater branching, was utilised as a source for
set seed early maturity. A source for short plant stature
trait was found among a few highly branched,
However, a number of other traits still need to be shattering, dark-seeded A. hybridus L. (‘Prima’
improved: grain type) accessions from Asia.
There are some traits such as plant height,
• Seed size days to reach maturity, plant architecture and
• Tolerance to insects and diseases drydown which are affected by environmental
• Seedling vigour influence. For example, the Nepal grain type
• Reduced seed shattering (accessions of A. hypochondriacus L.) when
• Resistance to lodging grown in a semiarid condition in Kenya produced
• Synchronous drydown of the plant and short plants (<1 m), with multiple flowering
seedhead stems that matured in 60 days with a high harvest
• Easy threshability index. When the same accession was grown in
Pennsylvania, it produced tall plants (>2 m) with are reduction of leafiness in the grain head and
a single stem that matured in >160 days and reduced plant height.
showed a low harvest index.
Information about incompatibility barriers In the cultivation of grain amaranth, seed shat-
preventing interspecific hybridisation are to be tering is one of the prime reasons for significant
collected. Few barriers seem to exist between loss in commercial grain production (Fitterer et al.
crosses of A. cruentus L., A. hypochondriacus, L. 1996). Amaranth seeds are prone to shatter. Each
and A. hybridus L. However, crosses between A. amaranth seed is enclosed in a separate papery
caudatus L. and any of the above species often utricle. The grain amaranths have circumscissile
result in nonviable progeny (Pal and Khoshoo utricle with a seam or abscission zone along the
1972). equator that opens at maturity. At maturity when
utricles open, the papery utricle cap or lid gets
separated along the abscission zone and fall off
8.3 Objectives in Amaranth causing removal of seeds. Most of the seeds get
Breeding trapped within the compact inflorescence.
However when the grain heads are combined or
A systematic and comprehensive breeding work bulk harvested, the brief shaking causes additional
for improvement of amaranths is yet to be seed loss. Plant breeders could reduce shattering
adopted. For long-term genetic improvement, by developing cultivars that have indehiscent utri-
few experimental approaches are essential. cle cap lacking abscission zone and therefore do
Experimental approaches and breeding objec- not separate from the lower part of the utricle
tives are quite different in vegetable and grain (Brenner and Hauptli 1990; Joshi and Rana 1991).
amaranths. The major objectives in improving Taxonomists including Sauer (1967) have
cultivars of grain amaranths are to raise yield, observed that some populations of A. powel-
increase pest resistance and improve harvestabil- lii S. Watson have non-circumscissile utricles. A.
ity. The modern cultivars tend to lodge, shatter powellii is a wild and weedy species closely related
and often mature early or late. Kauffman (1992) to the grain amaranths. Non-circumscissile utri-
working at Rodale Research Centre (RRC), cles have also been observed infrequently in culti-
Pennsylvania, identified several useful breeding vated amaranths (Hauptli et al. 1980; Joshi 1981a,
traits like increased seed size, synchronised dry- b), and few such populations are also indicated in
down of plant and seedhead, resistance to seed the GRIN database (USDA-ARS 2001). Jain et al.
shattering, improved pest resistance/tolerance (1984) used the symbol Dh and dh in A. hypo-
and increased seed protein and functional traits. chondriacus, with dh representing a recessive non-
He also subsequently released several lines in the circumscissile allele and Dh representing the
1980s. Williams and Brenner (1995) emphasised dominant circumscissile allele. They noted that the
three internationally recognised breeding objec- trait has potential for use in breeding programmes.
tives for grain amaranths, viz. reduced plant The non-shattering trait from A. powellii (Pl
height and high yield, enhanced food quality and 572261) has been possible to transfer to both A.
non-shattering of seed. A concise list of goal or cruentus and A. hypochondriacus applying tradi-
objectives which should be considered in breed- tional breeding procedures (CAD 2009). Three
ing programme of amaranth is mentioned below: interspecific hybrid populations of grain amaranth
with non-shattering (seed retaining) utri-
1. Harvestability of grain: The prime aspects cles, namely, DB 92226, DB 9350 and DB 98246,
which are associated with the proper and ade- were developed at Agronomy Department, Iowa
quate harvest of grains include seed shatter- State University of the North Central Regional
ing, lodging, timing of maturity and uniformity Plant Introduction Station (NCRPIS). These popu-
of maturity and drydown of plant at seed lations showed little or no abscission at the equator
maturity. The aspects with lesser significance of the utricle or beneath. The first two are intended
8.3 Objectives in Amaranth Breeding 113
to be crossed with standard cultivars and appeared when grown in fertile well-drained soil (Weber
as a source of shattering resistance in newly devel- and Kauffman 1990). Dwarf plants are short
oped cultivars. The cultivar DB 98246 was devel- due to reduced internode length. One source of
oped with an intention of biomass production. The dwarf statute was identified in A. caudatus
populations were released by Iowa State (Fig. 8.1). In segregating progenies this was
University during 1999 and 2000. determined by a single recessive gene (Kulakow
Lodging causes the whole plant to lean or fall 1987). Reduced leafiness in the inflorescence
over. This may or may not be related to head size. head is an added attribute to improve harvesta-
The RRC line that has been shown as the highest bility. Most amaranth cultivar has leaves within
lodging resistant is D136 that may partially be inflorescence. Having no leaves in the inflores-
due to its later maturity (Myers 1996). cence and also limiting flowering to the stalk
Timing of seed maturity is an important trait. terminus may be the appropriate goal for grain
Inflorescences of the grain amaranths are highly amaranths (Kauffman 1992). The dense inflo-
branched. Flowering progressively proceeds rescence types can increase grain mould prob-
from bottom to the top of the inflorescence so as lems by retaining moisture, and the drooping
the maturity of seeds. As a result seeds reach types are difficult to harvest (Sumar et al.
maturity at different times on a single plant. 1992).
Adjacent plants in the field may also differ in
maturity. Flowering over an extended period of 2. Seedling vigour: Amaranthus is a rapid
time has an adverse effect on harvestability. grower, but after emergence seedling initially
However breeding for this trait should avoid grows very slowly during the first few weeks;
developing amaranth cultivar dependent on thereafter its growth gets momentum. This
timely rain during a narrow flowering span. initial slow growth rate leaves amaranth
Amaranth grain matures much earlier and the
plant dries up quite late. If the inflorescence
heads are allowed to remain till the plants dry,
heavy shattering can lead to grain loss. Vegetative
drydown in relation to seed maturity affects
harvest ability. Delay in drydown exacerbates
lodging and seed shattering. Even after leaf drop,
stems and heads may still contain too much mois-
ture for efficient threshing. Care must be taken to
avoid loss of stem strength for types that dry
down too quickly.
Amaranth breeding lines lack uniformity for
characters such as plant height and grain-head
size. The height of individual plants in a field
can be quite varied. Some plants that are nearly
of full height have much smaller seedhead and
thinner stem, while others usually have larger
seedheads. Cultivars that are more uniform in
height and grain size could improve the harvest
ability of the crop. Breeding work at RRC suc-
ceeded (Kauffman 1992) in developing shorter
stature line. Shorter stature is helpful to prevent
lodging. Dwarfism has been recovered from
several crosses though some of the RRC breed- Fig. 8.1 Dwarf form of Amaranthus caudatus with erect
ing lines can attain excessive height (≥2 m) terminal inflorescence
vulnerable to weed competitors. The fragile ble. Teutonico and Knorr (1985a) developed
nature of amaranth seedling makes them sus- an improved method to select for low oxalate
ceptible to damage from wind-borne soil. by screening cultured amaranth cell.
Greater seedling vigour, which could come 6. Stress tolerance: Amaranths can be cultivated
from larger seed size, might allow amaranth to on different types of soil even in marginal areas.
be planted deeper, which could be desirable It is tolerant to high temperature. Cold-tolerant
with certain soil and weather condition. seedlings could be a significant improvement,
3. Seed weight: Larger seeds would improve though there is no clear evidence for the exis-
seedling vigour, ease of handling and popping. tence of cold tolerance. Germplasm screened
Wild species are potential source of genes for for cold tolerance and tolerance to aluminium
larger seeds. Artificial polyploids can also be toxicity in acidic soil showed variability.
used to increase the seed size. Seed size can 7. Disease and pest: Amaranthus tricolor is the
be increased by 41–159 % with polyploidy worst sufferer among amaranths from fungal
(Pal and Khoshoo 1968; Sun and Yue 1993), disease, often prematurely killed by a fungus
with significant changes in the nutritional that aggressively colonises the leaves and
value (Misra et al. 1971) and improved pop- stems (Bansal 1996). Most of the accessions
ping (Pal and Khoshoo 1968). Polyploidy is appear to be very susceptible. Finding of an
an established mean to increase seed size. improved resistant cultivar to the pathogen is
4. Grain yield: Like other crops higher yields are desirable. Evaluation of this resistance among
also desirable in amaranths. Increase in grain diverse amaranth accessions is in progress.
yield is quite feasible in amaranths. Cultivars Amaranths are susceptible to a number of
occasionally have yielded much higher in rep- insects (El-Aydam and Burki 1997). The grain
licated research trial (≥3000 kg/ha) (Myers amaranth can recover after insect feeding, but
1996) in comparison with commonly seen in vegetable amaranths, the damage is severe
farmer yield (≤1000 kg/ha). Heterosis may be enough to lower the yield heavily. The insect
a key factor to boost grain yield in amaranths. sucking can damage the developing seeds and
5. Taste and nutritive value: Few people like the reduce the harvest up to 80 %. In worst situa-
taste of amaranth, while others find it unac- tion insect can disrupt the vascular supply in
ceptable. It is blended with other cereal prod- inflorescence, destroying the whole branches.
ucts at such a low percentage that its taste is Wilson et al. (1989) identified some potential
masked by other grains in the mixture. A sources of tarnished plant bug resistance eval-
milder or widely accepted taste profile would uating the germplasm for resistance at North
favour the acceptance of amaranths in food Central Regional Plant Introduction Station
marketplace. Research on amaranths in the (NCRPIS), Iowa. In Mexico, the stem borer
last few decades has emphasised mainly its Sciara is a problem, but a resistance was
good nutritional profile, more precisely its found in the Aztec grain race (Espitia 1994).
high seed protein level and high-lysine Resistance to root-knot nematodes is available
content. Amaranth germplasms show diver- in some accession of A. cruentus and A. dubius
sity in protein level and other nutritional val- (Babatola and Awoderu 1986).
ues. Few cultivars of A. hypochondriacus
were successfully selected for higher seed Based on the results obtained and problem of
protein content in India. Plant breeders can amaranth cultivation, the following course of
select for nutritionally superior vegetable or actions was proposed:
forage cultivars. The anti-nutrients like oxa-
late and nitrate contents vary between geno- 1. Detailed genetic studies involving diallel
types (Devadas et al. 1984). Selection of analysis in order to evolve high-yielding gen-
variant with low oxalate and nitrate content otypes with lower content of anti-nutrient fac-
will widen its acceptability as herbage vegeta- tor and high average nutritive value with late
8.3 Objectives in Amaranth Breeding 115
bolting habit, lodging resistant and non-

shattering traits.
2. Standardisation of the agrotechniques to
obtain maximum yield, low content of anti-
nutrient factor and late bolting habit. 3 major
3. Fundamental studies on metabolism of anti- crops
(Maize, Rice &
nutrient factors in amaranths. Wheat)
4. Standardisation of plant protection measures
using herbal or safer insecticides with mini-
30 feed the world
mum residual toxicity and waiting period.
5. Amaranth is an ideal crop for biocycling of
200 entering statistics
urban waste and for minimising soil, water
and environmental pollution. Development of
7000 used
high-lysine lines in grain amaranths is yet
another important area of research. 30,000 edible
Genetic variability is very high in the popula- 30,000 documented

tions of wild amaranths and fairly high in culti-
400,000 plant species estimated worldwide
vated species. It is possible to make use of the
unexploited germplasm resources of the wild
Fig. 8.2 The use of crop species diversity in agriculture
species for crop improvement and cultivar breed- (FAO 1996a, b, c)
ing of this underutilised but promising crop.
According to Kauffman (1984), the traits useful
for successful agronomic use of amaranths are ing a widely adaptable elite varieties (Fig. 8.3).
(1) synchronised drydown of the entire plant at The decline of genetic variability both at the
maturity; (2) reduced plant height and reduced inter- and intraspecific levels is associated with
branching; (3) non-shattering seeds that facilitate several risk factors like epidemics of pests and
easy harvesting; (4) development of cultivar with diseases due to greater genetic vulnerability, the
light-coloured seeds with high-protein content, lack of adaptability to climate-change-related
fast growth and high water-use efficiency; (5) stresses or other environmental stresses, the lack
production of cultivars resistant to fungal or of genetic variation for specific quality trait and
insect diseases; and (6) more branching in vege- reaching performance plateaus or plateau of stag-
table amaranth for multiple cuttings. nation. Therefore a more efficient utilisation of
In our present agricultural system, both inter- plant genetic diversity is a prerequisite to meet
and intraspecific diversities are declining. Only the challenges endangering food security and
30 edible plant species have the prime responsi- poverty alleviation (FAO 1996b). General aims
bility to feed the world though there is an esti- and objectives of using Plant Genetic Resources
mated total of 30,000 (FAO 1996a) to 50,000 (PGR) in crop improvement are:
(Sánchez-Monge 2002) edible plants known to
exist. Out of 30 edible species, only three major 1. Development of cultivars specifically adapted
crops being maize (Zea mays), wheat (Triticum to abiotic or biotic stresses
aestivum) and rice (Oryza sativa) performing the 2. Assurance of sustained production with
major role (FAO 1996a) (Fig. 8.2). The plant increased nutrient and water efficiency in con-
breeders involved in amaranth breeding contrib- venient environment through reduced applica-
uted towards utilisation of diversity at the intra- tion of agrochemicals
specific level developing novel breeding 3. Development of alternatives for farmers
population, selecting best genotypes, developing through development of industrial, energy or
genetically homogeneous cultivars and introduc- pharmaceutical crops
Fig. 8.3 The diversity

triangle of plant breeders
(FAO 1996a, b, c)
Homogeneous, indistinguishable
wide adaptations
Improved
Decreasing diversity
Increasing diversity
Cultivars
Variety development, release &
seed production
Adapted breeding programme
Evaluation before breeding
Materials in the collection
Discovery & Collection
Available diversity of the crop species

(in situ and ex-situ)
Breeding systems of Amaranthus species are India-coordinated research project on underuti-

complex because they are influenced by both lised and underexploited crop plants for its fur-
genetic and environmental variations (Hauptli ther improvement involving a large number of
and Jain 1985; Jain et al. 1982). Breeding work research stations in different parts of India. This
on grain amaranth has just began and needs fur- integrated work has made some progress, and
ther research for drought resistance, yield some high-yielding varieties have been identified
improvement and maturation of grain (Brenner and became available under commercial cultiva-
et al. 2000; Joshi and Rana 1991; Williams and tion in the rainfed areas.
Brenner 1995). The breeding of new varieties has Field trials of commercial amaranths have
just begun in Europe in the UK, the Netherlands, been carried out throughout the world, like in
Germany, Austria, the Czech Republic and China, Iowa, Arizona, India, Kenya, Peru,
Poland. New breeding lines with potential for Montana, Utah, Mexico and Thailand. Through
high grain and biomass yield should be investi- field trials cultivars with desirable traits have
gated for performance under different climatic been selected. High yield and uniform lines could
conditions. be selected only after a few years of selection
(Yue and Sun 1993). Optimum time of planting,
type and application rate of fertiliser and response
8.4 Conventional Breeding to fertiliser, planting density, irrigation and
by Selection control of pest and disease were investigated and
standardised in different field conditions. In all
The work on the genetic improvement of grain experimental works conducted, only a limited
amaranths, especially in India, has been achieved selection of diversity of species and cultivars was
through conventional selection methods from represented. The research would be more effi-
local collections of landraces available at differ- cient and meaningful if representative collections
ent experimental stations. The selection process of most diverse and productive cultivars are
has been proved to be valuable as it has resulted selected for evaluation.
availability of varieties that are cultivated at pres- A reduction in plant height and branching with
ent and has provided a foundation from which high harvest index is of great value in crop
significant advances in yield can be achieved. improvement. Breeding system in grain ama-
Recently ICAR (Indian Council of Agricultural ranths varies from high rates of selfing (over
Research) has included crop amaranths under all 90 %) to mixed mating with as much as 30 % out-
8.4 Conventional Breeding by Selection 117
crossing. Outcrossing rates vary significantly with pure line selection from Koilpatli A 38 gives high
genotype and environmental parameters. A bidi- yield of succulent leaves and stems and matures
rectional selection based on male/female flower in about 8 weeks. A dual-purpose vegetable or
ratio per plant suggested that breeding system is cereal selection from Ooty A 50 gives good
under genetic control and thus can be modified to greens 3 weeks after sowing with medium leaves,
suit breeding procedure for either inbreeding or and stems developed. A yield of 300 kg/ha of
outcrossing species (Joshi and Rana 1991). Thirty nutritious grain was obtained. A high-yielding
five genotypes of grain amaranths were examined variety ‘Annapurna’ has been developed for culti-
for their adaptability and yield performance at vation (Joshi 1988) from the germplasm col-
three different locations (lowland rainfed areas at lected from Pauri in Uttrakhand, India, through
300 m above the sea level, upland 1000 m above pure line selection, and it has given an average
the sea level and highland more than 1000 m grain yield of 22 kg/ha under multilocation test.
above the sea level) in Chiang Mai, Thailand, Vivekananda Laboratory, Almora, Uttrakhand,
from 1982 to 1987 by Senthong (1986). The has developed VL-21 a local selection. Likewise
different types of grain amaranths showed marked NBPGR, New Delhi, has developed a high-grain-
variability in yield stability and adaptation. yielding selection IC-5564 for cultivation in
‘African’ and ‘Mexican’ morphological group plain. It would be worthwhile to resort to the
exhibited higher yield and more stability than most suitable breeding procedure such as bipa-
other grain amaranth types. The ‘South American’ rental mating, recurrent selection or diallel selec-
group was least well adapted to lower latitude but tive mating (Jensen 1970).
performed well in highland atmosphere. These Research on amaranth breeding for crop
observations show probable effect of drought improvement at North Central Regional Plant
stress in low rainfed regions and by cold atmo- Introduction Station, Iowa State University,
sphere at highland areas. resulted in the release of five improved breeding
Two populations, A. cruentus UC 87 and A. lines including traits like non-shattering seed
hypochondriacus UC 99, were subjected to S1 cases and large stems with good resistance to
family analysis for agronomic characters lodging for biomass. The non-shattering trait
(Ayiecho 1986). For mass and recurrent S1 selec- resulted in the better seed retention and subse-
tion for yield, height ratio and harvest index were quent increase in yield and improved ornamental
applied, and selections were advanced to the lines for cut-flower use. Accessions have been
second-generation mass selection in 1983. In identified with traits like male sterility, non-
addition, S1 selection for plant height and for shattering seeds, heavy seeds and dark red foli-
days to flower was applied to UC 87 and UC 99, age. Accessions having red foliage pigmentation
respectively. UC 87 showed high direct and indi- can be useful for colouring food. Incompatibility
rect selection gains indicating the presence of in crossing or minimal crossing with weedy spe-
additive effect. UC 99 gave the similar result. In cies is another useful aspect to maintain pure
both selections, mass selection was more effi- commercial breeding line. The accessions having
cient than S selection. minimum crossability with weedy amaranth are
Grain amaranth landrace accessions UCC 192 to be documented as the next goal.
(A. cruentus) and UCH 213 (A. hypochondria- The knowledge about interrelationship
cus) were used by Vaidya and Jain (1987). In a between various agronomic and quality charac-
mass selection experiment, involving both the ters along with direct and indirect influence of
populations, only 5 % of the tallest and highest- component characters on yield can be very effec-
yielding plants in both populations were selected tive to improve the foliage yield of vegetable
for three cycles. Selection gain was highest in the amaranths. Shukla et al. (2010) conducted an
first cycle for both the traits. The results favoured investigation to explore the interrelationship
the previous observation about the existence of among various agronomic traits and quality traits
genetic variability within landrace population. A and their direct and indirect effect on foliage
yield in 39 distinct cultivars of vegetable ama- pletely developed interspecific barrier, usually
ranth (A. tricolor). Among the agronomic traits, via a partially sterile F1 hybrid, by means of
plant height and number of inflorescence were repeated backcrossing and selection of well-
positively correlated with foliage yield, while adapted types’. Though the success of introgres-
chlorophyll a, chlorophyll b, carotenoid, fibre sive hybridisation in overlapping habitats is
and ascorbic acid showed positive correlation dependent upon the availability of a suitable eco-
with foliage yield. Chlorophyll a and chlorophyll logical niche for the establishment of introgressed
b showed significant positive correlation with types, introgressive hybridisation is supposed to
carotenoid, fibre and ascorbic acid, while ascor- have played a vital role in the origin and diversi-
bic acid was positively correlated with fibre and fication of amaranths especially in grain ama-
carotenoid. Protein content was linked with plant ranths. Improvement of heritable qualitative traits
height, branches per plant and 500 seed weight. that are governed by one or a few major genes or
Foliage yield showed direct positive correlation gene complexes is one of the prime objectives of
with chlorophyll a, carotenoid and inflorescence introgression. Generally, the conventional back-
length but revealed negative correlation with crossing procedure is applied to introgress traits
branches per plant, leaf size, seed yield, chloro- like resistances or restorer genes from wild rela-
phyll b, moisture content and ascorbic acid. tives into recipient breeding materials (= the
Suitable traits have been marked out to enhance recurrent parent).
foliage yield in vegetable amaranth. Hybridisation studies are very significant to
establish evolutionary linkage and accessible
gene flow for traditional breeding programme.
8.5 Hybridisation and Heterosis Murray (1940) was one of the first persons to
in Amaranths classify interspecific hybridisation within the
genus Amaranthus. He divided monoecious spe-
Hybridisation has been proved to be the most cies according to the arrangement of the male
efficient breeding procedure to create new varia- flower in the inflorescence. Two categories of
tions both under domestication and natural con- plants were recognised – type I plants having
ditions. Inter-varietal and intraspecific crosses male and female flowers intermingled with each
are the most successful in the sense that such other and type II plants having male flowers
hybrids are viable and fertile. In contrast hybrids arranged at the apical part of inflorescence. He
from wide crosses either suffer from nonviability performed quite a few crosses between and
or sterility or from both, as such promptly elimi- among type I monoecious species (e.g. A. cauda-
nated by the forces of natural selection. Very tus, A. hybridus, A. retroflexus and A. powellii),
often occasional hybridisation in nature between type II monoecious species (A. spinosus) and
incompletely isolated species (geographical dioecious species. Crosses between monoecious
races) leads to the development of hybrid swarms, species produced hybrids with different ease.
i.e. highly variable populations consisting of Crosses involving type I and type II plant showed
segregating progenies of species hybrid at geo- most difficulty in hybrid production. Crosses
graphical boundaries that separates distribution between type I monoecious species and type I
regions of species. Such an interspecific hybridi- dioecious species readily produced hybrids indi-
sation (that apparently looks like crossing cating a phylogenetic proximity between these
between two habitats) producing partially fertile taxa. Crossing involving A. hybridus and A. cau-
hybrids is called introgressive hybridisation or datus was among the most consistent with the
just introgression (Anderson and Hubricht 1938). weak prezygotic isolation indicating close affin-
Introgression can be defined as ‘the infiltration of ity. Interestingly, crosses between A. hybridus
germplasm from one species to another by and A. caudatus with A. tuberculatus were simi-
repeated backcrossing’ or more precisely ‘the larly prolific and consistent predicting evolution-
transfer of genetic materials across an incom- ary relationship.
8.5 Hybridisation and Heterosis in Amaranths 119
Natural hybrids of grain amaranths were ration of 13 different crop-wild and wild-wild
detected by characters intermediate between A. spontaneous hybrids was studied by Greizerstein
caudatus or A. cruentus and species of A. hybri- and Poggio (1995), and the information was used
dus complex. In some cases morphology of to formulate the first set of genomic configura-
hybrid reflects three-way hybridisation among tion for those species (Brenner et al. 2000). Crop-
the species (Tucker and Sauer 1958). The hybrid wild hybrids have been produced to fulfil all
forms may be found in nature which indicates major breeding objectives:
that first-generation hybrids were fertile enough
to advance to more stable subsequent genera- 1. Increase in grain yield
tions. Khoshoo and Pal (1972) involved A. hypo- 2. Increased tolerance against pest
chondriacus (as male parent) in crosses with A. 3. Increase in protein content
hybridus and A. caudatus, all having 32 chromo- 4. Early maturity and improvement in growth
somes. Hybrids that evolved from these crosses 5. Improvement in harvestability
showed the formation of 16 bivalents. Hybrids 6. Reduction in grain shattering
derived from crosses between A. hypochondria-
cus and A. hybridus showed much higher pollen One of the most widely used grain amaranth vari-
fertility than hybrids produced through crosses eties in the USA is A. hypochondriacus var.
between A. hypochondriacus and A. cruentus. Plainsman. The variety was derived from a cross
Surprisingly crosses between A. hybridus and A. between A. hypochondriacus and a Pakistani A.
caudatus resulted in lethal seedlings. Hybrid fer- hybridus accession (Baltensperger et al. 1992).
tility in this study consolidated the concept that Biomass heterosis and combining ability with
domestication of grain amaranths has occurred domesticated species have been measured using
independently and evolutionary A. hybridus interspecific crosses between A. hypochondria-
should be placed much closer to A. hypochon- cus and A. hybridus (Lehman et al. 1991). Useful
driacus and A. caudatus. traits inherent in wild species have been trans-
Grain amaranths and their putative progenitor ferred to crop species through hybridisation, e.g.
showed two basic chromosome numbers, i.e. crop-wild hybrids have been developed to trans-
2n = 32 (A. hybridus, A. hypochondriacus, A. fer non-dehiscence property of A. powellii to A.
caudatus, A. quitensis) and 2n = 34 (A. cruentus cruentus and A. hypochondriacus breeding lines
and A. powellii). Pal and Khoshoo (1982) tried to in an effort to reduce grain shattering.
explore the phylogenetic relationship between Hybridisation with A. cannabinus, a wild dioe-
these two basic numbers applying dibasic cross cious species, might be beneficial to get hybrids
between A. hypochondriacus with 2n = 32 and an with greater seed size (Brenner et al. 2000).
African race of A. hybridus with 2n = 34 chromo- Introduction of herbicide resistance from wild
somes. At the metaphase I, most of the meiotic species to cultivated species is of great implica-
cells of the interspecific F1 hybrids showed 15 tion. Herbicide resistance evolved in A. hybridus
bivalents and one trivalent association of chro- has been transferred to elite breeding lines of A.
mosomes. F2 hybrid progeny showed 1:2:1 segre- hypochondriacus and A. cruentus. Research
gation pattern for 32, 33 and 34 somatic related to hybridisation involving wild-crop rela-
chromosomes, respectively. This type of meiotic tives of amaranths subsequently emphasised the
configuration in dibasic hybrids indicated that study of gene flow between two problematic
n = 17 may have evolved by aneuploidy probably weeds, viz. A. hybridus and A. tuberculatus
involving reciprocal translocation resulting in a (Trucco et al. 2006a). This study has attracted
decrease in chromosome number from n = 17 to breeder’s interest because A. hybridus is almost
n = 16. Many authors studied the meiotic behav- unequivocally regarded as the common progeni-
iour in various crop-wild hybrids and provided tor of the domesticated grain species. The fre-
clues to understand gene pool accessibility and quent occurrence of herbicide resistance in
phylogenetic relationship. The meiotic configu- amaranth weed suggested that it should be pos-
sible to transfer the disease-resistant trait from showed even higher degree of hybridisation
weed to crop via hybridisation or alternatively to defect (about 90 % pollen sterility). Hybridisation
select that same trait in cultivated crops. between closely related species like A. edulis and
Interspecific hybridisation among most of the A. caudatus yields a hybrid showing hybrid
Amaranthus species has been investigated (Sauer vigour, probably as a result of overwhelming
1950), and some of them have been even treated effect of minor genetic disharmony by superior-
as ‘promiscuous’ (Trucco et al. 2005b). Though ity of heterozygous genotypes of the hybrids.
hybridisation occurs frequently between Intraspecific hybridisation between different cul-
Amaranthus tuberculatus and A. hybridus, tivars of A. hypochondriacus and A. cruentus has
genetic introgression takes place only in one shown no heterosis in the hybrids, yet interspe-
direction, from A. hybridus to A. tuberculatus cific hybridisation among grain species showed
(Trucco et al. 2009). Interspecific hybridisation biomass heterosis. Interspecific hybridisation
was experimentally documented under field con- rather than intraspecific hybridisation can
ditions for these two species (Trucco et al. increase biomass productivity in the hybrids
2005a,b). Chromosome numbers are variable (Lehman et al. 1991). Interspecific hybridisation
among weedy Amaranthus species. Documents between grain and vegetable amaranths often
on interspecific hybridisations in Amaranthus produces hybrids with deformities, high pollen
reveal that equal chromosome number is not a sterility and chromosomal aberrations indicating
prerequisite for hybridisation to take place, but the presence of a great incompatibility barrier
hybrid progeny appears to be more viable and between species of sect. Amaranthus and sect.
stabilised when their parental species have the Blitopsis (Mohideen and Irulappan 1993).
same chromosome number, as in the case of Natural hybrids of A. spinosus (n = 17) and A.
hybrids derived from crosses between A. hybri- dubius (2n = 32) are easily formed in areas where
dus and A. tuberculatus (Trucco et al. 2009). the two species grow side by side. Genome anal-
Floral structure may also be an influencing factor ysis of the resultant triploid suggested that 17
in hybridisation, as pollen grains from other chromosomes homologous to the chromosome
species have to face greater pollen competition complement of A. spinosus are present in A.
in self-pollinating species than in dioecious dubius (Behera and Patnaik 1982). However
species. study on characteristics of the amphidiploid A.
Amaranths are usually wind pollinated with dubio-spinosus (2n = 49) does not support this
the exception of grain amaranths which are insect claim (Sreelathakumary and Peter 1993). From
pollinated. Species included under section the present knowledge of chromosome number
Blitopsis are predominantly self-pollinating due relating to relationship within the genus, it
to the number of staminate flowers per glomerule appears that aneuploid condition in all probability
and small axillary inflorescence. On the other has emerged as a result of interspecific hybridisa-
hand, single staminate flower per glomerule and tion (Grant 1959a, b, c). Aneuploidy remains a
big inflorescence are responsible for more cross- major factor responsible for species variation,
pollination in sect. Amaranthus (the grain ama- speciation and taxonomic ambiguity of the genus.
ranths). Interspecific hybrids are formed as a Three species of amaranths (A. hypochondriacus,
result of cross-pollination of two different spe- A. caudatus and A. edulis) were rendered tetra-
cies. Eight naturally occurring interspecific ploid (Pal and Khoshoo 1977). In comparison
hybrids have been reported in vegetable ama- with diploid, the raw autotetraploids are shorter,
ranths species with varying degree of sterility. In sterile and thus non-lodging.
the case of natural hybridisation involving A. Eight hybrids derived from interspecific
edulis and A. hybridus, complete chromosome crosses between A. edulis and A. hypochondria-
pairing was observed, but the pollen fertility was cus, A. edulis and A. caudatus, A. edulis and A.
less than 50 % (Sreelathakumary and Peter 1993). caudatus var. atropurpureus, A. caudatus and A.
Interspecific hybrids of A. tricolor and A. lividus hybridus, A. edulis and A. hybridus, A. caudatus
and A. hypochondriacus, A. hybridus and A. gene than susceptible individual. The resistant
hypochondriacus and A. powellii and A. hypo- species acquire a rare and unique trait of rapid
chondriacus were investigated. The crosses were amplification of EPSPS gene that enables them to
successful when crossing made between A. pow- resist the herbicide glyphosate. This unique trait
ellii and A. hypochondriacus, A. edulis and A. may be transferred from resistant species to other
hypochondriacus, A. caudatus and A. hypochon- related species exposed to similar selective pro-
driacus, A. edulis and A. caudatus, A. edulis and cess through interspecific hybridisation.
A. caudatus var. atropurpureus and A. hybridus According to Heap (2011), glyphosate resistance
and A. hypochondriacus (Pal and Khoshoo has been reported in 21 species globally.
1973a, b). Hybrids evolved from crossing Amplification of the 5′-enolpyruvylshikimate-3-
between A. hypochondriacus (as the male parent) phosphate synthase gene has recently been
and A. hybridus, and A. caudatus showed the for- reported in Amaranthus palmeri which has
mation of 16 bivalents. However, hybrids pro- enabled them to resist glyphosate. This evolved
duced from crosses between A. hypochondriacus property could be introgressed to other weedy
and A. hybridus showed much greater pollen fer- Amaranthus species through interspecific hybrid-
tility than hybrids derived from crosses between isation. Gaines et al. (2012) evaluated the feasi-
A. hypochondriacus and A. caudatus. bility of interspecific hybridisation as a measure
Interestingly, in hybrids between A. hybridus and to transfer this trait from A. palmeri to other
A. caudatus, seedling was lethal (Trucco and weedy amaranths like Amaranthus hybridus,
Tranel 2011). Ranade et al. (1997) found that the Amaranthus powellii, Amaranthus retroflexus,
hybrid of A. edulis and A. caudatus is associated Amaranthus spinosus and Amaranthus tubercu-
with A. caudatus in a single cluster, while the latus via pollen grains of A. palmeri. Crossing
hybrid of A. hybridus and A. hypochondriacus is was conducted under both the field and green-
included in the cluster of the latter species. The house conditions using glyphosate-resistant male
low genetic distance values between these A. palmeri as pollen donors and the other weed
hybrids and other accessions of A. caudatus and Amaranthus species as pollen recipients.
A. hypochondriacus, respectively, indicated the Hybridisation between A. palmeri and A. spino-
fact that these are not strongly and completely sus occurred with frequencies of <0.01–0.4 % in
differentiated genetically. Hybrids produced field crosses and 1.4 % in greenhouse crosses. A
through crosses with wild species have addressed majority of the hybrids of A. spinosus and A.
all major breeding objectives including yield palmeri were monoecious, grown to flowering
improvement, pest management and grain har- and produced viable seed. Hybridisation fre-
vestability (Brenner et al. 2000). The crop-wild quency in field condition between A. palmeri and
hybrids have been produced to transfer non- A. tuberculatus was <0.2 % and between A.
dehiscence trait from A. powellii to A. cruentus palmeri and A. hybridus <0.01 %. This is consid-
and A. hypochondriacus breeding lines, with an ered as the first documentation of hybridisation
intention to reduce grain shattering and improve between A. palmeri and both A. spinosus and A.
grain harvestability. hybridus.
Herbicide resistance in weed represents an A key evolutionary concept is the relative
excellent model system to study evolutionary importance of acquiring a unique adaptive trait
concept. The most common herbicide used to via interspecific gene transfer versus evolution of
control weed in the world is glyphosate. But the same adaptive trait within a species.
simultaneous evolution of resistance against this Interspecific hybridisation can be influential on
herbicide is a problem in agriculture. This resis- evolution if hybrid genotypes are more superior
tance is due to an overexpression of a novel gene to one or both parents (Abbott 1992; Barton
5′-enolpyruvylshikimate-3-phosphate synthase 2001). As far as the introgression of herbicide
(EPSPS gene). The resistant plant possesses 40- resistance genes are concerned, the hybrid would
to more than 100-fold more copies of EPSPS clearly have higher fitness than the susceptible
parental species when exposed to herbicide selec- dise with A. palmeri may create management
tion. Much effort has been devoted to study the problem in agricultural field in the future.
transfer of herbicide-resistant genes from crop Both Amaranthus tuberculatus and A. hybri-
species to related weed species (e.g. Ellstrand dus are phylogenetically linked to the A. hybri-
et al. 1999; Legere 2005; Gaines et al. 2008), but dus-A. powellii-A. retroflexus complex (Franssen
information regarding interspecific hybridisation et al. 2001b; Trucco et al. 2007) regarding acqui-
between weed species are still not adequate. sition of glyphosate-resistant trait from A. palm-
Intraspecific hybridisation has been proved to be eri through interspecific hybridisation, prior to
useful in the transfer of herbicide resistance at the the evolution of glyphosate resistance within
commercial field scale in Lolium (Busi et al. those species. Though glyphosate is used inten-
2008) and Amaranthus (Sosnoskie et al. 2009). sively on Amaranthus sp., only A. tuberculatus
Glyphosate resistance has evolved in A. and A. palmeri have evolved glyphosate resis-
palmeri due to overexpression of EPSPS tance (Culpepper et al. 2006; Legleiter and
(5′-enolpyruvylshikimate-3-phosphate synthase) Bradley 2008; Norsworthy et al. 2008; Steckel
gene. This trait is transmissible to other weedy et al. 2008). It indicates the preference of inter-
Amaranthus species, particularly A. spinosus via specific hybridisation as significant evolutionary
pollen-mediated gene flow. The hybridisation avenue for glyphosate-resistant trait to occur in
rate was found higher in field condition because related Amaranthus species more rapidly than by
such condition represents ideal natural situation independent evolution within each species
where A. palmeri coexists with other Amaranthus because interspecific hybridisation occurs more
species. Thus, interspecific hybridisation may be rapidly than independent evolution. This also
anticipated to occur in field conditions and repre- indicates that the major troublesome Amaranthus
sent a potential route of evolutionary adaptive weed species would not be manageable with
change for related species currently lacking this glyphosate (Trucco et al. 2009). Depending on
novel resistance mechanism, with considerable the similarity on chromosome number (2n = 34),
agronomic relevance for managing Amaranthus pollen morphology (Franssen et al. 2001a) and
weeds. Evolved glyphosate resistance mecha- genome size (Rayburn et al. 2005), it can be pre-
nism in A. palmeri is widespread and is of great dicted that A. palmeri and A. spinosus may have
potential for the rapid selection and spread of evolved from a more recent common ancestor
glyphosate resistance in other troublesome than the other Amaranthus species examined.
Amaranthus species via interspecific hybridisa- The rapid evolution of herbicide resistance in
tion with A. palmeri. In the case of hybrids with palmer amaranth due to multiple copies of the
A. spinosus, glyphosate-resistant (GR) hybrid EPSPS genes suggests that the genome of it may
progeny are self-fertile, and their progeny are be rearranged. Epigenetic responses to a chang-
viable and GR. Amaranthus spinosus a common ing environment based on alterations in genome
weed is a close relative of A. palmeri, a problem- architecture rather than changes in the underlying
atic agricultural weed with widespread glypho- DNA base sequence could explain at least some
sate resistance. These two species are known to of its capacity to adapt rapidly (Ward et al. 2013).
hybridise facilitating transfer of glyphosate resis- A large number of non-hybrid progeny appeared
tance. According to Nandula et al. (2014), from controlled crosses made between dioecious
glyphosate resistance in A. spinosus is caused by A. palmeri (palmer amaranth) and A. tubercula-
the amplification of EPSPS gene. Part of EPSPS tus (common water-hemp). All the non-hybrids
amplicon is found to be present in GR A. palmeri. were examined for two interspecific crosses, and
This is due to hybridisation between A. spinosus all plants were female and had DNA content very
and GR A. palmeri. Though A. spinosus is not of similar to the female (palmer amaranths). Among
such stature of A. palmeri and A. tuberculatus, the hybrids eight were nonviable (lethal or neu-
still its ability to germinate over a broad range of ter), and only one hybrid (triploid) continued
temperature, profuse seed production and hybri- gene movement (Trucco et al. 2006b). As a dioe-
cious species, palmer amaranth is an obligate ters have also been applied to establish the pater-
outcrossing species. Grant (1959b) observed that nity of amaranth hybrids (Wassom and Tranel
heteromorphic sex chromosomes are not present 2005). The domesticated amaranths function as
in the karyotype of any dioecious amaranth maternal parent characterised by white seed-coat
species. Apparent agamospermy has been colour trait which is recessive to brown seed-
reported in female palmer amaranth plant polli- coat colour. Hybrids derived from crosses with
nated by common water-hemp (Trucco et al. dioecious amaranth species are distinct from
2007). This requires further investigations. those hybrids that have evolved from crosses
Wetzel et al. (1999b) reported the transfer of ALS with monoecious species, since dioecism is con-
resistance via hybridisation and backcrossing sidered as a dominant trait (Trucco et al. 2006b).
between palmer amaranth and common water- Furthermore F1 hybrids obtained from crosses
hemp. Gaines et al. (2012) also reported low level between grain amaranth cultivar and monoe-
(˂0.2 %) of interspecific hybridisation between cious weedy amaranths are often fertile, but
palmer amaranth and common water-hemp but hybrids from crosses between monoecious spe-
the highest level of fruitful hybridisation between cies and dioecious male show reduced fertility
palmer amaranth and spiny amaranth producing but close resemblance to dioecious weedy parent
viable and fertile F1 progeny. (Murray 1940; Trucco et al. 2006b). Two grain
Some weedy amaranth species can hybridise amaranth cultivars, Amaranthus ‘D136-1’ (PI
with the cultivated grain amaranths (Brenner 538327), an interspecific hybrid developed at
et al. 2000) because the crossability barrier Rodale Institute (USDA 2013) with pale green
between grain amaranths and their weedy rela- inflorescence, and Amaranthus hypochondriacus
tives is very fragile. In the USA the common ‘Plainsman’ (PI 558499) developed at Rodale
weed which is cross-compatible with grain ama- Institute and University of Nebraska’s Panhandle
ranths is either dioecious (A. palmeri, A. arenic- Research and Extension Centre (Baltensperger
ola, A. tuberculatus) or monoecious (A. hybridus, et al. 1992; USDA, 2013) with maroon inflores-
A. powellii, A. retroflexus) (Wassom and Tranel cence, were evaluated for crossability with
2005). Weed-crop hybrids appear as weeds in weedy members (Brenner et al. 2013).
grain amaranth field and produce off-type brown Outcrossing rates with weed amaranths were
seeds in place of white seed lots of grain ama- assessed at nine locations in the USA. The
ranths (Hauptli and Jain 1984). These off-type ‘Plainsman’ has about ten times more weed
brown seeds though nutritionally similar to white hybrids than D136-1. The proportions of hybrid
seeds (Pond et al. 1991) create a cosmetic prob- progeny which were either monoecious or dioe-
lem that reduces the marketability (Sooby et al. cious were generally similar between D136-1
1998). Weed-crop hybrids are still a major seed and ‘Plainsman’ variety. Both the accessions had
purity problem. The frequency of outcrossing one male flower in each glomerule sampled in
between the cultivated grain amaranths ranges contrast to the relationship between varied male
from 0 % (on highly male fertile) to 100 % (on frequencies and outcrossing that was observed
male-sterile plant) and is heritable (Hauptli and with another amaranth population (Hauptli and
Jain 1985). Outcrossing rate in the available Jain 1985). The biological mechanism underly-
landraces of grain amaranths varies between ing the significant differences in outcrossing
7.6 % and 41.4 % (Espitia 1994) which is respon- across various locations could not be explained.
sible for frequency variability of cultivar-weed The low outcrossing can be helpful to select cul-
outcrossing. tivars that can be maintained more easily for the
The weed-crop hybrids are phenotypically production of high-value grain with few brown
distinguishable from amaranth cultivars as seed contaminants in field. There could be other
hybrids typically express the trait of dominant grain amaranth cultivars having even more
weed (Brenner et al. 2000). Molecular parame- extreme differences in outcrossing.
Interspecific and intraspecific factorial mat- mass of the parents. As forage, interspecific
ings among grain amaranths, from varied origins hybrids could substantially increase biomass
and types (vegetable, grain and weed), were yields over open-pollinated varieties. As the grain
tested for heterosis and combining ability. An amaranths possess a photosynthetic pathway
intraspecific mating within A. hybridus exhibited with an optimum temperature above 40 °C
highly significant (P = 0.01) accession heterosis (El-Sharkawy et al. 1968) and higher water-use
for biomass and GCA (general combining abil- efficiencies (Miller et al. 1984), their biomass
ity) and SCA (specific combining ability) effects. heterosis might be utilised in hot, arid environ-
No hybrid yielded more biomass than its best ments. Before the value of interspecific amaranth
parent, and mid-parent heterosis ranged from hybrids can be understood, their fodder quality,
−36 to 29 %. Date of maturity had little effect on hybrid seed production and cropping niches need
biomass heterosis in A. hypochondriacus mat- assessment. Male sterility sources (Peters and
ings, whereas late flowering of the accessions per Jain 1987; Gudu and Gupta 1988a) may be help-
se did cause greater biomass. The magnitude of ful in these studies.
biomass heterosis in intraspecific A. hypochon- Biomass heterosis has long been observed in
driacus crosses suggests that the Asian acces- Amaranthus. Interspecific crosses among the
sions may not be useful as parents in biomass or grain amaranths species and their weedy relatives
forage breeding programmes. However, among yielded sterile hybrids but with increased bio-
the factorial matings, only the intraspecific A. mass. The putatively palaeopolyploid nature of
hypochondriacus mating exhibited SCA effects amaranths coupled with two gametic numbers in
for biomass. This result is consistent with the the genus (n = 16 or 17) is probably responsible
idea that the ‘Hindustani’ region is a secondary for interspecific hybrid sterility (Pal et al. 1982).
centre of diversity (Grubben and van Sloten According to Lehman et al. (1991), interspecific
1981). An intraspecific mating between A. cruen- crosses among diverse amaranth germplasms
tus grain and vegetable types showed significant produced biomass yield of 0–57 % above the
(P−0.05) GCA effects for the grain types when mid-parent value. Biomass yield can be simply
used as male parents, but average heterosis was and rapidly estimated in hybrid amaranth
zero. Mid-parent heterosis ranged from 0 to 57 % (Lehman 1990). Basal stem diameter taken at
for intraspecific A. cruentus hybrids. The small harvest measured by hand callipers predicts the
variation for flowering and modest biomass het- dry weight when harvest sample size is at least 40
erosis in A. cruentus may be due to a narrowed plants and plant populations are high.
genetic base. Two theories that explain this nar- Grain protein percent was studied by Pandey
row genetic base are either (1) a post-Cortez and Pal (1985) in the F1 and F2 of a diallel cross
introduction of the crop to tropical Africa with of six A. hypochondriacus genotypes showing
subsequent selection or geographic isolation or additive and nonadditive gene effects. Data from
(2) an evolutionally recent, sexual isolation of A. a six-parent diallel without reciprocal studied by
cruentus from the other grain amaranths. An Pandey and Pal (1985) indicated that hybrids
interspecific factorial mating displayed high bio- from six crosses exceeded the mean parental
mass productivity, with an average of over 60 % value for protein content of the grain, and hybrids
more biomass accumulated when A. hypochon- from three of these crosses exceeded the better
driacus rather than A. cruentus was the male par- parent. Heterosis with respect to the six parents
ent. Date of flowering had a major influence on was 15 %.
biomass production. Grain yield heterosis has been estimated in A.
The use of interspecific amaranth hybrids hypochondriacus (Pandey 1984a). Using six-
seems to be a promising way to increase biomass parent, nonreciprocal, diallel crosses that created
productivity of Amaranthus. When vegetable 30 hybrid progenies, seven of these hybrids
types are used as parents, heterosis from interspe- outyielded their better parents by 33–71 %. This
cific hybrids may double to quadruple the bio- result indicated that increased heterosis is obtain-
8.6 Male Sterility in Amaranth Breeding 125
able by employing genetically diverse parents. duce nonviable pollen grains through
High general and specific combining ability in microsporogenesis. The entire process of micro-
parents suggested that inbreed line development sporogenesis is delicately balanced under the
might follow the maize pattern in which distinct genetic control of many loci, and mutation at any
lineages are maintained that produce consistent one locus may upset the entire process and result
hybrid vigour. Pandey (1984b) further analysed in the formation of non-functional pollen, i.e.
yield-contributing traits in the F1 and F2 of A. male sterility. As the sterility is a genetic abnor-
hypochondriacus and determined that both addi- mality in a natural population, hence sterile
tive and nonadditive genetic variances were plants are eliminated by selection forces.
important. However if properly maintained under domesti-
Reports on grain yield are lacking in A. cruen- cation, they can prove to be an asset to plant
tus, though Kulakow and Jain (1987) found breeding providing a natural and effective means
appreciable inbreeding depression in comparison for genetic emasculation of plants. Therefore
of F1 and F2 generation in terms of anthesis, plant male sterility simplifies the hybridisation on
height, petiole length, leaf length, leaf width, commercial scale. It is of more importance in
panicle length and weight. They imposed two crops where flower structure is an obstacle to
cycles of selection and found rapid gains for artificial emasculation or in monoecious plants.
anthesis time and leaf length, suggesting a large Male sterility can be utilised for developing mass
additive terms in the total genetic variance. reservoirs, particularly in outbreeders, for the
Overall heterosis for either grain yield or forage conservation of variability and also as a tester
yield appears to be promising in Amaranthus. genotype for assessing the combining ability of a
Male sterility is available in A. hypochondriacus large number of stocks, making hybrid products a
only but not in other species. A. hypochondriacus requirement for certain breeding programmes
(n = 16) is the most likely species to get benefit and genetic studies.
from initial heterosis with male sterility to The floral structure in amaranth with an initial
enhance breeding and hybrid seed production. A staminate flower followed by an indefinite num-
source of male sterility in Amaranthus cruentus ber of pistillate flowers in compact cluster creates
(n = 17) is needed. The latter may cater to enhance a definite problem in pollination studies.
interspecific forage crosses and permit reciprocal Development of a male-sterile line is very helpful
analysis of mating. to facilitate effective crossing. Most of the basic
Amaranth is still categorised as an underuti- breeding works on amaranths have been done at
lised crop. The main attributes responsible for its the Organic Gardening and Farming Research
limited domestication are the seed shattering Centre, PA, USA (Kauffman 1980). The
before harvest, tiny seeds, lack of public aware- phenomenon of male sterility in higher plant is
ness on its food value, lower demand and produc- due to either recessive or dominant ms genes or it
tion than conventional cereals, lack of production has cytoplasmic causes. In cross-fertilisation it is
research and development programmes, difficulty of considerable agronomic importance in con-
in maintaining pure stands due to cross-pollination nection with the utilisation of the heterosis effect.
by black-seeded wild species and lack of special- It is not a regular phenomenon in amaranths.
ised breeding projects (Rana et al. 2005). Male sterility is available only in Amaranthus
hypochondriacus (Peters and Jain 1987; Gudu
and Gupta 1988a and Brenner 1993) but not in
8.6 Male Sterility in Amaranth other Amaranthus species. Peters and Jain
Breeding descried the diagnostic features of male sterility
and identified some cytoplasmic male sterility
Male sterility is a reproductive deficiency of few type. Brenner (1993) found the unusual off-type
plants where male reproductive organs in her- black and translucent seeds in ‘Plainsman’ seed
maphrodite flowers are non-functional and pro- lots that frequently developed male sterility and
other aberrant types. These aberrant types may be 8.7 Wild Gene Pool, Landraces
the result of hybridisation with wild species and/ and Heritability
or seed immaturity (Brenner et al. 2000). The
source of male sterility is available through the There are few problems in amaranth breeding
USDA ARS (1999) – these include selection of like high degree of heterozygosity, low heritability
Peters and Jain (1987) and Brenner (1993) and of some trait and susceptibility to some diseases
other spontaneous male sterility found in land- specially in A. caudatus which are to be addressed
race germplasm population. and solved (Flores et al. 1982). Numerous major
Gudu and Gupta (1988a) identified twenty genes have been identified that may be useful for
male-sterile plants in a normal population of mutation breeding in amaranth, such as genes
Jumla variety (ex Nepal). They could be distin- coding flower, embryo and seed pigmentation,
guished from normal plants by the colour and leaf characters, type of starch in perisperm, early/
morphology of their inflorescence. The sterility late flowering, inflorescence architecture and
was conditioned by a single recessive nuclear vegetative architecture (Table 8.1). Goals in
gene, ms. Male sterility was discovered by Peters improving cultivars of grain amaranth are similar
and Jain (1987) in selfed families of several to those in other grain crops – improving and sta-
open-pollinated as well as interpopulation hybrid bilising the yield, increasing pest resistance and
individuals in grain amaranth (A. hypochondria- improving harvestability (Brenner et al. 2000).
cus). Segregation patterns in the F2 generation of Several desired traits were identified, viz. vigor-
numerous crosses involving male-sterile plants ous seedling growth, determination of the plant
clearly suggested gene-cytoplasmic mode of growth, timing and uniformity of flowering and
inheritance. Segregation ratios provided evidence seed maturation within plants, synchronous dry-
for one or two restorer nuclear genes in different ing of plant and inflorescence, reduction of leafi-
populations. Cytological studies showed male ness in the inflorescence area, reduction in seed
sterility to be associated with abnormal tapetal retention, increasing size of seeds, pale seed pig-
cell functioning and microsporogenesis failure mentation and enhanced food quality traits
prior to the first metaphase, leading to abortive (increasing seed proteins).
anthers. Further work on genetic selection and Grain amaranth cultivars are generally uni-
mode of gene expression is needed to explain the form, but due to high phenotypic plasticity, it
relative deficiency of male sterility in certain appears to be heterogamous in field plantings
segregating families. (Guillen et al. 1999). The high phenotypic vari-
A comparative study of microsporogenesis in ability displayed by these cultivars indicates the
male-sterile and male-fertile grain amaranth was ability of amaranths to adjust to environmental
conducted using electron-microscopy (Fang et al. variations, but it can also make selecting within
1996). The onset of microsporogenesis gets cultivars unreproductive (Guillen et al. 1999). In
deviated in male sterility at the mononuclear contrast to single-plant selections, some grain
pollen stage following the release of microspores amaranth landraces have been shown to be genet-
from tetrads. Abnormality in the behaviour of ically variable (Hauptli and Jain 1984) and have
degenerated tapetum was observed, which failed responded to selection.
to envelop individual microspores after their A landrace is a local variety, regional ecotype
release, leading to an abnormal vacuolation in of a domesticated plant species which has evolved
the mononuclear pollen grains. As a result, the over time largely to adapt in the natural and cul-
normal thickening of pollen wall could not occur, tural environment in which it exists. It differs
and pollen grains could not engorge, causing from conventional cultivars which have been
male sterility. selectively bred to conform to a particular stan-
8.7 Wild Gene Pool, Landraces and Heritability 127
Table 8.1 Several traits with associated hypothecated gene in amaranths

Trait Hypothetical gene Source References
Embryo colour Pe/pe homozygous A. caudatus Kulakow (1987)
Pink/pale prevent expression of
Pe
Floral feature
1. Male fertile/male sterile Ms1/ms1 A. hypochondriacus Gudu and Gupta
(1988a, b)
2. Cytoplasmic inheritance of male S1/S2 cytoplasmic A. hypochondriacus Peters and Jain
sterility genes; Rf1/rf1, Rf2/rf2 (1987)
and Rf3/rf3 nuclear
genes
3. Early flowering (temperate) Ea/ea Amaranthus sp. x A. Murray (1960)
hybridus
2. Late flowering (tropical) Ea/ea Amaranthus sp. x A. Murray (1960)
hybridus
3. Early flowering/late flowering Ea/ea A. retroflexus x A. cruentus Kulakow and Jain
(1985)
Inflorescence
1. Determinate/indeterminate Dt/dt A. caudatus Jain et al. (1984),
Kulakow (1987)
2. Erect/drooping Pd1/pd1, Pd2/pd2 and A. caudatus Kulakow (1987)
modifying factors
Hybridisation barrier abnormal Ah1/ah1, Ah2/ah2 A. caudatus x A. Jain et al. (1984)
hybrid/normal hybrid hypochondriacus
Herbicide resistance Triazine With chloroplast A. caudatus Cheung et al.
tolerant/non-tolerant (1988) Jordan
(1996)
Leaf and other markings A. hypochondriacus Jain et al. (1984)
1. Blade ‘V’ mark/no mark Vm1/vm1, Vm2/vm2 A. retroflexus Kulakow et al.
(1985)
2. Blade spot/no spot Ls/ls A. hypochondriacus Kulakow et al.
(1985); Gupta and
Gudu (1990)
3. Blade/red, vein/green R1/r A. tricolor Matsumura (1938)
4. Blade/red, vein/green R2/r A. tricolor Matsumura (1938)
5. Blade/pale red/green R3/r A. tricolor Matsumura (1938)
6. Blade of both surfaces red/green R4/r A. tricolor Matsumura (1938)
7. Blade red/green A/a, B/b, C/c A. tricolor Deutsch (1977)
8. Stem red/green A/a A. tricolor Deutsch (1977)
Perispermic starch glutinous/ Gl/gl A. caudatus Okuno and
non-glutinous Sakaguchi (1982)
Pigmentation
Red/green R/r Several species Jain et al. (1984)
Orange/green O/o A. tricolor Jain et al. (1984)
A. caudatus Jain et al. (1984)
A. cruentus Kulakow et al.
(1985)
Chlorophyll present/absent Three genes A. caudatus x A. Jain et al. (1984)
hypochondriacus
(continued)
Table 8.1 (continued)

Trait Hypothetical gene Source References
Normal/chlorophyll variegated Three genes A. caudatus x A. Jain et al. (1984)
hypochondriacus
Yellow (flavonoid)/nonyellow C/c A. tricolor Matsumura (1938)
Red seedling/flowering Lp/lp A. caudatus A. Jain et al. (1984)
hypochondriacus
Red whole plant/base of the plant Bd/bd A. hypochondriacus A. Jain et al. (1984)
tricolor and various other
species
Red normal/diminished One gene A. caudatus Jain et al. (1984)
Red not intense/intense One gene Jain et al. (1984)
Seed-coat colour P/p A. caudatus Kulakow et al.
(1985)
Black/pale A. hypochondriacus Coons 1982
Y/y A. caudatus Kulakow et al.
(1985)
A. hypochondriacus Gupta and Guru
(1990)
Brown/pale Br/br A. caudatus Kulakow et al.
(1985)
Black/dark medium light brown/pale Two genes A. cruentus x A. retroflexus Jain et al. (1984),
Kulakow et al.
(1985)
Shattering
Dehiscent utricle/indehiscent utricle Dh/dh A. hypochondriacus Jain et al. (1984)
Nonpersistent utricle/persistent Few genes Various species Jain et al. (1984)
utricle
Vegetative structure
Tall/dwarf Dw/dw A. hypochondriacus Pandy (1982)
A. caudatus Jain et al. (1984)
Cotyledon long/short Multiple genes A. caudatus Walton (1968)
Ok/ok A. cruentus x A. retroflexus Jain et al. (1984)
Narrow blade/normal blade V/v A. tricolor Kihara and
Matsumura (1935)
dard of characteristics. Specimens of a landrace lation but not all become susceptible to a new
tend to be relatively genetically uniform, but are pests or diseases. Landraces are quite distinct
more diverse than members of a standardised or from both ancestral wild species of modern stock
formal breed. Landrace populations often show and separate species or subspecies originated
variability in morphological appearance, but they from the same ancestor as modern domestic
have a certain genetic similarity and can be stock. Landraces are not all derived from ancient
identified by their appearance. Landraces have a stock, largely unaltered by human breeding inter-
genetic continuity with improved varieties. The ests. Amaranthus has a rich stock of landraces
relatively high level of genetic variation of land- which can be a rich source of wild gene pool that
races is one of the advantages that these can have can be utilised in amaranth breeding. The collec-
over improved varieties. Stability of landraces tions of landrace germplasms are the backbone of
against adverse conditions is typically high, but the varietal improvement programme which aims
yields may not be as high as improved varieties. at raising agronomically acceptable lines using
As a result, some individuals of a landrace popu- conventional plant breeding and selection meth-
ods. To generate varieties resistant to abiotic The presence of high variability in this crop
stress, certain types of germplasm-landraces, offers much scope for its improvement. Cultivated
wild relatives or wild progenitors may play a landraces of the three species of grain amaranth
basic role in the success of a breeding pro- have a wide degree of genetic diversity which can
gramme. Landraces are often capable to produce be utilised to develop selections of improved
some yield even in difficult situation where the lines with suitable characteristics for grain pro-
modern varieties fail or proved less reliable. duction. Traditional landraces of grain amaranth
Landraces of self-pollinated species like that are still extant in various agroecosystems
Amaranthus spp. are mixture of a great number may provide the genetic variability needed to
of homozygote genotypes (Ceccarelli et al. diversify the gene pool of improved amaranth
2004). Therefore landraces are the rich sources of varieties. These landraces can therefore be
readily usable genetic variation. Selection within exploited to significantly enhance grain amaranth
landraces is one of the easiest, oldest and cheapest productivity in various agroecosystems. Systematic
methods of plant breeding. Landraces as the survey, collection and evaluation of local
source of trait like drought tolerance is well doc- amaranth germplasm are of great importance
umented in the case of barley in the Syrian Arab for current and future agronomic and genetic
Republic. Landraces are capable of yielding yield improvement of this crop.
more than modern cultivars under low input and Yield and its component traits are controlled
stress condition. Genetic variation in landraces by polygenes, whose expression is greatly influ-
was studies by Jain (1985) applying qualitative enced by the environment. Sufficient genetic
markers and quantitative traits and allozyme vari- variability available in the germplasm material is
ation. The New World collections were found to a necessary prerequisite for yield improvement
be varied in the amount of genetic variation from through proper selection method (Ali et al. 2008).
region to region and between species. Most land- The genetic variance of any quantitative trait is
races seem to be highly homozygous and carry a made up of additive variance (heritable) and non-
significant amount of variation for quantitative additive variance which comprise dominance
traits such as plant height, branching, flowering and epitasis (nonallelic interaction). Therefore, it
time, head length and harvesting index. Accessions becomes compulsory to divide the observed
showed no or little allozymic variation within phenotypic variability into its heritable and non-
and among population, though they showed high heritable compartments with suitable parameters
morphological variability. The heterozygosity such as phenotypic and genotypic coefficient of
and interspecific gene transfer in northern Indian variation, heritability and genetic advance. To
population were found to be higher in compari- initiate and sustain an effective long-term plant
son with New World landraces. Landrace popula- breeding programme, genetic variability which is
tion from two states of India was studied in a heritable difference among genotypes is needed
greenhouse by Vaidya (1984) for the evaluation in an appreciable level within a population.
of genetic variation of few qualitative and quanti- Heritability denotes the proportion of phenotypic
tative characters. variance that is due to genotype which is herita-
Genetic variability in the germplasm is the ble. Heritability is known to differ to some extent
prerequisite for the success in any crop improve- depending on the populations handled. The
ment programme. Several researchers have estimates of heritability serve as a useful guide to
studied the potentials of improvement of grain the breeder. Selection for this trait would be fairly
amaranth through the study of its genetic easy as there would be close correspondence
resources (Pandey and Singh 2010; Prashantha between genotype and phenotype due to a very
and Nagaraja 2011). A wide range of variability little contribution of environment to the phenotype.
has been observed in respect of agroeconomic But for a character of low heritability, selection
traits in grain amaranths (Pandey and Singh may be considerably difficult or virtually imprac-
2010; Prashantha and Nagaraja 2011). tical due to the masking effect of environment on
the genotypic effect. The degree of gain in a trait 526832, IC 585672, IC 588820 and IC 588811,
obtained under a given selection pressure is which were found to be superior over the checks
expressed as genetic advance which is another (Co-1, Arikarai and RNA-1), are useful for
important criterion that guides the breeder to making selection to fix the desirable characters to
choose a selection programme (Hamdi et al. be exploited for developing suitable parental
2003; Shukla et al. 2004). High heritability and materials. The future of amaranth is dependent
high genetic advance for a given trait are indica- on the careful recombinant selection of genotypes
tive of the fact that it is governed by additive gene from the germplasm collections.
action and therefore offers the most effective The extent of genetic variability among
condition for selection (Tazeen et al. 2009). different Amaranthus species was investigated by
Germplasm characterisation and evaluation are Khurana et al. (2013). The investigation also
being conducted at a regular basis at a number of aimed at the estimation of the heritability values,
locations around the world. Extensive, well-doc- correlation coefficients and their partitioning into
umented amaranth germplasm characterisation direct and indirect effects by path analysis.
has been conducted by several organisations in Differences among genotypes in respect of all the
India, Peru and Mexico (Joshi 1981a, b; Sumar characters under study were highly significant.
et al. 1983; Espitia 1986). Phenotypic and Estimates of heritability and coefficients of
genotypic variances, phenotypic and genotypic genotypic and phenotypic variability in both, i.e.
coefficient of variation, heritability and genetic summer and rainy season, crops were found to be
advance have been used to assess the magnitude high for traits, namely, total green yield, number
of variance in grain amaranth germplasm (Rana of leaves per plant and leaf area index. Genetic
et al. 2005; Shukla et al. 2010; Pandey and Singh advance expressed in terms of percent of mean
2011; Prashantha and Nagaraja 2011). was found to be higher in characters like leaf area
Genetic parameters for grain yield and its index and leaf width in both summer and rainy
associated traits were studied (Sravanti et al. season. Traits like plant height, number of
2012) by evaluating 40 germplasm lines along branches per plant, leaf length, leaf width, number
with three checks (Co-1, Arikarai and RNA-1) of of leaves per plant, leaf area index and protein
amaranth (Amaranthus spp.) in a randomised content were significantly and positively corre-
block design with three replications at the lated with total green yield in both summer and
National Bureau of Plant Genetic Resources, rainy seasons.
Regional Station Hyderabad, Andhra Pradesh, Genotypic variability and character association
India, during kharif, 2010. Analysis of variance in grain amaranth genotypes for agronomically
revealed highly significant differences for geno- useful and yield-contributing traits were evaluated
types, indicating magnificent variation among the (Yadav et al. 2014). The experiment was con-
genotypes for all the 19 characters taken into ducted at the National Bureau of Plant Genetic
consideration, viz. plant height, stem girth, stem Resources, Regional Station, Shimla, Himachal
weight, number of branches per plant, leaf area, Pradesh, during 2013–2014. The materials for
leaf length, leaf width, petiole length, total leaf study consisted of 27 grain amaranth germplasm
weight, number of leaves per plant, days to 50 % accessions. The characters like width of leaf
flowering, inflorescence length, lateral spikelet blade, length of lateral spikelet and grain yield
length, days to 80 % maturity of seed, leaf/stem per plant showed high genotypic coefficient of
ratio, 1000 seed weight, protein content, dry variation (GCV) and phenotypic coefficient of
matter content and seed yield per plant. High variation (PCV) values. The differences between
heritability and genetic advance were observed PCV and GCV were very less for all the charac-
for all the characters, which suggested that they ters taken into consideration. This showed the
are controlled by few genes. On the basis of mean close proximity between the corresponding
performance for grain yield and its components, estimates of PCV and GCV in almost all the
five germplasm lines, namely, IC 588812, IC characters except petiole length, which suggested
that environmental parameters have very little estimates of broad-sense heritability indicate that
influence on the expression of these characters, improvement through selection would be limited.
and variability was caused by genetic constitu- Genetic advance expressed as percentage of
tion only. High degree of heritability associated mean was high for seed yield, leaf length, inflo-
with high genetic advances was recorded for the rescence length, number of leaves and was mod-
characters, like plant height, inflorescence length erate to low for other characters. Relationship of
and days to 80 % maturity, which suggested that heritability and genetic advance also gives an
these characters can be recognised as favourable idea about the type of gene action. It appeared
features and as an indication of additive gene that both the additive and nonadditive gene
action. Similar types of results were also reported actions influence all the characters, which sug-
by Mulugeta et al. (2012). On the basis of evi- gests that simple selection methods alone would
dence, most of the traits appeared to be correlated not be effective; rather hybridisation followed by
and associated with grain yield and intercorre- selection would be a better option for amaranth
lated among themselves. Significant positive improvement. Other workers have also reported
correlation was found in few cases at phenotypic the role of additive and nonadditive gene action
level like positive correlation between seed yield and emphasised recombinant breeding for grain
per plant and days to 80 % maturity and plant amaranth improvement. Direct positive effects of
height with days to 50 % flowering. Though the inflorescence length, number of leaves and leaf
rest of the features showed nonsignificant corre- length and negative direct effects of oil content,
lation, plant height, inflorescence length, lateral plant height and leaf width were observed on
spikelet length and days to 50 % flowering were seed yield. A character showing positive correla-
recorded with high GCV, PCV, heritability and tion may not have direct effect on seed yield but
genetic advance percentage of mean, which sug- may contribute to yield via other characters.
gested that maximum emphasis should be given Besides seed yield, inflorescence length, number
on the above features during the selection of of leaves and plant height were found to be
grain amaranth with higher grain yield. important characters for selecting better yielding
Hybridisation and subsequent phenotype-based genotypes.
selection could be helpful in the transfer of these Several genes have been identified governing
characters to the progeny. On the other hand, different traits in amaranths like pigmentation,
concomitant selection based on high inflores- determinant terminal inflorescence, drooping
cence length, plant height and high leaf blade inflorescence, embryo colour, early flowering,
width would be an effective selection method for male sterility, etc. Amaranths show variability in
the improvement of grain amaranth. pigmentation. The red, yellow and orange pig-
One hundred accessions (50 from India and 50 ments in amaranths are due to the presence of
from exotic sources) of grain amaranth betacyanin pigments (Cai et al. 1998). Among the
(Amaranthus hypochondriacus) germplasm were grain amaranths, six major genes have been iden-
grown in complete randomised block design. tified that control the pigmentation pattern
Data were recorded on six quantitative and two (Kulakow 1986). Diversity in variegated pattern
qualitative characters, viz. protein content (deter- may be due to quantitative inheritance.
mined by the conventional Kjeldahl method), oil Pigmentation markers are useful for varietal iden-
content (determined by nondestructive method tification, estimation of outcrossing, controlled
using New Port NML), plant height, number of hybridisation and aesthetic value. Wild and vege-
leaves, leaf length, leaf width, inflorescence table amaranths generally have black or dark
length and seed yield/plant (Rana et al. 2005). brown seeds, while grain amaranths typically
Heritability was high for protein content, oil con- have yellow- and white-coloured seeds.
tent and seed yield; moderate for inflorescence Polymorphism in seed-coat colour in controlled
length and leaf length; and low for plant height, progenies has suggested for one or two genes
leaf width and number of leaves. Moderate to low (Kulakow et al. 1985) with indications of multiple
alleles and additional loci for brown seed coat in coloured embryo is determined by two comple-
some crosses (Kulakow and Jain 1985, 1990b). mentary epistatic genes with one locus
The genetic control over four developmental determining the presence or absence of red beta-
characters in Amaranthus caudatus, viz. determi- cyanin pigment and the other regulating its
nance, panicle orientation, dwarfism and embryo expression. All these genes are responsible for
colour, and their inheritance patterns were stud- some of the largest morphological differences in
ied (Kulakow 1986, 1987). The grain amaranth A. caudatus. An accession of weedy A. retro-
species and various landraces show variability in flexus has rapid flowering under both short-day
the structure of terminal inflorescence. The deter- length (8 h) and long-day length (16 h). Hybrids
minant inflorescence trait that has been found in and backcrosses between A. cruentus and A. ret-
both A. caudatus and A. hypochondriacus is reg- roflexus showed that the early flowering is regu-
ulated by a single gene having the allele for the lated by a single gene with the dominant allele
common indeterminate inflorescence completely determining the earliness (Kulakow and Jain
dominant over determinant allele. Drooping ori- 1985). Isolation of a single gene determining
entation of inflorescence typically found in A. very early flowering may be useful for develop-
caudatus landraces is determined by two major ing short-season varieties facilitating rapid
genes with erect panicle incompletely dominant cycling of crosses (Kulakow and Jain 1985).
over drooping panicle, while other minor genes
are modifying panicle orientation. A single reces-
sive gene determines dwarfism. Pleiotropy was 8.8 Mutation Breeding
responsible for abnormal growth of dwarf plant.
Some genotypes of A. caudatus have pink- Conventional plant breeding involves three major
coloured embryos where pigmentation appears steps: the screening of relatively large popula-
during embryo development. This trait is deter- tions, utilisation of genetic variability and selec-
mined by one gene with the intensity of the pig- tion of the desired genotypes. There are simple
mentation and is modified by other genes and efficient techniques to induce genetic varia-
(Kulakow 1987). This is the first trait known in tion, and the use of radiation is one of such tech-
amaranths where seedling genotypes can be iden- niques. Application of radiation in appropriate
tified on the maternal plant. It is useful in detect- dose followed by selection for desired traits has
ing hybridisation in some population. Grain become a common practice in plant breeding.
amaranth A. hypochondriacus reported to have Mutation breeding in amaranths mostly empha-
both the glutinous and non-glutinous types of sised the use of radiation mutagenesis to enhance
periplasmic starch (Okuno and Sakaguchi 1982). quality and quantity of amaranth grain and to
In grain amaranth, starch granules are stored in evaluate the performance of selected mutants and
the perisperm of the diploid cells derived from parent lines. For this purpose the seeds of two
the nucleus. Therefore the starch property in the genotypes of grain amaranth Amaranthus cruen-
perisperm of grain amaranths is maternally inher- tus, genotype ‘Ficha’ and hybrid K-433 (products
ited. Two types of starch property at least in A. of interspecific hybridisation between A. hypo-
hypochondriacus differentiated in Mexico and A. chondriacus and A. hybridus) were irradiated by
hypochondriacus with both glutinous and non- γ radiation dose 175 Gy (Gajdošová et al. 2008)
glutinous perispermic starches were propagated followed by positive selection performed during
from Mexico to Nepal and India. Non-glutinous eight mutant generations (1998–2008). The mor-
plant believed to be a natural hybrid between glu- phological observations were recorded and selec-
tinous and non-glutinous plant was selfed by tion on desired traits was done starting from M2
Okuno and Sakaguchi (1982). Segregation ratio generation. The plant with negative features like
among 500 progenies and in the F3 derived from weak seedling growth, undeterminated plant
them indicated that a single major dominant gene growth, abundant leafiness in the inflorescence
determines the non-glutinous starch. Pink- area, nonuniform flowering and seed maturation,
8.8 Mutation Breeding 133
low seed size, etc., was removed from the field, eral generations under both field and greenhouse
and only plants with positive traits were col- conditions, and seven promising drought-tolerant
lected. The weight of 1000 seeds (WTS) was lines were selected. Further physiological and
determined using seed counter Contador (2 × 500 morphological interpretation on two of these
seeds), recorded and statistically evaluated. mutant Amaranthus lines were reported by Kgang
Finally, several putative mutant lines of A. cruen- et al. (2008)). These mutant lines showed
tus and hybrid K-433 were selected and charac- increased recovery after withdrawal of water for
terised by highly and significantly increased 2 weeks and also showed increased protein con-
WTS (in comparison with control) showing grad- tent per gramme of dry weight in comparison
ual tendency towards stabilisation of this trait with the wild type. Improvement of the grains
when compared with the samples of the previous and development of new varieties of A. tricolor
generations. All those selected plants are plants were previously achieved at the ARC
expected to have genetically fixed Vegetable and Ornamental Plant Institute, South
WTS. Therefore, these plant materials can be Africa, where seeds were treated with different
considered as valuable mutation lines useful in doses of gamma radiation to induce increased
future amaranth breeding programme. drought tolerance. Radiation is known to cause
The genetic variability is a prerequisite for changes to the genome, but plant performance of
developing new cultivars, and this variability is both the wild-type and mutant Amaranthus plants
either induced or is a natural phenomenon that may be similar (Jie et al. 1993). Hence, this study
occurs spontaneously. The rate of spontaneous asks for the comparison of the phenotypic perfor-
mutation is quite low and can’t be utilised for mance of irradiated and wild-type Amaranthus
breeding programme; for this reason mutations lines during drought conditions. Two mutant
are artificially induced treating with physical and lines were compared to a wild type, and the
chemical mutagen. Several useful genetic parameters considered were plant height, relative
changes have been induced artificially by muta- water content, protein content of the leaves and
gen treatment like high yield, flower colour, dis- genomic differences between the two mutants
ease resistance and early maturation and so on in and the wild type applying RAPD analysis.
crops, vegetables, medicinal herbs, fruits and In order to increase food availability and
ornamental plants. So far, over 3000 mutant vari- household incomes of families in the Andean
eties of rice, wheat, barley, sorghum, legumes, region of Peru, induced mutation techniques
cotton, edible oil, ornamental plants and fruits were applied to improve barley (Hordeum vul-
(www-mvd.iaea.org) have been officially gare) and kiwicha (Amaranthus caudatus) culti-
released over 60 countries. India and China are vars (Gomez-Pando et al. 2009). Kiwicha is a
the two major producers of mutant varieties in native and an ancient crop of the Andean region
the world to feed their ever-growing human but has been rediscovered as a promising crop
population. with high-quality seed protein and a
Amaranthus tricolor is a nutritious vegetable drought-tolerant habit. To improve the traditional
crop that is used as a subsistence or cash crop in cultivar Ancash of Amaranthus caudatus, physi-
the rural areas in Africa, Asia and especially in cal mutagen gamma rays were utilised. Dried
India. Its yield and production are severely lim- seeds of the traditional cultivar ‘Selection
ited by abiotic stresses such as drought. Mutation Ancash’, previously purified in isolated condi-
technology was previously used as a tool to cre- tions, were treated with gamma rays at doses of
ate genetic variation and to select for lines with 100, 200, 300, 400, 600, 800 and 1,000 Gy. In M2
improved drought tolerance (Kgang et al. 2008). generation, two types of mutations were identi-
A. tricolor seeds were subjected to different fied from the materials treated with gamma rays
doses of gamma radiation, and to ensure subse- at a dose of 400 Gy. In M3 and M4 generation,
quent seed germination, 160 Gy was selected. mutants with different yield potentials were
The evolved mutant lines were screened over sev- selected among the 36 mutant lines. Five mutant
lines appeared to have greater yield potential than Plant tissue culture techniques have made it
the parental cultivar. From these high-yielding possible to regenerate plants from all major food
lines, Centenario cultivar was selected and and horticultural crops in aseptic culture.
released in March 2006 with similar quality, bet- Micropropagation via organogenesis is now rou-
ter yield and different plant colours than the tinely used for clonal propagation of ornamental
parental material. The yield varied between 3500 and other vegetatively propagated plants.
and 5500 kg/ha at farmer location in the coastal Explants like shoot meristem, adventitious buds
region, but the range was 2500–3700 kg/ha in the and microspores can be directly treated with
highland areas. Few salient features of Centenario mutagen, and shoots can also be directly regener-
cultivar like the better yield potential, tolerance ated from the treated explants followed by root
to Sclerotinia sp., colour and size of the grains formation (Suprasanna et al. 2010). Regenerated
were contributing factors in the preference of plants are maintained in the greenhouse and then
Centenario over other commercial cultivar. put under the selection pressure. Similarly
Slabbert et al. (2003) investigated the dryland somatic embryos of vegetatively propagated
yield and growth, community acceptance and crops like banana, date palm, cassava and others
palatability of seven selected amaranth can also be readily induced. The mutagen treat-
(Amaranthus tricolor) and seven cowpea (Vigna ment of embryogenic cell suspension can either
unguiculata) mutant lines. The study indicated eliminate or reduce the chimaeras drastically and
mutant line A19 as the highest yielding and enables to get mutant somatic embryos which
mutant line A5 as the least yielding. The mutant can be subsequently regenerated into plantlets in
lines showed significant morphological variabili- appropriate culture media. Embryogenic cells are
ties. Linear regression analysis indicated the plated on a filter paper and put on agar solidified
amaranth mutant lines A19 and A2 well adapted medium; cells are treated with gamma radiation
to environmental/seasonal changes while A6, followed by the transfer on culture medium and
A550 and A993 poorly adapted. Mutant line A5 allowed to form somatic embryos. The treated
appeared to be the most stable in yield during cells can also be put under the selection pressure
environmental changes, while A554 showed to in order to isolate mutants, e.g. disease-, salt- and
be the most sensitive to environmental changes. drought-tolerant mutants. The selected mutant
Community members opined that A550 was the plants are transferred in the greenhouse and
tastiest of the amaranth mutant lines. finally to the field evaluation and used for cross-
Mutations are induced by physical mutagens ing with other varieties. The lethal dose (LD50
(like gamma radiation, high- and low-energy dose) for each experimental plant should be
beams, etc.) and chemical mutagen (like ethyl determined following the radiosensitive curve to
methane sulphonate or EMS) treatment of both avoid too high or too low dosage. Moreover,
seed and vegetatively propagated crops. Among plants and even varieties differ in radio sensitiv-
physical mutagens, gamma radiation has been ity. Low dose of gamma radiation has promoted
widely applied to induce mutation for both seed growth in Citrus depending on cultivar, main-
and vegetatively propagated crops. Recently ion tained embryogenic nature of date palm for
energy technology – heavy ion beam (HIB) and 2–3 years, promoted growth in orchids, enhanced
low-energy ion beam (LIB) – is being applied to secondary metabolites in medicinal plants and
induce mutation in wide range of crops. HIB is fre- also improved the shelf life of postharvest prod-
quently used for inducing mutations in plants (Jain ucts. Collaborative research programme under
2010). They participate in linear energy transfer the Food and Agriculture Organization and
(LET) and enhance the induction of higher bio- International Atomic Energy Agency (FAO/
logical effects. Several Arabidopsis mutants have IAEA) has given emphasis on crop improvement
been generated through deletions, insertions and by induced mutation using nuclear techniques
chromosomal translocations by HIB. (Jain 2000) intended to produce strains of cereals
8.8 Mutation Breeding 135
with higher concentrations of micronutrients and ‘Genetic Improvement of Underutilised and

improved bioavailability by reduction in the Neglected Crops in LIFDCs through Irradiation
phytic acid concentration. and Related Techniques’ in 1998. The overall
Several mutant genes have been successfully objective was to safeguard food security, enhance
introduced into commercial crop varieties of nutritional balance and promote sustainable agri-
maize, barley, soybean and sunflower to enhance culture in LIFDCs (IAEA-TECHDOC.1426, 2004,
the nutritive value. Traditionally, micronutrient Jain 2009). The species which were given empha-
and trace element content of crops can be sis for study comprised medicinal and aromatic
improved by using field fortification strategies, plants that are important for the West Asia and
applying enriched fertilisers in the soil. North Africa [e.g. argel (Solenostemma argel),
Biotechnological tools have opened up new ave- caper (Capparis spp.), oregano (Origanum syria-
nues to improve the amount and availability of cum), mint (Mentha piperita), liquorice
nutrients in crop plant. These involve simple (Glycyrrhiza glabra), aloe (Aloe spp.), coriander
selection of varieties with high nutrient concen- (Coriandrum sativum), cumin (Cuminum cymi-
tration in the seeds, followed by crossbreeding to num) and henna (Lawsonia inermis)], Andean
incorporate a desired trait within a plant and ulti- grain crops for Latin America [e.g. quinoa
mately genetic engineering for the manipulation (Chenopodium quinoa), canihua (C. pallidicaule)
of the nutrient content of the plant (King 2002). and amaranth (Amaranthus caudatus)] and nutri-
Production of ‘Golden rice’ represents an exam- tive millets for Asia [e.g. finger millet (Eleusine
ple of successful biotechnological application in coracana), Italian millet (Setaria italica) and little
breeding where a gene necessary for the accumu- millet (Panicum miliare)] (Jain 2009).
lation of carotenoids (vitamin A precursors) in The FAO/IAEA Mutant Variety Database or
the endosperm that are not available in the rice MVD collects information on plant mutant vari-
gene pool has been successfully transferred. eties (cultivars) released officially or commer-
Recent advances in plant genomics have cially worldwide. The information includes data
opened new possibilities for application of muta- on the mutagen and dose used, the characters
tion techniques in crop improvement. Using the improved and agronomic data if available. The
reverse genetic strategy called TILLING purpose of the database is to demonstrate the sig-
(Targeting Induced Local Lesion IN Genomes), it nificance of mutation breeding as an efficient
is possible to induce a series of alleles in a target way for safeguarding and promoting global food
locus provided that its sequence is known security, to serve as a platform for breeders to
(McCallum et al. 2000). TILLING strategy was represent their varieties to a global audience and
at first developed for model plant and animal spe- to stimulate germplasm transfer for cultivation,
cies as an approach towards functional genomics, breeding or genomic studies. By 2000, the
but now it has become a valuable parameter in Mutant Variety Database (MVD) had informa-
crop breeding as an alternative to the transgenic tion on 2252 mutant varieties or cultivars evolved
approach. The TILLING techniques require a by mutation and officially released in 59 coun-
high frequency of mutations induced by chemical tries worldwide, mostly in Asia (1 142), Europe
mutagen in combination with a thorough screen- (847) and North America (160). Most of the
ing method for single nucleotide polymorphism desired genetic variations explored in breeding
(SNPs) in the targeted sequence. The applicabil- programmes have occurred naturally and are pre-
ity of this technique for generating a series of served in different germplasm collections.
new alleles in a gene of interest has already been However, when these collections of germplasms
already exemplified in barley, maize and wheat. fail to offer a source for a particular trait, it is
FAO/IAEA has initiated a programme on necessary to depend on other sources of varia-
genetic improvement of underutilised and neglected tion. In such cases, mutation techniques can be
species through a Coordinated Research Project on helpful for the rapid emergence of desired traits.
Though the great majority of induced mutations becomes very expensive for traits that have to be
are recessive and deleterious in nature from a evaluated through tedious phenotypic analysis.
breeder’s viewpoint, it is possible to detect Genetic analysis of mutation-induced changes
desired genotypes from adequately large mutated is another aspect. The irradiation-induced
populations using proper selection tools. Due to changes that occurred in the genome of mutant
these unique probabilities, mutation breeding amaranth can be identified at molecular level
techniques have significantly contributed to plant employing molecular markers. Amaranth acces-
improvement worldwide, have a profound impact sions consisting of both control genotypes and
on the productivity of some crops and gained mutant lines were analysed with microsatellite
popularity among the plant breeders. markers which yielded reproducible polymor-
Three mutant varieties of amaranth have been phic band profile clearly differentiating both the
developed and registered in MVD, viz. Centenario species-specific band profile and mutant line-
in Peru (mutant variety ID No. 2950), New specific profile (Razna et al. 2012). There is a
Asutake in Japan (mutant ID No. 2474) and Sterk possibility to study molecular markers to distin-
in Russian Federation (mutant ID No. 226). The guish γ-radiation-induced changes on the molec-
mutant variety Centenario was evolved by irra- ular level. Amaranthus accessions of control
diation of seed with gamma rays (400 Gy) and genotypes and mutant lines when analysed by
approved in 2006. The improved feature of the microsatellite markers produced reproducible
mutant variety was improved seed production polymorphic banding patterns with a potential to
trait. The variety New Asutake was officially distinguish both the species specificity and
approved in 2006 and developed by irradiation mutant lines specificity, too (Razna et al. 2012).
with gamma rays (500 Gy). Improved trait of the
mutant variety was early maturity. The Sterk
mutant variety was developed by the treatment of 8.9 Polyploidy in Amaranth
hybrid seeds of A. paniculatus and A. nutans by Breeding
watery solution of chemical mutagen. The
improved attribute of the mutant variety was Polyploidy has a great implication to bring about
resistance to drought and medium resistance to sudden abrupt changes in the phenotypes. Some
low temperature. of these may be useful to plant breeding.
Mutation breeding has some advantages along According to Stebbins (1947) ‘polyploidy, one of
with few limitations. The advantages include the best known natural forces of evolution, is the
development of new gene alleles that do not exist most rapid method of producing radically differ-
in collected germplasm (seeds) pools and intro- ent but vigorous and well adapted genotypes’,
duction of that new gene allele for a commercial hence suitable for development as a technique in
variety. So new varieties with desired mutated plant breeding. Polyploids are of wide occur-
alleles can be directly used as a commercial vari- rence, estimated to be around one-third of angio-
ety. The limited genetic changes of any single sperm in nature. Among domesticated species,
plant of a mutated population mostly of recessive polyploids are frequent. Stebbins (1957) pointed
nature enable breeders to develop a new variety out that many of the cultivated crops have origi-
in a short breeding cycle. The disadvantage of nated through polyploidy. Natural hybridisation
mutation breeding is its limitation in generating has played a key role in it. Artificially induced
the desired dominant alleles. It is also less effec- polyploidy is useful in plant breeding in many
tive than crossbreeding for a trait needs a combi- ways, with a definite assumption that polyploidy
nation of multiple alleles, such as tolerance to would revolutionise breeding methods and would
abiotic stresses. The low frequency of mutation help sudden or rapid development of seemingly
needs growing and screening of a large popula- superior genotypes. Proper handling of poly-
tion for selection of desired mutants. This ploids has opened new vistas in plant breeding,
8.10 Genetic Engineering and Biotechnological Approach in Genetic Improvement 137
like (1) rapid development of inbreed lines from and Khoshoo (1977) observed some plants were
haploid, (2) development of tetraploid to sustain either predominantly male or female in the C1
hybrid vigour, (3) rendering of sterile and fertile generation and suggested that later generations
hybrids through amphidiploidy and (4) produc- should be studied to evaluate the stability of grain
tion of intrinsically superior polyploids. tetraploids. Pal and Pandey (1982) found that in
Plant breeders have tried to increase the per- C10 generation tetraploid grain amaranths per-
formance through inducing polyploidy in grain formed similarly to early generations and con-
amaranths (Pal and Khoshoo 1977; Behera et al. cluded that polyploids are stable in Amaranthus.
1974; Sun and Yue 1993) and vegetable ama-
ranths (Madhusoodanan and Pal 1984; Behera
and Patnaik 1975). Polyploids are induced treat- 8.10 Genetic Engineering
ing the seeds with colchicine or by applying it to and Biotechnological
the growing point according to the method of Approach in Genetic
Murray (1940) and Behera et al. (1974). Murray Improvement
(1940) used colchicine solution and observed
that one application of 0.25 % aqueous solution Genetic engineering is not just an extension of
of colchicine, applied to the growing point of A. conventional breeding, but it is also a comple-
caudatus, gave positive results. Madhusoodanan mentary part of it. In conventional breeding
and Pal (1984) generated triploid vegetable A. tri- method, new plant varieties are developed
color with desirable performance, but they were through the selection process and require the
impractical to propagate vegetatively or by large- expression of genetic material which is already
scale production of triploid seeds from diploid- existing within species. Conventional breeding
tetraploid crosses. Pal and Khoshoo (1968) involves process that occurs in nature. Genetic
worked on agronomic disadvantages in grain pro- engineering primarily involves artificial insertion
ducing tetraploids and found a 7 % reduction in of foreign genetic material followed by selection.
the number of female flowers that matured into Conventional breeding believes in mixing of fea-
seeds and the appearance of ‘stray’ predomi- tures from different populations within a species
nantly male plants. Polyploids are shorter and and then selecting plant complemented with dif-
thicker stemmed than normal diploid, and seed ferent features from a population or natural
size increased by 42–159 % with polyploidy (Pal genetic element. However genetic engineering
and Khoshoo 1977; Sun and Yue 1993). relies on incorporation of genetic element in ran-
Tetraploids showed increase in size of various dom location that can disrupt complex gene inter-
plant organs though there was very little differ- actions. Many of the products of such random
ences in plant size; tetraploids flowered a week insertion may yield unexpected effect.
later than the diploids. Amphidiploidy was The amount and availability of nutrient in
induced in sterile F1 hybrid between A. caudatus plant can now be improved applying biotechno-
(n = 16) and A. retroflexus (n = 17). All amphidip- logical strategies. Those strategies include sim-
loids were perfectly fertile and phenotypically ple plant selection for varieties with nutrient-rich
alike. The amphidiploids were much longer than seeds, crossbreeding to incorporate a desired
tetraploid race. The tetraploid of A. caudatus trait within a plant and genetic engineering to
showed 50 % increase in the seed weight. manipulate the nutrient content of the plant. In
Polyploidy was induced in A. hypochondriacus, plant crossbreeding, all genes of the parent
A. edulis and A. caudatus by Misra et al. (1971) plants are combined together, and the progeny
with colchicine treatment. Tetraploids of A. cau- has both desirable and undesirable traits. To
datus had 60 % more protein, more lysine and eliminate undesirable traits, plant breeders used
threonine than diploids. It is concluded that poly- to ‘backcross’ the new plant varieties with other
ploidy has increased grain size and weight with- plants over several generations. Using the
out decreasing the fertility or nutritive value. Pal genetic engineering techniques, the gene(s)
responsible for a desired trait(s) are introduced culture of single cells and meristems can be
in a precise and controlled manner within a rela- effectively used to eradicate pathogens from
tively short period of time. Golden rice, contain- planting material and thereby dramatically
ing carotenoids, and rice with higher amount of improve the yield of established cultivars. Large-
iron are two classical examples of genetically scale micropropagation laboratories are already
engineered plants enriched with nutrients. established supplying millions of plants for the
However, public concerns regarding safety, commercial ornamental market and the clonally
appearance and ethics must be overcome before propagated crop market.
these products can be effectively introduced into
the food supply.
Advancement in molecular biology has 8.11 Genetic Transformation
encouraged the significant development in bio- in Breeding Amaranths
technology which has dramatically enriched our
knowledge to understand plant genomics and Technological innovation in plant biotechnology
manipulate plant gene pool. Marker-assisted is an important catalyst in any crop enrichment
selection (MAS) is one of such techniques which programme. Stable Agrobacterium-mediated
has enhanced the efficiency of breeding for genetic transformation offers advantages in
specific trait, and genetic engineering has made it transferring one or few copies of DNA fragments
possible to transfer genes from almost any carrying the genes of interest. It can be carried
organism into a crop species. The practice of out on the whole plant by applying either tissue
plant breeding has been shifted from the public infiltration techniques (Tague and Mantis 2006)
domain, and thus, selection criteria are increas- or floral dip (Yasmeen et al. 2009). This method
ingly vulnerable to being dominated by private is advantageous over formal tissue culture-based
profit motives rather than public good motives techniques as it directly results into transformed
(Simmonds 1990). seed by passing the lengthy tissue culture process
Plant tissue culture is a promising applied and somaclonal variations (Yang et al. 2009).
branch of plant biotechnology which includes a A protocol for Agrobacterium-mediated
set of in vitro techniques, methods and strategies. genetic transformation of Amaranthus tricolor
Tissue culture techniques are actually part of a was generated through cocultivation of explant
large group of strategies and technologies, with Agrobacterium rhizogenes (Swain et al.
ranging from molecular genetics, recombinant 2010). Bacteria plant-specific factors which
DNA technologies, genome characterisation, influenced transformation were optimised. Of the
gene transfer techniques to aseptic growth of two Agrobacterium strains used, LBA9402 was
cells, tissues, organs and in vitro regeneration of found to be more infectious compared to A4. Few
plants. Tissue culture protocols are available for significant observations were recorded, like
most crop species, though standardisation or greater response of explants from garden-grown
optimisation of methodologies for specific pur- plants to in vitro culture than those from in vitro
pose is yet to be achieved for many crops, espe- cultures and better response of stem internodes
cially cereals and woody plants. Tissue culture than leaves. Significantly higher transformation
methods, in combination with molecular tech- frequency was achieved by immersing pre-pricked
niques, have been successfully utilised to incor- explants in bacterial suspension rather than the
porate specific traits through gene transfer. In direct injection method. The infection of inter-
vitro culture of isolated protoplasts, anthers, pol- node explants with the LBA9402 strain followed
len grains, ovules and embryos has been used to by cocultivation on growth regulator-free MS
create new genetic variation in the breeding lines, medium (MS0) for 5 days resulted in the emer-
often via haploid production. Cell culture has gence of hairy roots up to a maximum frequency
also produced somaclonal and gametoclonal of 97.22 %. Roots were individually cultured in
variants with crop improvement potential. The MS0, but fortified with bactericidal antibiotic
8.11 Genetic Transformation in Breeding Amaranths 139
(500 μg ml−1 cefotaxime). Rhizoclones showing dip has been evolved and successfully applied in
prolific growth were maintained through succes- the popular model plant such as Arabidopsis thali-
sive subcultures in MS0. Opine gene was intro- ana and Medicago truncatula. Agrobacterium-
duced, and expression of transgene was revealed mediated transformation through floral dip
by positive agropine and mannopine synthesis in followed by rapid selection process after trans-
all selected transformed rhizoclones. Shoot genic event has become a phenomenal approach as
regenerated from root clones showed capability it will help to overcome the difficulties anticipated
of auxin-independent growth and opine proficiency. during lengthy tissue culture procedures and
Shoot regeneration was stimulated in MS aug- screening transformed progenies. Munusamy
mented with 2.0 mg l−1 zeatin. Pal et al. (2013) et al. (2013) developed three constructs ( p5b5
have optimised a genetic transformation protocol (14,289 bp), p5d9 (15,330 bp) and p5f7
for major leafy vegetable crop, Amaranthus (15,380 bp)) in pDRB6b vector, with hygromycin
tricolor L., using epicotyl explant cocultivated as a selectable marker gene, which were intro-
with Agrobacterium tumefaciens. Two disarmed duced individually into Agrobacterium tumefa-
A. tumefaciens strains EHA 105 and LBA 4404, ciens strain (AGL1). The bacterial cell suspension
both carrying the binary plasmid p35SGUSINT was applied to Amaranthus inflorescence drop by
containing the neomycin phosphotransferase II drop and left to produce seeds (T1). The T1 seeds
gene (nptII) and the β-glucuronidase gene (gus), were germinated and grown to raise the seedlings
were evaluated as vector. The former exhibited a under non-sterile condition. Hygromycin selection
higher transforming efficiency. Several key fac- of seedling on cotyledonary leaves resulted in
tors influencing the transformation events were identification of 12 putative transformants, three
optimised. from construct p5b5, four from construct p5d9 and
Mature embryos of Amaranthus hypochon- five from construct p5f7. All positive putative
driacus (amaranth) were involved to develop an transformants that were selected at the first stage
in vitro culture system to be utilised for plant or primary screening through hygromycin spray-
regeneration and genetic transformation (Jofre- ing also showed positive result in leaf disc hygro-
Garfias et al. 1997). Plants were regenerated from mycin assay. A ~750 bp amplified hygromycin
embryo-derived callus cultured on Murashige gene was further verified through sequencing. The
and Skoog medium supplemented with 10 μM results indicated that Amaranthus inflorescences
2,4-dichlorophenoxyacetic acid or 3,6-dichloro- could be transformed easily, and the transformed
2-methoxybenzoic acid and 10 % coconut liquid progenies could be identified through a combina-
endosperm. Transgenic plants were generated by tion of simple and rapid methods.
infection of mature embryo explants with a dis- The plant metabolite oxalic acid is commonly
armed Agrobacterium strain containing the plas- considered as a food toxin having negative effects
mid pGV2260 (pEsc4), harbouring the genes on human nutrition. Oxalic acid is present as
encoding neomycin phosphotransferase type II nutritional stress in many crop plants like ama-
and β-glucuronidase. The presence of transgenes ranth and Lathyrus. Oxalic acid has also been
in the genome of transformed amaranth plants reported to be involved in the attacking mecha-
and in their progeny was detected by Southern nism of several phytopathogenic fungi. So
blot hybridisation. A pea chlorophyll a/b-binding degradation of oxalic acid in the crop plants
protein promoter was used for tissue-specific and through introduction of a specific gene would
light-inducible expression in transgenic amaranth have twofold advantages: firstly, nutritional stress
plants and their progeny. will be minimised, and secondly resistance to
The success in the transformation of fungal pathogen will be conferred. Oxalic acid is
Amaranthus by floral dip would broaden the pos- degraded by two major pathways, i.e. decarbox-
sibilities for the exploitation of the available ylation and oxidation. Decarboxylation of the
reported method on many other alternative crops. oxalic acid takes place either by conversion of
Agrobacterium-mediated transformation by floral oxalic acid to oxalyl CoA by means of oxalyl-
CoA decarboxylase or directly to CO2 and formic the transgene showed oxalate decarboxylase activ-
acid by oxalate decarboxylase. In oxidation pro- ity both in vitro and in vivo conditions. Transgenic
cess oxalic acid is broken down to CO2 and H2O2 tobacco and tomato plants expressing oxalate
that has been detected in plants. Mehta and Datta decarboxylase activity showed significant resis-
(1991) earlier reported purification and partial tance to fungus Sclerotinia sclerotiorum that uses
cDNA cloning of oxalate decarboxylase from oxalic acid during infestation. The work also dem-
Collybia velutipes. The enzyme has several onstrated a novel approach to develop resistance
advantages over other oxalate degrading against fungal infection through transgenesis
enzymes. Firstly, OXDC1 is specific for oxalate, (Kesarwani et al. 2000; Chakraborty et al. 2013).
and it catabolises oxalic acid to formic acid The transformation procedure using the cDNA of
(nontoxic organic acid) and CO2 in a single step oxalate decarboxylase followed in the case of
without requiring a cofactor. Secondly, the tomato and tobacco can be considered as standard
enzyme is active at low pH which would be protocol to transform vegetable amaranths where
helpful as most of the oxalates are localised in oxalate content to some extent is lowering its
plant cell vacuoles, where pH is low. acceptability.
Decarboxylative breakdown of oxalic acid is Protein is essential for normal physical growth
catalysed by oxalate decarboxylase (OXDC) in a and development. So deficiency of protein is
substrate-specific reaction, producing formic the most crucial factor regarding increase of
acid and carbon dioxide. Little success has been morbidity and mortality especially in developing
achieved in attempts aimed at reducing oxalic acid countries. Efforts to improve protein quality
levels and understanding of the biological signifi- and quantity have received limited success.
cance of OXDC in crop plants. To study the func- Improvement in nutritive values of crop plants, in
tion of OXDC and the metabolic consequences of terms of amino acid composition and availability
oxalate downregulation in a heterotrophic, oxalic of essential amino acids, has been a major long-
acid-accumulating fruit, Kesarwani et al. (2000) term objective of plant breeding programmes.
generated transgenic tobacco and tomato (Solanum Raina and Datta (1992) reported earlier the clon-
lycopersicum) plants expressing an OXDC ing of a novel seed albumin gene AmA1 from
(FvOXDC) isolated from the fungus Flammulina Amaranthus hypochondriacus. The AmA1 pro-
velutipes specifically in the fruit. These fruits of tein was purified to homogeneity. It is nonaller-
transgenic plant showed up to a 90 % reduction in genic in nature and rich in all essential amino
oxalate content, with concomitant increases in cal- acids, and the amino acid composition comple-
cium, iron and citrate. A full-length cDNA for ments well with standard for optimal human
oxalate decarboxylase was isolated by using nutrition as proposed by the World Health
5′-rapid amplification of cDNA ends polymerase Organisation. The protein is a 35 kDa monomer
chain reaction of a partial cDNA as cloned earlier with four isoforms that can be separated by chro-
(Mehta and Datta 1991). Genomic library matography. A full-length cDNA of AmA1 pro-
from Collybia velutipes was screened with this tein was characterised and isolated. The AmA1
cDNA as a probe, and a genomic clone was iso- gene is expressed during early embryogenesis.
lated and sequenced. Genomic sequence was ana- Germinated seeds did not contain any AmA1
lysed and compared with the cDNA sequence mRNA; no RNA was detected in leaves, root or
which revealed that the cDNA was interrupted other parts of the plant indicating the tissue-
with 17 small introns. The cDNA has been suc- specific expression of the gene. The AmA1-
cessfully expressed in cytosol and vacuole of coding sequence was successfully introduced in
transgenic tobacco and tomato plants. The trans- potato to improve the nutritive quality of potato,
genic plants showed normal phenotype, transfor- and expression of the gene was tuber-specific and
mation was stable and the transferred trait was constitutive nature. The experiment showed
stably inherited to the next generation. The recom- unique outcome. The transgenic population
binant enzyme showed partial glycosylation, and showed a significant increase in growth and
8.11 Genetic Transformation in Breeding Amaranths 141
production of tubers, as well as both qualitative quality or yield. The applicability of genetically
and quantitative improvements of protein. altered plants on commercial scale depends on
Chakraborty et al. (2000) reported the develop- several factors like stable integration and
ment of transgenic potatoes with enhanced expression of the transgene under the different
nutritive value by tuber-specific expression of genotypic backgrounds of the host species, wider
an AmA1 seed albumin protein gene, in seven environmental adaptability and sustainable pro-
genotypic backgrounds suitable for cultivation in duction, including food safety. In vitro and
different agro-climatic regions. Transgenic tubers in vivo studies on experimental animals demon-
showed up to 60 % increase in total protein con- strated that the transgenic tubers are also safe for
tent and a significant increase in the concentra- human consumption.
tions of several essential amino acids which are The nutritive value of wheat is mainly charac-
otherwise limited in potato. Moreover, this terised with low content of essential amino acid
genetic manipulation also exhibited enhanced like arginine, lysine, hystidine and threonine.
photosynthetic activity, moderate increase in During milling process with the removal of aleu-
tuber yield and a concomitant increase in total rone layer and embryo tissue, high amount of
biomass. The comparison of protein profile sug- essential amino acids is lost. Several attempts
gested that the proteome rebalancing might have have been made to improve the amino acid con-
caused increased protein content in transgenic tent by conventional breeding methods. Recently
tubers (Chakraborty et al. 2010). Furthermore, as a pioneering achievement, herbicide-resistant
the data on field performance and safety evalua- transformed wheat has been developed through
tion indicated that the transgenic potatoes are microprojectile bombardment of regenerable
suitable for commercial cultivation. embryonic callus (Vasil et al. 1992) which has
Transgenic potato plant with engineered opened up a new avenue to improve agronomic
AmA1 seed storage protein gene in tuber might traits of transgenic crops, as well as the grain
have utilised the free amino acids for the synthe- composition and quality. An attempt was made to
sis and accumulation of the storage protein caus- express recombinant storage protein with the
ing a depletion in endogenous amino acid pool. appropriate amino acid profiles in the wheat
Such depletion of endogenous free amino acid endosperm tissue to improve the essential amino
might have triggered photosynthetic activity and acid content in wheat flour (Cecilia 2009). The
increased the photosynthetic rate. It is evidenced transgene AmA1 was stably integrated into the
that increased photosynthesis is responsible for wheat genome by a competent wheat endosperm-
the increase in protein synthesis and yield. Thus specific promoter (1Bx17 HMW-GS), expressed
in transgenic potato, the increased rate of photo- in the wheat endosperm and transmitted to the
synthesis increased total protein content. Seed next three generations. The transformation
storage proteins are considered to act as the sink pattern of the foreign gene showed tissue speci-
to control the movement of photosynthate into ficity. The amount of in vivo expression of AmA1
developing organs. It is suggested that AmA1 protein gene showed simultaneous increase in the
seed storage protein probably acts as a sink pro- essential amino acid content of the transgenic
tein in transgenics, thereby regulating the move- flour with no significant change in the total pro-
ment of metabolites, specially the amino acids, tein content. Nutritive value of several other
into the developing tuber where they are utilised crops can be enriched by transformation with the
to synthesise new proteins and promote growth novel AmA1 protein gene.
(Chakraborty et al. 2010). This study can be All the experimental outcome emphasised the
treated as a major breakthrough in gene expression fact that expression of novel AmA1 gene might
and translational research in which a genetically be a potential strategy for the nutritional improve-
engineered plant with a transgene of seed storage ment of food crops. Transformation of other
protein has led to nutritional improvement with crops with this unique AmA1 protein gene is quite
essentially no negative collateral effects on crop feasible.
Unwanted transgene escape occurs in several (RFLPs), microsatellites, simple sequence

crops like oilseed, rapeseed, maize, cotton, etc. repeats (SSRs), etc., satisfy the above parame-
They are found in the variant cultures and wild ters. The development of molecular markers
types as well as in hybrids. The consequence of like microsatellites and SSRs requires a prior
the continuous transgene escape in the cultivation knowledge of DNA sequence. A few dominant
field cannot be predicted reliably, because that and universal molecular markers such as random
depends surely on the ability of the transgene to amplified polymorphic DNA (RAPD) and ampli-
influence the fitness of the plant containing the fied fragment length polymorphisms (AFLPs)
introgressed gene. It would not be logical to wait have also been employed to analyse the variability
until something adverse happens. Transgene of in Plant Genetic Resources.
flows destroy the gene pool at its centre of origin A heterotic clustering of genotypes appropriate
and domestication. Care must be taken to minimise for hybrid breeding represents a different
the transgene escape into the wild as effectively approach of diversity analysis at the molecular
as possible applying all available measures during level with a principle of searching out a positive
developing genetically modified (GM) crops. correlation between genetic distance and hetero-
sis, i.e. the more the distance between two geno-
types of a crop species are, the more will be the
8.12 Molecular Markers heterozygosity genetically. Hybrids originating
and Genome Research from crosses between them are expected to show
for Using Genetic Resources heterosis (Melchinger et al. 1999; Reif et al.
in Plant Breeding 2003a, b). Since high degree of heterosis does
not necessarily mean high hybrid yield, so the
The utility of molecular markers and genome crossing effect on heterosis and hybrid perfor-
research regarding the utilisation of Plant Genetic mance needs to be differentiated and evaluated
Resources (PGR) for crop improvement include separately. Recent studies have shown that if the
several aspects: molecular markers applied for genetic diversity
assessment are linked to performance QTLs,
1. Studies on diversity to differentiate genetically rather than from using neutral markers, the cor-
similar or distinct accessions and to determine relation between diversity measures and hybrid
individual degrees of heterozygosity and het- performance appears more consolidated and
erogeneity within PGR populations conclusive (Vuylsteke 1999; Vuylsteke et al.
2. Genetic mapping to identify markers in close 2000; Jordan et al. 2003).
proximity to genetic factors affecting quanti- Marker-assisted selection (MAS) can be helpful
tative trait loci (QTLs), followed by marker- (i) to select individuals carrying molecular mark-
assisted selection (MAS) of desired genotypes ers which are linked to the trait of interest, instead
in segregating populations of exercising extensive phenotypic evaluation
3. Exploitation of valuable QTLs from PGR (foreground selection), and (ii) to minimise
4. Studies on the allelic variability of PGR collec- unwanted parts of the donor genome, including
tions and identification of those alleles which the linkage drag (background selection).
are beneficial for important agronomic traits Foreground selection needs a strong linkage
between the trait of interest and its flanking
Molecular markers should be selectively neutral, markers for which one is selecting. Background
highly polymorphic, co-dominant, well dispersed selection requires genotyping with a larger
throughout the genome and cost- and labour- number of markers, covering the whole genome.
efficient to be utilised efficiently in diversity MAS has been proved efficient for the transfer of
analysis (Bretting and Widrlechner 1995). Protein simply inherited qualitative traits from germ-
markers like isozymes and DNA markers like plasm resources into elite breeding lines applying
restriction fragment length polymorphisms backcross procedures. It is particularly useful for
8.13 Molecular Breeding in Amaranths 143
the traits which are recessive in nature, can be damage and repair, we should be able to design
assessed only after flowering or that are very dif- strategies of mutagenic treatment to significantly
ficult and expensive to assess. Through combined increase the mutation frequency. When the genes
application of both foreground and background responsible for DNA damage repair are deleted
selections, the transfer of monogenic trait from a or mutated, these breeding lines could become
germplasm resource into an elite breeding line highly vulnerable and sensitive to mutagenic
may be completed within 3–4 generations in treatment, and frequency of mutation could be
place of usual six generations of conventional significantly enhanced. Such lines are commonly
laborious backcrossing with the same proportion termed as ‘super mutable genetic lines’. Once
of the recurrent parent genome (Ragot et al. 1995; mutation of important traits is induced and iden-
Frisch et al. 1999). MAS for multigenic, quanti- tified in such ‘super mutable genetic lines’, the
tative traits at first requires the identification of transgenes and mutated genes can be separated
the genomic regions (QTLs) that affect the trait through self-crossing as they are not linked to
of interest. In classical QTL mapping, a segregat- one another; hence non-transgenic stable mutant
ing population (e.g. F2, F3 or recombinant inbred lines could be generated. In conventional muta-
population) is developed from two inbred lines. tion breeding, mutations generated through
This mapping population is evaluated for the induced mutation are used either directly or indi-
trait(s) of interest. Simultaneously, the popula- rectly (through crosses with other varieties) to
tion is genotyped with a number of markers, and develop a new variety or elite breeding line. It is
a genetic map is constructed from the marker now possible to tag the mutated genes, accumu-
data. In the final QTL analysis, data is analysed late them into a single elite breeding line and fol-
for co-segregation of particular markers with the low them up in subsequent breeding
trait of interest which is ultimately followed by programmes.
transfer of favourable QTL alleles into elite Improved knowledge on molecular genetics
breeding lines via pure MAS or MAS in combi- and genomics and better understanding of geno-
nation with phenotypic selection. typic variability and rapidly emerging molecular
techniques have enabled plant breeders to apply
induced mutation techniques in plant breeding to
8.13 Molecular Breeding evolve novel varieties more judiciously and pre-
in Amaranths cisely than ever before. Plant molecular mutation
breeding has emerged as a strong, efficient proce-
In general, plant molecular breeding depends on dure for plant improvement in a very short period
the understanding of the genetic control of target of time in which genomic and molecular infor-
traits of interest at the molecular level. mation are utilised in framing the breeding strate-
Information on molecular genetic is also of great gies, screening, selection and verification of
help in framing a proper mutation breeding strat- induced mutants and in the subsequent utilisation
egy. Firstly, it is important to assess the feasibil- of mutated genes in the breeding process. It is
ity and potentiality to induce a mutation of based on the knowledge of DNA damage, repair
interest. Secondly, a mutation may have pleiotro- and mutagenesis, plant molecular genetics and
pic effects if the gene is situated at the upstream genomics of important agronomic traits as well
or at the middle of a long biosynthetic pathway. as induced mutations.
Thirdly, knowledge of genes controlling a trait of Responses of different genotypes and plant
interest would constitute the very basis of the tissues to different mutagens, usually measured
TILLING (Targeting Induced Limited Lesions using lethal doses (LD), genetic chimaeras
IN Genomes) method. Induction of mutation is formed due to mutagenic treatment and their
the starting point in mutation breeding, and a low effect on transmission of mutated alleles and seg-
frequency of mutation has been a severe limiting regation in the following generation. Such knowl-
factor. Equipped with knowledge of the DNA edge of genetics is very significant to determine
proper doses and modes of mutagenic treatment, relates well with expression level of DHDPS
as well as for the methodology of growing M2 gene, suggesting eQTL for high-lysine pheno-
populations and harvesting. type, though expression profiles of all other
In spite of great potentiality of grain ama- enzymes comprising AK show no correlation
ranths as a source of nutritious food, little efforts with high-lysine content in seeds. Annotations of
have been devoted to explore its genomics. A all the genes in the lysine biosynthetic pathway
partial transcriptome of A. hypochondriacus has using comparative genomics and expression
been reported for the understanding of the mech- analysis offer insight into high-lysine phenotype.
anism adopted by the species of the family The proposed draft genome (Sunil et al. 2014)
Amaranthaceae in response to the environmental can be very useful for the advancement of the
stress (Delano-frier et al. 2011). A limited understanding of diverse phenotypes that are
sequencing of the genome of A. tuberculatus has unique to amaranths like rich nutritional profile,
been done to understand the mechanism applied rapid growth, drought resistance, wide adaptabil-
by the species to resist three herbicides. A more ity and characterisations of genes for betalain
systematic and concrete approach towards pigment.
exploring the amaranth genome includes creation It has been well documented that DNA is sub-
of BAC libraries for all three grain amaranths jected to continuous damage, and the cell has its
(Maughan et al. 2008), development of microsat- own machinery to address such injury. There are
ellite markers (Mallory et al. 2008) and construc- different ways for the repair of DNA damages
tion of linkage map using single nucleotide caused by different types of mutagen. This
polymorphism (Maughan et al. 2010). Studies on knowledge is very important for proper framing
the genomics of grain amaranths in respect to of mutagenic experiments, so that an enhanced
their unique nutritional properties and C4 photo- mutation frequency can be achieved. Cells with
synthesis among the edible dicots are limited. damaged DNA will survive only when these
Keeping in view these objectives, Sunil et al. damages are repaired either correctly or errone-
(2014) working on genome and transcriptome of ously. The repair done erroneously will be fixed
A. hypochondriacus reported a draft genome as in the genome as induced mutations. The nature
the first of its kind of grain species highlighting of DNA damage caused by different types of
the high-lysine phenotypes and C4 evolution mutagens to a great extent determines the molec-
under Caryophyllales. Out of 411 linkages, sin- ular feature of induced mutations.
gle nucleotide polymorphisms (SNPs) derived
from homozygous region of the grain amaranths,
355 SNPs (86 %) were reported to be present in 8.14 Plant Tissue Culture
scaffolds and 74 % of the 8.6 billion bases of the Techniques in Amaranth
sequenced transcriptome map to the genomic Breeding
scaffold. The genome of A. hypochondriacus
reported codes for at least 24829 proteins and Currently, there are not much published reports
contains 13.76 % of repeat elements. They have on the tissue culture of Amaranthus. Probably the
also placed two workable hypotheses for high- first report on in vitro growth and morphogenetic
lysine phenotype including gene number poly- response was published by Flores et al. (1982).
morphism for AK (aspartate kinase) and eQTL of He has developed an in vitro culture system for
DHDPS (dihydrodipicolinate synthase). Lysine both grain and vegetable amaranths. Leaf discs
biosynthesis is mainly regulated by two allosteric and hypocotyl segments from 2 to 3 weeks old
enzymes aspartate kinase and dihydrodipico- seedlings of A. hypochondriacus, A. cruentus and
linate synthase. Any kind of polymorphism A. tricolor were cultured in MS and B5 media
within these two gene loci is expected to have a supplemented with 2,4-dichlorophenoxyacetic
correlation with high-lysine phenotype in ama- acid, α-naphthyl acetic acid, benzyladenine and
ranths. The high free lysine content in seeds cor- zeatin in various combinations. The tissue culture
8.14 Plant Tissue Culture Techniques in Amaranth Breeding 145
showed various responses like rapid callus ements, 3 % sucrose, 0.8 % agar noble, pH 5,8,
growth, abnormal root formed on the leaf disc without the addition of hormones. Chemical
and embryo-like structure developed from the compositions of the culture media containing
surface and veins on the disc with slight modifi- inorganic salts and hormones have an important
cation of media. Viable protoplasts from primary role to play for successful aseptic in vitro culture
leaf mesophyll tissue were isolated from all the of explants. Preferential depletion of some ele-
three species enzymatically. Protoplast culture is ments leads to symptoms of deficiency or toxic-
of great potential for genetic manipulation of ity, sometimes with necrosis of the inoculum.
plant. Primary result indicated that protoplast The underexploited crops, including amaranth,
from A. tricolor can expand and form a new cell offer a special challenge for the use of in vitro
wall. This was a pioneering work in amaranth tis- approaches, because extensive efforts are
sue culture forming a base for future biotechno- required from the plant breeders to select and
logical work like in vitro propagation, germplasm improve this plant material. In vitro systems have
preservation and transfer, and agronomic important practical applications not only for
improvement through genetic manipulation. rapid breeding of this rediscovered crop but also
Shoot tip-derived plantlet of A. caudatus, A. tri- for producing cell biomass to be used as source
color, A. hypochondriacus, A. retroflexus and A. of phytochemicals of practical interest.
viridis flowered in vitro following 8–32 weeks of Tissue culture studies in amaranths are just
culture. Shoot tip was cultured on MS medium beginning. There are several areas which can be
containing salt and 30 g/l sucrose, 100 mg/l myo- emphasised, like the control of oxalate/nitrate
inositol, 0.4 mg/l thiamine and 8 g/l agar. content in cell culture may provide clues regard-
Addition of NAA (0.1 mg/l) enhanced inflores- ing regulation of nitrogen assimilation and con-
cence production but was not necessary for sequently may lead to the selection of low-oxalate
flower production. The fruits produced by A. cell/plant. The wide variability available in the
gangeticus and A. retroflexus dehisced, and their germplasm can be used to identify cell lines more
seeds were dropped on the surface of agar tolerant to stress and also to involve in plant
medium, where they germinated immediately. regeneration system. Amaranth germplasm rep-
Bennici et al. (1992) studied on several species resents a potent source of herbicide-resistant
and varieties of Amaranthus and showed its trait. This resistance could be introduced in culti-
potential with regard to dedifferentiation and vated form (grain and vegetable) via protoplast
morphogenetic responses in vitro. On the back- fusion with their weedy relatives.
ground of these studies, Guidea et al. (2012) In spite of significant potential, the absence of
reported micropropagation of selected genotypes adequate knowledge on basic process governing
and their subsequent exploitation, rescue of the cell division and differentiation pattern in ama-
genetic variation or induction of new variation, ranths is creating obstacles before the application
phytoremediation studies and use of cell biomass of cell and tissue culture techniques for their
to obtain phytochemicals of practical interest. improvement. Conditions required for induction
The study was to evaluate the growth and mor- of somatic embryogenesis are determined empir-
phogenetic responses of various types of explants ically. Definite guideline is yet to be standardised,
from three varieties of Amaranthus species: except for few model systems. Research is slowly
Amaranthus cruentus ‘Amont’, Amaranthus gaining momentum towards a primary under-
hypochondriacus ‘Intense Purple’ and standing of the process involved, through the use
Amaranthus ssp. ‘Plenitude’. The seeds belong- of mutant cell lines (Sung and Dudits 1981).
ing to these three varieties of Amaranthus species Though application of tissue cultural methodolo-
were germinated in vitro in aseptic conditions on gies in amaranth is limited, still there are several
MS basal medium containing half strength as areas where tissue culture can be of great
regards the concentration of macro- and microel- potential.
Evolution of Sexuality
in Amaranths 9
9.1 General these derivatives having sexual compatibility,

gynomonoecious member might have originated.
The genus Amaranthus is unique in showing a Monoecy in amaranths might have transmitted
wide range of sexuality from gynomonoecy, from cosexual species of Chenopodiaceae-
monoecy to dioecy and also variability in mating Amaranthaceae alliance or Chen-Am alliance.
behaviour from self-compatibility to obligate Molecular analysis of the Caryophyllales estab-
outcrossing, i.e. from monomorphic reproductive lished Chen-Am alliance as a monophyletic
system to dimorphic reproductive system. lineage.
Bisexual flower represents the ancestral condi-
tion in angiosperm, and monoecy is considered to
have been derived from bisexual condition 9.2 Trend of Sexuality in Plants
through intermediate gynomonoecious or andro-
monoecious forms. Dioecy results in a division of Flowers are the most varied structure in the flow-
labour between sexes and may ultimately lead to ering plants. Such variability is a key factor and
greater reproductive efficiency. A variety of primarily instrumental to achieve mating success.
hypotheses have been proposed for evolution of Selective forces are responsible for origin and
dioecy from hermaphroditism through monoecy, maintenance of sexual diversity in flowering
gynodioecy and androdioecy. Monoecy in the plants. Flowering plants are mostly hermaphro-
grain amaranths are supposed to have originated dite producing bisexual flowers. Few flowering
from the dominant cosexual form having exclu- plants (−10 %) have unisexual flowers with a
sively bisexual flower through sterility mutation wide spectrum of gender strategies that involve
followed by subsequent specialisation in flower varied combinations of female, male and her-
form and position. Dioecy may have evolved maphrodite flowers at the plant population level.
from monoecy through disruptive selection on Though dioecy is widely distributed in flowering
male and female reproductive allocation, fol- plants occurring in nearly half of all families,
lowed by gender specialisation ultimately lead- overall frequency of dioecious species is only
ing to unisexual plants. In the case of vegetable 6 % (Renner and Ricklefs 1995) signifying its
amaranths, genetic modification in female fertil- rare association with successful evolutionary
ity of hermaphrodite population might have diversification. Dioecy is commonly associated
resulted in the formation of bisexual member with unspecialised pollination system that
with non-functional gynoecium. Later male ste- involves wind, water or generalised pollinator
rility gene might have played a role to give rise to rather than the more specialised pollinator that
female member. Subsequent inbreeding between commonly drives floral diversification and

148 9 Evolution of Sexuality in Amaranths
reproductive isolation in many cosexual flower- transition between these two systems. Establishment
ing plants (Johnson et al. 1998). Sexual structure of monoecy requires specialisation in the shape,
can be spatially separated in a flower or can size and positioning of the male and female
become functional at different times reducing flower (Shmida et al. 2000). But before such spe-
the likelihood of self-pollination. Gradual cialisation, a mutant with some unisexual flowers
sexual diversity has emphasised the mechanism needs to be established first. Only after the uni-
that aimed at promoting outcrossing and sexual mutant is firmly established, subsequent
reducing likelihood of inbreeding depression. mutation related to specialisation in flower form
Conventionally floral traits have been considered and position has given rise to monoecy.
as anti-selfing mechanism that encourages cross- Many investigations have been directed
pollination preventing self-pollination towards exploring traits leading to dioecy (Renner
Strong empirical evidences indicate that and Won 2001; Gleiser and Verdu 2005; Case
bisexual flower represents the ancestral condition et al. 2008). A variety of hypotheses have been
in angiosperm (Richard 1997; Doyle 1998; proposed for evolution of dioecy from hermaph-
Endress 2001). Monoecy with separate male and roditism through monoecy, gynodioecy and
female flowers on the same plant is considered to androdioecy. But not all these pathways have
be derived condition from bisexual condition received the same theoretical attention. There are
(Mitchell and Diggle 2005). Monoecy has tra- no well-authenticated cases of androdioecy as an
ditionally been considered as originating from intermediate stage in the evolution of dioecy and
gynomonoecy or andromonoecy (Wilson 1979; very limited evidence for the evolution of andro-
Bawa and Beach 1981; Bertin 1982). Many dioecy from hermaphroditism. Two fundamental
research have been directed to explore the evolu- evolutionary pathways for the origin of dioecy
tionary transition giving rise to dioecious condi- through monoecy and gynodioecy are generally
tion, i.e. male and female flowers on separate recognised. Both involve the transition from gen-
plants (Renner and Won 2001; Gleiser and Verdu der monomorphism to dimorphism. In the gyno-
2005; Case et al. 2008). Dioecy results in a dioecy pathway male sterility genes spread in
division of labour between sexes and may ulti- bisexual population, resulting in an intermediate
mately lead to greater reproductive efficiency stage that involves females and hermaphrodites.
(Lloyd 1982). Male and female reproductive Genetic modifiers of female fertility gradually
function may be optimised through different convert hermaphrodites to male resulting in
selective processes ultimately leading to pheno- dioecy. The monoecy pathway is less well inves-
logical dimorphism in male and female plant. tigated, assumed to involve disruptive selection
Despite seemingly complex pattern of sexual on male and female allocation in monoecious
diversity in flowering plants, two broad and fun- population which gradually increased gender
damentally distinct patterns of gender variation specialisation until unisexual plants originated.
or sexual system have been recognised – mono- Monoecy has originated several times from the
morphic sexual system and dimorphic sexual dominant cosexual condition in angiosperm
system (Bawa and Beach 1981). In the former population having exclusively hermaphrodite
system, species bears bisexual female and/or flower. This might have occurred through sterility
male unisexual flowers on the same individual, mutation that produced unisexual flowers.
leading to monoecious, gynomonoecious, andro- A lot of investigations have been done on the
monoecious and trimonoecious condition. In the gynodioecy, but till date little attention has been
latter system, species have dimorphic sexual given to monoecy pathway. It is not clearly
system, i.e. unisexual individual such as dioecy, resolved whether the transition is determined
gynodioecy and androdioecy. solely by selection or quantitative genetic varia-
Despite the simultaneous/common occurrence tion governing sex allocation or whether major
of hermaphroditism and monoecy, little attention male sterility genes are also having a role. A
has been paid to the factors favouring evolutionary recent molecular phylogenetic study (Renner and
9.3 Sexuality in Amaranths 149
Won 2001) provides convincing evidence of sexuality from gynomonoecy, monoecy to dio-
multiple origins of dioecy from monoecy in the ecy. Dioecy is the rare breeding system among
primary Neotropical shrub family Siparunaceae. angiosperm. It may affect the ability of a lineage
It is very vital to determine the circumstances to avoid extinction or encourage speciation.
under which selfing rate increased resulting in Monoecy is the predominant phenomena in
inbreeding depression in ancestral cosexual amaranths. Grain amaranths are exclusively
population. Such condition facilitates the spread monoecious, while vegetable amaranths are pre-
of unisexual variants that favour outcrossing. A dominantly monoecious with exception. It would
link between origin of gender dimorphism and be logical to presume that both grain and vegeta-
conditions that promote inbreeding depression ble amaranths were evolved and domesticated in
might occur through the action of polyploidy. their respective centre of origin from their weed
Chromosome doubling in plants can result in the progenitor having wide range of variability in
inhibition of self-incompatibility leading to self- sexuality ranging from monoecy to dioecy.
compatibility (Chawla et al. 1997) providing Amaranths followed the general evolutionary
opportunities for self-fertilisation. Molecular phy- trend in sexuality, i.e. from monoecy to Dioecy. It
logeny in North American Lycium (Solanaceae) can be presumed that monoecy in amaranths
indicates that gender dimorphism has evolved might have transmitted from cosexual species of
only in species that are polyploid and self-com- Chenopodiaceae-Amaranthaceae alliance or
patible but their close relatives are diploid and Chen-Am alliance. The Chen-Am alliance is of
self-incompatible. Such pattern also found to have worldwide distribution, comprising 2400 species.
originated independently in South African species The alliance is noted for the evolution of C4
of Lycium. Polyploidy might have acted as a trig- photosynthesis, halophytism, xerophytism and a
ger for the evolution of gender dimorphism variety of breeding system. The close relation-
(Miller and Venable 2000). Large plant size might ship of Chenopodiaceae and Amaranthaceae has
be another condition that can potentially lead to been recognised based on core floral formula
gender dimorphism (de Jong and Klinkhamer consisting of five tepals, five stamens and 2–3
1994). In plant species with large stature (like carpels (Hershkovitz 1989). Molecular analysis
shrubs and trees), a considerable amount of self- of the Caryophyllales (Manhert and Rettig 1994;
ing can occur due to the presence of many open Downie and Palmer 1994; Downie et. al. 1997;
flowers on a plant at the same time leading to Cuenoud et. al. 2002) established Chen-Am
inbreeding depression. Recent marker gene analy- alliance as a monophyletic lineage.
sis of the clonal aquatic plant Sagittaria latifolia The grain amaranths are grown and consumed
offers evidences that indicate that geitonogamous both as green vegetable and pseudocereal. Though
selfing and strong inbreeding depression have grain amaranths are crops of America but later
influence in evolutionary transition from mon- they migrated to Asia and Southeast Asia, Europe
oecy to dioecy (Dorken et al. 2002). Population of and North America. Vegetable amaranths spe-
both sexual systems occur together in this species cially Amaranthus tricolor and A. dubius proba-
making it an excellent model for studying the evo- bly originated and domesticated in Asia and
lution of combined versus separate sexes. Southeast Asia (Grubben and Van Sloten 1981)
and later spread throughout tropical and temperate
regions of Africa, Central America and Europe by
9.3 Sexuality in Amaranths the immigrants (Martin and Telek 1979).
Amaranthus blitum aggregate included in sect.
The genus Amaranthus is unique in mating Blitopsis of subgen. Albersia, comprising A. bli-
behaviour ranging from self-compatibility to tum and A. emarginatus, was studied morphomet-
obligate outcrossing, i.e. from monomorphic rically (Das and Iamonico 2014). Amaranthus
reproductive system to dimorphic reproductive blitum with two varieties are supposed to have
system. It shows a wide range of variability in originated and domesticated in Mediterranean
150 9 Evolution of Sexuality in Amaranths
Fig. 9.1 Evolution of

sexuality in vegetable
amaranths
Influence of Male Modification of
Sterility gene female fertility
Hermaphrodite plant of
Chen-Am alliance
Inbreeding
Female plant Hermaphrodite plant with
non-functional gynoecium
Gynomonoecious plant
Optimization of reproduction
through various selection process
Monoecious plant
Basin, Europe and North America and A. emar- hermaphrodite Chen-Am member through various
ginatus in tropical America. The member of ‘bli- processes like spread of male sterility genes and
tum complex’ later might have migrated to Asia, gender modifier gene of female fertility in her-
Southeast Asia and other parts of the world. maphrodite population followed by optimisation
Amaranthus bengalense, a new variant of A. bli- of male and female reproductive function through
tum, was reported from West Bengal, India (Das different selective processes. In case of vegetable
and Iamonico 2014). A new gynomonoecious spe- amaranths, genetic modification in female fertility
cies Amaranthus parganensis Saubhik Das was of hermaphrodite population might have resulted
discovered from Lower Gangetic Plain of West in the formation of bisexual member with non-
Bengal that closely resembles A. tricolor of sect. functional gynoecium. Later male sterility gene
Pyxidium subgen. Albersia (Das 2015). The new might have played a role in giving rise to female
species shows structural gynomonoecy with rudi- member. All these derivatives originated through
mentary gynoecia in bisexual flowers and pro- genetic modification and have sexual compatibil-
vided a clue regarding origin of monoecy through ity. Subsequently, through inbreeding process
intermediate sexual system. Dioecious species are gynomonoecious member might have originated.
confined to a small area in North America (Sauer Monoecy is a derived condition originated from
1957), though sporadic appearance has been gynomonoecious condition through optimisation
reported from time to time (Brenan 1961). of male and female reproduction through different
Monoecy in amaranths may have evolved from selection processes (Fig. 9.1). Further evolution of
9.3 Sexuality in Amaranths 151
sexuality is not observed in vegetable amaranths. oecy and gynodioecy are identified, of which
On the basis of available morphological and phy- monoecy line might have been followed by grain
togeographical evidences, Mosyakin (2005) sug- amaranths. Gynodioecious pathway is not accept-
gested that dioecy would have evolved in plants able due to lack of any member with intermediate
growing in open habitats such as coastal areas, sexuality. Origin of dioecy from monoecy may
river valleys, disturbed plant communities, des- have evolved through disruptive selection on
erts, semi-desert and prairies. It was accompanied male and female reproductive allocation, fol-
by development of many adaptive morphological lowed by gender specialisation ultimately lead-
traits in some groups (perianth reduction for more ing to unisexual plants.
successful cross-pollination through anemophily, Increased selfing rates resulted in inbreeding
indehiscent utricles for more successful hydro- depression in ancestral cosexual population.
and anemochory, etc.). Dioecious amaranths prob- Such condition facilitates the spread of unisexual
ably independently evolved from monoecious variants that favours the outcrossing. Frequent
ancestor at least twice questioning the monophyly inbreeding in monoecious vegetable amaranths
of the subgen. Acnida (Mosyakin 2005). has resulted in a large number of morphotypes.
Monoecy in the grain amaranths are supposed Relatively greater self-pollination in members of
to have originated from the dominant cosexual sect. Blitopsis and members of sect. Pyxidium
form having exclusively bisexual flower like has resulted in breeding depression that favoured
most of the angiosperms through sterility muta- the spread of outcrossing. Such outcrossing
tion followed by subsequent specialisation in might have resulted in formation of gynomonoe-
flower form and position. Two main evolutionary cious member. Grain amaranths show relatively
pathways for the origin of dioecy though mon- greater outcrossing forming number of variants.
Cultivation of Amaranths
10
10.1 General within 4–5 days after sowing. Amaranths do not

require high amount of nitrogen like maize but
Amaranths are a type of crop that can be grown respond well to fertilisers. There are a number
on any type of soil even in marginal areas in the of obstacles in the breeding of amaranth – like
wild as an escape from cultivation; they need no nonavailability of improved variety, heavy lodg-
special agricultural attention and can tolerate ing and seed shattering and lack of information
environmental stress. Collection and preservation about agrotechniques. Few attributes related with
of seeds are the primary aspect of cultivation. higher seed production were evaluated like mixed
Seeds should be dried below 12 % moisture con- cropping, spacing and population trials, trans-
tent to store for at least 2 years before regenera- plantation versus direct sowing, etc., with posi-
tion. Common amaranth seeds remain dormant tive result. Vegetable amaranths are harvested by
when shed; dormancy is lost after 2–3 months of uprooting or repeated clipping. In multicut
storage in dry condition. Recommended temper- method, the first cutting is done 25–35 days after
ature for germination is 20°–30°, while germina- sowing; subsequent cutting is done with an
tion is poor below 20° temperature. Temperature interval of 7–10 days. Quality and yield get
above 25° is optimum for growth, but growth deteriorated after flowering. For greater nutritional
ceases at temperature below 18°. Lower tempera- value, amaranth leaves are to be harvested at a
ture and short-day length favours flowering. young stage (20 days after sowing). On average
Seeds of grain amaranths deteriorate significantly grain amaranths have longer growth period than
after 1 year of storage. Germination in field weedy species. In grain species, terminal inflo-
depends upon age of seed and depth of sowing. rescences are cut when the plants are still green,
Leafy amaranths are warm-season crop adapted sundried for 6–7 days in thrashing yard and
to hot and humid climate, whereas grain ama- thrashed by biting to collect the grains.
ranths are grown in tropical lowland to an altitude Amaranths are a crop of marginal areas that
of 3500 mt. Amaranths can be grown on various require no special agricultural requirement.
types of well-drained soil. Grain amaranths are Being a pseudocereal they are adapted to fragile
drought-tolerant plant, but leafy amaranths environmental condition and can tolerate difficult
require frequent irrigation to keep soil moist. condition and environmental stress, specially the
Most of the amaranths are day neutral but differ vegetable amaranths. They can grow in the wild,
in their day-length requirement and respond also as an escape from cultivation. Vegetable
differently to changes in photo- and thermoperi- amaranth species are reported to be tolerant to
odism. Seeds are broadcasted on farm but well- adverse environmental effects (Dieleman et al.
pulverised seedbed; germination takes place 1996; Ghorbani et al. 1999). Amaranths are being

154 10 Cultivation of Amaranths
cultivated from prehistoric period till now in germplasm and to increase quality and retain the
almost all environmental conditions ranging from genetic integrity of the germplasm. The viability
the true tropics to semiarid regions and also in of seeds in storage condition is a good indicator
few highest farms in the world. Ecotypes having of seed quality and vigour in many crops. Seeds
the ability to withstand alkaline sandy soils with may be stored for a short period as required
pH as high as 8.5, as well as the acidic clays of for carry-over seeds or for considerably longer
hillside slash-and-burn fields of the tropics, have period as in the case of germplasm accessions
been evolved. Although traditionally amaranths and high-value seed stocks. One can get full ben-
are cultivated within 30° latitude of the equator, it efit from storage system only when the seeds
can be grown in higher latitudes using strains that have high-initial quality. Therefore in germplasm
flower in spite of the longer-day length (photope- preservation, maximum seed quality and vigour
riod) than that of the tropics. Most of the grain are of paramount importance. Amaranth seeds
amaranths have been concentrated in highland start losing genetic integrity when germination
valleys, such as the Sierra Madre, Andes and capacity is below 40 %. Under ambient condition
Himalayas. Compared to the cereals, the yield of storage period should not exceed 3 months for
grain amaranths has been very low, and the dif- best performance of amaranth seeds. Seed preser-
ference continues to widen. Factors responsible vation is the most common method of ex situ
for the low grain production in the Himalayas are maintenance of genetic resources of about 70 %
many. The available varieties of these crops in of the Earth’s plant species. A variety of prob-
this region are the mixture of landraces, the crop lems has been encountered to be associated with
takes a long time to mature, and their growing ex situ conservation strategies, one of which is
season overlaps with that of the cereals. High the problem of genetic changes in storage due to
degree of seed shattering and lodging of plants at ageing and/or field rejuvenation (Roberts 1988).
maturity makes it less productive. Other factors Chromosomal aberrations occur during seed age-
responsible for the poor yields are: (1) crops are ing in storage.
generally grown in lands of marginal productiv- Seed viability equation developed by Ellis and
ity without applying adequate amount of fertilis- Roberts (1980) can be used as a valuable guide-
ers and pesticides, and (2) there is lack of line for estimating deterioration of seeds in stor-
information on scientific cultural practices for age especially when initial viability levels are
this crop to take maximum advantages of its yield high (Mead and Gray 1999). The following equa-
potential. tion reflects seed longevity over a range of stor-
age temperature and moisture content:
V = K i − P / 10 KL − C1 m − C2 t
10.2 S
eed Dormancy and Seed
Viability of Amaranthus This equation relates probit percentage viability
(V) at any time (P), to any combination of mois-
Cultivation of any crop plants and preservation of ture content (m) and temperature (t). Once the
germplasm require a profound knowledge about values of the species’ constants KL, C1 and C2 are
seed dormancy and viability. The primary objec- known, longevity of seeds of that species can be
tive of storing seeds is to maintain the genetic calculated for specific storage condition with
integrity of the preserved accessions as long as certain temperature and seed moisture content.
possible. Deterioration of seeds in storage may The initial viability of individual seed lot is rep-
cause low vigour, reduced number of viable seeds resented by their Ki value which is dependent on
and genetic drift. The precise guideline is the genotype, the prestorage environment and
required to protect or reduce the decline in viabil- their interactions (Ellis and Roberts 1980). The
ity of conserved germplasms. This would help equation enables to predict percent viability
gene-bank personnels to frame the viability test- expected after any given period under different
ing and estimate regeneration intervals on stored combinations of storage temperature and
10.2 Seed Dormancy and Seed Viability of Amaranthus 155
oisture content within medium-term gene-bank

m Wild species and few vegetable accessions of
storage. It has been shown that usefulness of this Amaranthus have seed dormancy. The cultivated
equation regarding storage behaviour persists as and especially white-seeded grain types lack seed
long as initial viability is high. Longevity differ- dormancy and will germinate in 3–4 days at
ence has been found in seeds of various species 21 °C or above (Myers 1996). Kigel (1994)
during storage at comparable temperature and reviewed the seed dormancy; Deno (1993) wrote
moisture content. It has been suggested that the about cool-moist treatment to overcome seed
differences between the seed chemical composi- dormancy of many plant species having the kind
tions affect longevity by influencing the water- of dormancy like amaranths. A month or more of
binding capacity and chemical activity in seeds, moist stratification at approximately 2–5 °C will
hence the rate of deterioration (Walters 1998). be enough to overcome seed dormancy for many
Seed collectors should dry amaranth seeds accessions. Years of dry storage and many chem-
during prolonged collection mission to avoid ical and scarification options can abolish dor-
probable high rate of deterioration. The seeds mancy (Kigel 1994), but the cool-moist treatment
should be dried below 12 % moisture content to is recommended because it is safe and reliable.
store for at least 2 years before regeneration. Germination can be achieved on blotter or sand
Normal storage by the farmers for over a year is (Baskin and Baskin 1998). After stratification
possible for seeds of amaranths without major efficient seed germination will takes place with
viability loss, at temperature ≤20 °C in any 20 °C (night) and 30–35 °C (day) temperature.
combination with ≤12 % moisture content. The Attempts to explain the reasons for low germinat-
species constants obtained can be used to give an ing capacity of amaranth seeds were made based
estimate of anticipated longevity of seeds for the on studies on different species comprising grain
species during storage within short- to medium- amaranths and different vegetable amaranths. To
term storage condition. combat the unevenness in germination, a number
Seeds in the state of rest do not germinate, in of methods for presowing seed processing have
spite of potentially favourable conditions for been employed. The lack of readiness of seeds to
germination, known as seed dormancy. Three germinate leads to unevenness of emergences,
types of seed dormancy are distinguished: and a number of methods for presowing seed pro-
(1) resulting from the external determinants of cessing were developed in order to stimulate
germination (2) determined by physical and them for better germination and emergence of
biochemical factors occurring inside the seeds, seedlings in the shortest period of time possible
but provoked by external factors, and (3) resulting under a wide range of environmental conditions
from internal physiological state of the germ, (Jisha et al. 2013).
independent on the surrounding conditions. Seed From the biological point of view, the seeds of
dormancy of most species can be primary amaranth are fruits and small nuts, in which the
(inborn) or secondary. Primary dormancy is asso- proper seed is surrounded with the hard pericarp.
ciated with natural maturing of seeds. In many Such structure of fruits favours durability of
species such a state is induced by the germination seeds. In their natural environment, plants some-
temperature that is improper for them, or not times have to wait for a long time for suitable
meeting the light conditions creating favourable subsoil moisture to germinate. Even when the
conditions for germination. Inborn dormancy climatic conditions and habitat are similar to
may be of physical and mechanical nature, asso- that required by amaranth, still plant emergences
ciated mainly with the hardness of the seed cover. sometimes become difficult, although seed ger-
Secondary dormancy occurs in nongerminating mination proceeds very fast – only in a few days.
seeds, in which the stage of absolute dormancy Amaranth seedlings are very small and the soil
has already been passed and the seeds are able to moisture in the surface layer determines their
germinate. survival. Almost all the methods known from
agricultural practice were used in experiments to methods of prechilling and KNO3 (ISTA 1985)
increase the germinating capacity of seeds of proved adequate for A. cruentus.
amaranths, viz. treatments affecting an increase Aufhammer et al. (1998) conducted an experi-
in permeability of the seed cover and soaking ment in incubator to find out the effects of several
seeds in water (Musa et al. 2014), in the alcohol factors on the germination of two amaranth culti-
solution of CaCl2 (Colmenarez de Ruiz and vars. The factors considered in the experiment
Bressani 1990) and in the sulphuric acid solution were year of harvest, crop type of the mother
(Soomarin et al. 2010), by cooling (Zharare plant, seed position on the mother plant, stage of
2012) and by the action of alternating magnetic maturity, temperature, light and seed dressing. In
field to seeds (Dziwulska-Hunek and the experiment percentage germination and
Kornarzyński 2009). Most of the above- germination speed were also recorded. Most
mentioned methods as presowing treatment of effects appeared in interaction with cultivars.
amaranth seeds achieved the expected results. Germination percentage was above 80 % when
The most spectacular effects were given by the the temperature was higher than 16 °C. The ger-
use of various treatments to seeds stored for sev- mination speed showed close relationship with
eral years, overwintered in soil and exposed to temperature, and speed decreased with decrease
the salt stress, for those which have naturally in temperature. Light or even a short illumination
weakened germination capacity (Moscova 2012). inhibited germination and slowed it down at tem-
In laboratory conditions, the germination energy peratures below 25 °C, while presoaking of seeds
and capacity of amaranth seeds of the cultivar accelerated germination. Seed stored for more
Rawa much more depend on the germination than 1 year showed decreased germination per-
temperature than on the kind of preparation in centage. An early harvest of homogenous and
which seeds are soaked. Germination capacity at dense amaranth crops is recommended for ama-
25 °C can be regarded as satisfactory, irrespective ranth seed production. More research regarding
of the method for seed material treatment. presowing treatment of amaranth seeds is
Effective stimulation of amaranth seed germina- required.
tion at 15 °C is induced by soaking them for 8 h Seeds of grain amaranths are considered
in 0.03 % water solution of Pol-Gibrescol, as well among the most sensitive seeds susceptible to
as in a mixture of 0.03 % Pol-Gibrescol and 2 % significant deterioration after storage of 1 year.
Betokson 025 Super SL. Seeds are required to be preserved in safe storage
Although ISTA (1985) recommended germi- since they are harvested in the preceding season
nation at either 20 °C or 20/30 °C for Amaranthus and generally used for sowing in the next season
sp., Grubben (1976) suggested that 20 °C was often after a time gap of 6 months or more. Seeds
approaching a germination temperature bound- gradually lose their vigour and germinability and
ary, and germination was poor at temperatures ultimately become less viable during the ageing
lower than 20 °C, particularly in the presence of process in storage (Maity et al. 2000). Losses in
white light. Germination of Amaranthus seed is seed quality also occur during field weathering,
photo inhibited. In the dark, germination is inde- harvesting and storage. Several intrinsic and
pendent of temperature, but in the light, germina- extrinsic factors influence the viability of seeds
tion at constant temperature increases with during storage. Among intrinsic and extrinsic
increasing temperature. In A. cruentus germina- factors, seed moisture content, relative humidity,
tion was usually greater at 20/30 °C than at temperature of storage, pests and diseases and
20 °C. Seed dormancy in Amaranthus spp. is a oxygen availability are more important. Storage
type of relative secondary dormancy, a photo dor- of seeds wrapped in polyethylene and aluminium
mancy (Kendrick and Frankland 1969) induced foil was found effective in preventing moisture
by prolonged exposure of seeds to white and far- uptake and maintaining seed viability, while stor-
red light. It can be broken by exposure to red age of seeds in paper and cloth-made containers
light, but the more conventional seed testing were found least effective (Wilson and McDonald
10.2 Seed Dormancy and Seed Viability of Amaranthus 157
1992). Seed deterioration is an inexorable and an ent plant have a definite effect on the level of
irreversible event. One of the symptoms of dormancy. Fresh seed collected from distinct
seed deterioration is membrane deterioration populations of common amaranth may show dif-
(Copeland and McDonald 1995). Seed deteriora- ference in their germination behaviour, but such
tion alters the semi-permeability properties of the differences are reduced during dry storage. The
membrane and membrane integrity (Berjack and tough seed coat does not rupture easily and,
Villiers 1972). Electrical conductivity of seed while water uptake is not prevented, seed germi-
leachate increased gradually over period of nation is prohibited by it. In laboratory studies,
storage. With increase in storage period, the the minimum temperature for germination was
germination capacity, vigour and protein content 10 °C (Wiese and Binning 1987; Ghorbani et al.
decrease, while the electrical conductivity 1999), while maximum germination occurred at
increases irrespective of treatments and storage 35–40 °C. The ripe seeds soon after harvest
containers. This may be probably due to increased showed some germination when incubated at
moisture content of the seeds. temperature of 40 °C. Nitrogen is suggested to
Weed species which form persistent seed promote germination, but the effect of light on
banks are of concern for future weed manage- germination is yet to be documented
ment (Egley and Chandler 1983). In the absence conclusively.
of viable seed production, a decline in amaranth Seed of common amaranth showed 50 % less
seed bank might be due to seed mortality through germination at 15 °C than at 25 °C (Chakraborty
physiological age and herbivory or microbial 1977). In laboratory experiment, when dry-stored
decay (Egley and Williams 1990; Buhler and seeds are sown on moist paper or soil in the light
Hartzler 2001). Persistent seed bank of a few at a constant temperature of 18–20 °C, 70 % ger-
Amaranthus species has been investigated earlier mination was achieved (Cross 1930–1933).
(Steckel et al. 2007). Palmer amaranth (A. palm- Germination was nearly 90 % when the seeds
eri S. Watson) is known as prolific seed producer, were incubated at alternating temperatures of
a single female plant may produce up to 600,000 0/30 °C or 8/20/30 °C. In other experiments, little
seeds (Keeley et al. 1987). Investigation on seed difference was observed between germinations
persistence of palmer amaranth is lacking. It under diffuse light or light filtered through a leaf
shows seasonal seed dormancy and an extended canopy to reduce the red light ratio compared
emergence period, resulting in a season-long with the far-red light (Taylorson and Borthwick
interference and severe yield reductions in sev- 1969).
eral crop plants. Amaranthus species have low The optimum depth of seedling emergence is
persistence in soil seed banks (Steckel et al. a crucial factor that varies from 0.6 to 1.3 cm
2007). The redroot pigweed (A. graecizans) depending on temperature (Mohler 1993). In the
exhibits a sharp decline in the viable seed bank laboratory condition best seed germination was
within 2 years of burial, with no viable seeds observed between 5 and 30 mm deep in soil
remaining in the upper 0–15 cm depth of soil (Ghorbani et al. 1999). Germination was found
after the third year (Egley and Williams 1990). better in clay soil than in sandy soil. Below the
Several studies on natural and artificial seed depth of 40 mm and on the soil surface, seed
banks reported that the rapid decline of germination in both the cases was much less.
Amaranthus seed bank within a year of burial However, when seeds are sown in trays of field
might be due to physiological death of seeds, soil subjected to different cultural treatments, the
fatal germination, seed herbivory or microbial highest germination was achieved from seeds left
seed deterioration in the soil (Cardina et al. 1996; on the soil surface (Chepil 1946). Seedling emer-
Kremer 1993). gence decreased with increasing burial depth,
Generally common amaranth seed remain and cultivation increases seedling emergence by
dormant when shed. The time of emergence and bringing seeds into the upper soil layers, but fatal
the growing conditions experienced by the par- germination did not occur and a greater number
of seeds simply remained nongerminated. light after a period of darkness for germination.
Seedling emergence was less in untilled soil than This trait helps them to adapt in disturbed or
in tilled soil even when seeds were maintained at overturned soil. The weedy amaranths are often
the same depth (Mohler and Galford 1997). Few called ‘pigweed’ because they would germinate
of the seeds sown in a 7.5 cm layer of soil in open in hordes in an area after pigs had passed and
cylinders in the field and stirred periodically turned the soil, exposing their seeds to the light
seedlings emerged soon after sowing in autumn so they could germinate. Amaranth seeds can be
(Roberts 1986). In the following year, the seed- stored safely for up to 3 years at a temperature
lings emerged from May to August. Germination below 8 °C and at 10 % relative humidity in a
appeared to require a high temperature. A gradu- tightly closed moisture-resistant container
ally reducing number of seedlings emerged in (Hartmann et al. 2011). Ideal containers are air-
subsequent years, but some viable seeds still tight such as a sealed glass jars, metal cans or foil
remained after 5 years. envelopes as they maintain seed water content
Common amaranth seeds spread in moist soil, best. Seed in containers should be stored in a
persist and can remain viable but dormant for cool, shady and dry place to extend seed shelf life
over 30 years (Crocker 1916). However, other (Hartmann et al. 2011).
investigations reported just 1 % seed viability Seeds from different crops can be stored for
after 5.5 years in soil (Egley and Chandler 1983). different periods of time after harvesting. Seed
In an experiment to study the relationship viability at the end of the storage period is deter-
between burial depth and germination percent- mined by the initial viability at harvest and the
age, it was found that seed buried at 20, 56 and rate of deterioration during storage. Deterioration
107 cm showed 9, 11 and 18 % germination, differs with seed species as well as the storage
respectively, after 1 year; 11, 36 and 48 % after conditions including temperature and relative
10 years; and none after 16 years (Toole 1946; humidity (Muyonga et al. 2008). Seeds can be
Goss 1924). After a dry storage at low tempera- classified as either recalcitrant or orthodox based
ture for 30 months, seeds retained full viability, on their genetic potential to tolerate storage.
but after burial in field soil for the same period, Recalcitrant seed are those that do not tolerate
viability was less than 13 % (Egley and Chandler seed moisture below 25 % after seed maturation,
1978). Common amaranth seeds sown in the field while orthodox seeds can tolerate drying from
and followed by cultivation of wheat in winter 10 % down to 4 % moisture content after seed
and barley in spring over a 5-year period showed development, and these differ in the length of
an annual decline in germination of around 40 % time they can tolerate storage (Barker and
(Barralis et al. 1988). Emerging seedlings repre- Duarte 1998). According to Hartmann et al.
sented only 8 % of the seed bank. The viability of (2011), orthodox seeds can be further divided
seeds recovered from 10 cm deep in soil declined into medium-lived and long-lived categories.
from 98 % to 90 % in a 12-month time period, Medium-lived seeds can remain viable for peri-
while the viability of seeds recovered from the ods of 2–5 years provided that seeds are stored at
soil surface declined from 93 % to 62 % over the relative low humidity and temperature. Seeds of
same time period (Omami et al. 1999). Soil solar- most vegetables, flowers and grain crops belong
isation is the main factor for speedy decline in to this group. It is important for seed storage to be
germinability and viability, and common ama- designed in such a way that it should not create
ranth seeds are very sensitive to soil solarisation. conditions that will negatively affect seed and/or
When the seeds were sown in pots of moist soil seedling vigour. During seed deterioration the
and treated with worm air for 6 h, germination seed first loses vigour or the ability to germinate
was reduced by 30 % at 47 °C, 60 % at 52 °C and when environmental conditions are not favourable.
80 % at 54 °C (Laude 1957). Seed deterioration during storage is stimulated
If germination is slow, soil surface needs to be by high respiration and other metabolic rates
lightly stirred; amaranth seeds require some sun- which injure the embryo (Hartmann et al. 2011).
10.3 Agroclimatic Condition for Cultivation of Amaranths 159
Moisture content in seeds is the most important do not germinate even if they are subjected to
factor in seed longevity and, therefore, is impor- favourable conditions that are normally beneficial
tant to consider during storage (Baskin and for germination (Mayer and Mayber 1995).
Baskin 1998). For example, seed having ortho- Essentially, dormancy is an adaptation mecha-
dox characteristics can be best stored at a non- nism that prevents seeds from germinating after it
fluctuating low moisture level as they can tolerate has been dispersed by mother plants and that only
low moisture content. Seed moisture content of permits germination when environmental condi-
about 4–6 % is suitable for prolonged storage of tions are favourable (Baskin and Baskin 1998).
seed from many vegetable species. However, Dormancy in amaranth seed is reported to be
many storage problems may arise when seed high at the time seeds are detached from mother
moisture content is elevated during storage plants, but decline as seed water content decreases
(Baskin and Baskin 1998): (1) at about 8–10 % (Baskin and Baskin 1998). Generally, dormancy
moisture content, several insects are active and is at its peak within 2–3 months after seed
can reproduce; (2) above 12 % seed moisture harvesting. In amaranth seed dormancy after ripen-
content, fungi are active and can multiply to ing is associated with naturally occurring com-
produce spores; and (3) at the higher seed mois- pounds present in the seed at maturity, and oven
ture content levels, respiration, germination and drying or naturally air-drying can help to reduce
disease activity are stimulated leading to reduced amaranth seed dormancy (Hartmann et al. 2011).
seed viability (Barker and Duarte 1998). On the
other hand, too low water content in some seeds
can have a reducing effect on seed viability and 10.3 Agroclimatic Condition
germination rate (Abdullah et al. 2011). For this for Cultivation of Amaranths
reason, hydration is necessary for seeds stored at
a humidity atmosphere below 2 %, to avoid seed 10.3.1 Climate
injury, as this can influence the moisture content
of the stored seed. Conversely, in the case of Leafy amaranths or vegetable amaranths are a
some species, dry climate increases seed longevity, warm-season crop adapted to hot-humid climatic
while high relative humidity (RH) results in conditions. It is grown throughout the year in
shorter seed life (Barker and Duarte 1998). tropics and in autumn, spring and summer sea-
Amaranth seeds can be preserved safely for up to sons in temperate regions. Amaranth can tolerate
3 years at a temperature below 8 °C and at 10 % full sun, drought conditions and high tempera-
RH in a tightly closed moisture-resistant con- tures. Grain amaranths are grown in wide geo-
tainer (Hartmann et al. 2011). Ideal containers graphic areas ranging from tropical lowlands to
are airtight such as sealed glass jars, metal cans 3500 m in the Himalayas. Altitudes above
or foil envelopes as they best maintain seed water 1000 m in the tropics are considered best for
content. Seed in containers should be stored in a grain amaranth cultivation though they are toler-
cool, shady and dry place to extend seed shelf life ant to drought condition and low soil fertility. It
(Hartmann et al. 2011). performs much better under conditions that are
As mentioned earlier, germination of all seed considered ideal for maize (corn). It can be inter-
is affected by the viability of the seeds, seed cropped with maize, beans, peppers or squash. In
dormancy and adequate environmental factors. If the Andes region, it is often intercropped with
one of the three aspects is not sufficiently consid- quinoa (Chenopodium quinoa) and other pseu-
ered, germination can be severely delayed or docereals by the local farmers. For grain ama-
inhibited, leading to secondary dormancy (Tucker ranth cultivation, altitude is not a severe limiting
1986). Seed viability represents the ability of factor. Amaranths can grow satisfactorily from
nondormant seeds to germinate, and viability sea level to above 3000 m; the only exception is
testing is essential in determining seed quality. A. caudatus which is known to grow at an alti-
Seed dormancy is the condition whereby seeds tude of 3000 m in the Andean region and
Himalayas. Some of the grain species of A. tricolor has demonstrated (Foy and Campbell
Amaranthus are s ensitive to day length. Some 1984) tolerance for soil with high aluminium
accessions of A. hypochondriacus will not set levels. Cultivation practices differ according to
flower in summer; however they do mature in the the methods of harvest, duration, growth pattern
greenhouse during short-day situation in winter. of variety, etc. Land is prepared to a fine tilth by
Some strains of A. cruentus remain vegetative thorough ploughing and harrowing. Well-
for a long time in its equatorial house. However, decomposed and powdered organic matter at
it starts to set seed very early when introduced 20–25 t/ha is incorporated with the soil at the
into the long-day conditions. A. caudatus thrives time of final ploughing. The poor amaranth
well in high hills and is familiar as a short-day stands in field can be attributed to various reasons
species. It generally flowers and sets seed only like soil crusting, low soil moisture, poor seed-to-
when day length is less than 8 h. Amaranth pop- soil contact, uneven planting depth and wind
ulation are easily miniaturised, and their flower- erosion in some areas. Proper addressing of such
ing cycle can be accelerated by controlling the primary reasons will result in a good stand of
environment (Brenner and Widrlechner 1998). amaranths (Sooby et al. 1998).
Amaranthus production guide (Sooby et al.
1998) is a useful reference for field management
of amaranth. 10.3.3 Temperature
Temperature is an important parameter for seed

10.3.2 Soil germination, growth, emergence of inflores-
cence, etc. Grain amaranths grow best when the
Amaranths can be grown on various types of soil daily high temperature is at least 21 °C. Various
including marginal soil. Amaranth comes up well accessions of grain amaranths have shown opti-
in well-drained loamy soil rich in organic matter. mal germination at temperature range of
The ideal pH is 5.5–7.5, but there are types which 16–35 °C. The speed of emergence is encouraged
can come up in soils with pH as high as 10.0. It at the higher end of the temperature range. The
can be grown on marginal areas with little soil grain species A. hypochondriacus and A. cruen-
management. It is evident from the field observa- tus are not frost hardy and can withstand high
tions that amaranth can grow well on soil con- temperature. Growth of the plant stops at about
taining variable amount of soil nutrients. Initial 8 °C and below 4 °C the plants get injured.
studies at Rodale Research Centre (RRC) in However, A. caudatus being native to Andes and
Pennsylvania showed that young grain amaranth high Himalayas is more resistant to chilling than
plants grow taller with fertiliser but with little the other two grain species. The optimum germi-
improvement in grain yield. On the other hand, nation temperature for amaranth has been
vegetable amaranth needs high soil fertility, par- reported to be between 20 and 30 °C (ISTA
ticularly potassium and nitrogen. Grain ama- 2010). The best temperature for germination and
ranths prefer well-drained cultivation field with early seedling growth ranges between 25–30 °C
neutral or basic soils (pH values above 6). for A. cruentus and 25–35 °C for A. hybridus,
However, this aspect has not been studied care- respectively. Amaranth seeds require soil-
fully considering the wealth of amaranth germ- temperature range of 18–25 °C for germination
plasm variability that exists. It is most likely that and an air temperature above 25 °C for optimum
types that can tolerate acidic condition can be growth. The growth ceases at temperatures below
identified for cultivation in tropical lowlands 18 °C. Lower temperatures associated with
where acid soils are common. The genus is not shorter-day length will induce flowering with a
known for high salt tolerance, but they have the subsequent reduction in leaf yield. As the crop
ability to withstand mild salinity and alkalinity as grows during summer with the onset of the rains,
apparent in some species of amaranth. Moreover, frost damage should not be a problem. However,
10.4 Cultivation Practices 161
harvesting of the crop may be effected by frost. of Amaranthus caudatus, such as the ornamental
As amaranth is an annual crop, it does not mature ‘love-lies-bleeding’, will set seed when the day
completely in areas having a short growing length is longer (El-Sharkawy et al. 1968).
season. Frost is necessary to terminate the
crop’s growth.
10.4 Cultivation Practices
10.3.4 Rainfall 10.4.1 Sowing of Seeds
Grain amaranths are drought-tolerant crops, but The field must be well levelled and two to three
vegetable amaranths require frequent irrigation to ploughings are sufficient for sowing of amaranth
keep soil moist. Frequency of irrigation depends seed. To initiate amaranth cultivation, a well-
on soil. Seeds of amaranths require well-moistened worked, firm and moist seedbed is required. It is
soil to germinate and establish roots, but once necessary to firm the soil over the seed to make
seedlings are established, grain amaranths perform good contact between the seed and the soil. A
well with limited water; in fact they grow best loose (but firmly packed), friable soil is preferred
under dry, warm conditions. Vegetable amaranths, in seedbed. Amaranth seeds are very small in
on the other hand, require moisture throughout the size, and the weight of 1000 seeds is varying
growing season. Grain amaranths have been grown from 0.7 to 0.9 grammes. It has been found that
in dry agricultural areas that receive as little as the largest amaranth seeds are only 1/16 in., and
200 mm of annual precipitation, while vegetable some varieties have seed as small as 1/23 in. in
amaranths are routinely grown in areas receiving diameter. As per the recommendation, 1.2–3.5 kg
3000 mm of annual rainfall. seed/ha is to be planted to an average depth of
1.3 cm (Webb et al. 1987). According to the
authors, planting depth needs to be monitored
10.3.5 Day Length carefully as the deeper range might delay and
decrease seedling emergence. On the other hand,
Most of the vegetable amaranths are day neutral shallower planting depth is also believed to
in habit but differ in their day-length require- decrease seed-soil contact, exposes the seeds to
ments and respond differently to changes in pests and increases the risk of being washed away
photo- and thermoperiodism. Grain types, A. by water (Stallknecht and Schulz-Schaeffer
caudatus, A. cruentus and A. edulis, are short-day 1993). Further, amaranth seed requires a firm
species, while A. hypochondriacus is day neutral. moist seedbed with a soil temperature above
Many of the amaranths are sensitive to day 15 °C to ensure proper seedling emergence and
length. For example, accessions of Amaranthus good plant establishment. Consequently, drying
hypochondriacus from the south of Mexico will of the top soil during the germination process can
not flower in the summer in Pennsylvania. reduce seed germination and subsequently seed-
However they will flower in the greenhouse ling establishment. Therefore, amaranth seed
during the short-day conditions in winter. The germination depends mostly on moisture and
reverse happens with Amaranthus cruentus from temperature, with an optimal temperature range
Nigeria. It remains vegetative for a long period in between 16 and 35 °C (Muthomi and Musyimi
its equatorial home. However, it sets seed very 2009). As is the case with all crops, amaranth
early when placed under the long-day conditions also experiences harsh abiotic and biotic stress
in Pennsylvania and can be used to breed for conditions when planted in the field. Water-
early-maturing traits. Amaranthus caudatus, on deficit stress and other environmental factors,
the other hand, is known to be a short-day spe- like stress of unfavourable temperature, have
cies. It usually flowers and sets seed only when been shown to reduce seed germination and
day length is less than 8 h. However, some accessions seedling development.
Seeds may be planted in a nursery bed for sub- The wider rows gave the highest yields. Amaranth
sequent transplanting in cultivation field or sown seeds, being small in size, are mixed with fine
directly in the field. Transplanting ensures a very sand and sown uniformly by broadcasting. The
efficient use of seeds and the growing area to be seeds are covered either by raking up soil and by
weed free just before the seedlings are trans- covering with a thin layer of sand or soil. This is
planted. If direct seeding is practised, sowing in followed by a light irrigation. Soil is kept moist
rows is recommended. Seed sown at a depth of by frequent irrigation. Grown-up seedlings are
1.3 cm (0.5 in) or less in soil with temperature of selectively pulled out at 30 days after sowing and
15 °C will establish a good plant stand. Seed will marketed in small bundles along with roots.
germinate within 3–4 days with soil temperature
of 20 °C (68 °F). Before sowing, application of
50 quintals of farmyard manure per hectare has 10.4.2 Seedling Growth
been found to be good to enhance yield. Plant and Interculture
density, i.e. spacing between plants, has a great
impact on yield and harvesting specially in the Vegetable amaranth species are reported to be
case of vegetable amaranths. One common prac- tolerant to adverse environmental effects
tice is to grow plants at a spacing of 5–10 cm (Dieleman et al. 1996; Ghorbani et al. 1999).
(2–4 in) and harvest by uprooting when the plants They have been growing wild in arid and semi-
are 5–7 weeks old. Another common method is arid regions, which mean that they could be more
to sow seed less densely at a spacing of 15–30 cm tolerant to low-water and high-temperature con-
(6–12 in) and harvest by cutting the stem tips and ditions (Hurro and Cees 1991; Modi 2007). Many
tender leaves periodically starting from 4 to African communities believe that because
6 weeks when the plants are about 15 cm tall. The amaranth is found in the wild, there is no need to
crop is generally sown in the first or second week cultivate these plants (van Rensburg et al. 2007).
of June just after the first monsoon shower. Seedling growth is highly dependent on both
However, amaranth can be propagated from seed moisture and temperature, with A. hybridus being
in the early summer (Muyonga et al. 2008). more tolerant to high-temperature and low-
Traditionally, the seeds are broadcast, but better moisture conditions than A. cruentus.
crop stand is achieved if seeding is done in rows. Seed germinates within 4–5 days after sowing
The depth of sowing should be less than 2 cm in and needs maximum care till it attains a height of
view of very small grain size, with 50 cm spacing about 25–30 cm. In fact this is the most critical
between rows and between plants. On a large stage for obtaining maximum yield potential of
scale, grain amaranth is directly seeded into the the crop. During this phase, it must be properly
field at a seed rate of 1–1/2 kg to 2 kg/ha for good spaced, made free from weeds and must receive
grain yield. Amaranth is harvested by pulling out adequate moisture. One more weeding is neces-
and by frequent clippings (multicut). sary after 30 days of sowing. At the seedling
Field studies have shown that amaranth yields stage, one spray of some fungicides to check the
remain constant across a range of 0.3–4.5 kg/ha attack of damping off is necessary. Once the
(0.25–4 lbs/a). There are approximately 1000– stand is established, then its maintenance is rela-
3000 amaranth seeds per gramme (1,000,000– tively easy. The land, after thorough ploughing
2,500,000 /kg) (Sooby et al. 1998). Plant spacing and levelling, is made into shallow trenches/
recommended for grain amaranth varies widely. basins of 50–60 cm width and convenient length.
One recommendation is to maintain a spacing of Well-decomposed farmyard manure is applied in
23 cm (9 in) between plants and 75 cm (30 in) trenches and thoroughly incorporated in soil by
between rows. This results into a planting density digging. Seedlings (20–25 days old) already
of approximately 38,000 plants per hectare raised in nursery are transplanted in trenches at
(15,400 per acre). If harvesting is to be done 20–25 × 10–15 cm spacing. Seed requirement for
manually, the less dense spacings are advisable. transplanted crop is only 500 g/ha. Amaranth is a
short-duration and shallow-rooted crop. A light these situations, satisfactory stands of amaranth
hoeing is needed to prevent soil crust formation were established applying readily soluble
after irrigation and to keep soil loose. Field also phosphorus fertiliser directly below the seeds.
should be kept weed free, especially during ini- Under hilly condition in India, NPK at the rate of
tial stages. 20:30:20 kg/ha is sufficient to meet the full
Direct seeding though requires much less requirements of the crop.
labour, but it invites a greater risk of poor stand Amaranths are heavy feeder and high-yielding
due to diseases and predators of young seedlings crop; 20–25 tonnes/ha of FYM (farm yard
and poor competition with existing weeds in the manure) and 50:25:20 kg NPK/ha are recom-
crucial initial couple of weeks. If direct seeding mended as basal dose. Under pulling-out method,
is practised, sowing should be done in rows to 20 kg nitrogen should be top-dressed twice
facilitate cultivation. Because of the shallow during subsequent pulling-out of seedlings. For
depth of plantation, special care must be taken to clipping varieties, a still higher dose of 75:25:25 is
prevent the soil from drying out or soil solarisa- advisable. Nitrogen should be applied after every
tion until plants are established. Transplanting or clipping or cutting. Foliar spray of 1 % urea or
thinning is vital, and it may be done within about diluted cow urine at every harvest is good for
2 weeks after sowing when plants are 5–10 cm promoting further growth and for high yield.
tall (2–4 in). However, any delay in transplanting
has an adverse effect on yield.
10.4.4 Obstacles in Productivity
in Grain Amaranths
10.4.3 Manuring
The major productivity constraints of the crop
Amaranth does not require a high amount of identified are:
nitrogen fertiliser like maize, but responds well
to fertilisation. A leguminous cover crop tilled 1. Nonavailability of improved variety for culti-
under prior to seeding is enough to provide suf- vation till recently
ficient nitrogen for amaranth. Animal manure or 2. The lack of information on agrotechniques for
chemical fertiliser at a rate of 135 kg per hectare high seed production
(135 lbs/a) is also sufficient. The requirement of 3. Heavy lodging and seed shattering
chemical fertilisers is less when the cultivation is 4. Little or no use of inorganic fertilisers
preceded by a leguminous crop such as beans or 5. Little or non-utilisation of seed for various
soybeans. Singh and Whitehead (1993) studied domestic and agroindustrial products by mass
the growth responses to three different pH levels consumers
6.4, 5.3 and 4.7. Results indicated that pH 6.4 is
6. The lack of public (consumer) awareness
the best for growth and growth decreases with the about the organoleptic taste and quality of both
increase in soil acidity. Stallknecht and Schulz- grains and greens for production of various
Schaeffer (1993) recommended soils with pH domestic, agroindustrial delicious products.
above 6.0 suitable for growth. Fertiliser like NPK
showed no significant effect on yield, but To identify and standardise the attributes for
increased the seed protein content in the two high seed production in grain amaranths, agro-
lowest-yielding ecotypes, while effects on seed nomic trials concerning five attributes were con-
fat content were inconsistent. Amaranth is not ducted at NBPGR, Regional Station, Shimla,
well adapted to soils markedly deficient in avail- with newly released variety Annapurna during
able phosphorus; amaranth seedlings have repeat- 1985 and 1986. The attributes were (1) spacing
edly failed to establish. In the same trials, cereals and population trial, (2) sowing date trial, (3) mixed
such as corn, sorghum, wheat, triticale and even cropping with French bean, (4) transplanting vs.
small-seeded millets established satisfactorily. In direct sowing and (5) leaf picking vs. nonpicking.
Plant-to-plant spacing of 20 cm has been observed stage is recommended for taking maximum
to be the optimum spacing for obtaining highest advantage of the crop in the hills.
grain yield of 34 q/ha. To standardise the opti-
mum time for sowing of seed in the hills and for
obtaining highest seed yield, a randomised block- 10.4.5 Harvesting
design trial with four sowing dates with an inter-
val of 15 days was conducted. The highest grain Vegetable amaranths are harvested by uprooting
yield of 16.1 q/ha was obtained when sowing was tender plant or by repeated clipping or cutting.
done on first date of sowing (June 2) followed by Harvesting by repeated clippings, with an interval
the second date of sowing (June 17), and the last of 2 or 3 weeks is common through the end of the
was recorded on the last date of sowing (July 17). season (usually the short-day length period of the
The reasons for low yield levels in this trials were year). Frequent clipping is helpful to increase
due to heavy lodging and seed shattering. The both the yield and quality of leaves. At the end of
delayed sowing beyond June highly reduced the vegetative growth, flowering begins, and subse-
plant height, inflorescence length and glomerule quently harvest becomes inferior both in quality
number per plant. It also induced tenderness to and quantity. Vegetable amaranth is harvested
the plants. Early sowing caused a big stem borer early in the morning by pulling out or by clipping.
problem as compared to late-sown plants which In the first method, grown-up plants are pulled out
were completely free from this disease. Highest at 30, 45 and 55 days after sowing, along with
yield of 21 q/ha was obtained under mixed crop- roots, washed and sent to market in small bundles.
ping when French bean (viny type) was grown In multicut method, first clipping or cutting is
between two rows of amaranth at 25 cm apart. done 25–35 days after sowing. Subsequent cut-
Mixed cropping with French bean not only pro- tings are made at 7–10-day intervals. Premature
vided the nitrogenous fertiliser to the amaranth flowering or bolting is a serious problem in culti-
plants but also held them by twining and prevent- vation of amaranth. Bolting is usually associated
ing lodging which is a serious problem in the with planting of short-day varieties during
hills. Highest grain yield of 24.3 q/ha was November to December, deficiency of nitrogen,
obtained in the case of direct sowing, and there extreme high temperature and poor soil erosion.
was a reduction of 7 q/ha under transplanting. Practices like raising of crop at ideal time depend-
Early maturity by a week time and reduction in ing on locality, frequent application of nitrogen
plant height, inflorescence length and leaf size fertilisers and manures and keeping soil loose by
were observed in the case of transplanted plants. light hoeing may be some of the preventive mea-
The farmers in the Himalayas generally pluck the sures. Prolonged flowering is a serious problem
leaves of grain amaranth at vegetative stage when in cultivation of amaranth. Quality and yield
the plant is tender for vegetable use and leave the are deteriorated after flowering. Flowering of
rest for grain yield. There was a reduction of Amaranthus species usually starts 4–8 weeks after
2.9 q/ha grain yield in the case of leaf-plucked showing (Grubben and Denton 2004). However
plant than the normal ones, but there was 20 q/ha the growth and development pattern are highly
additional foliage yield for vegetable use. In the variable between species to species and cultivars
case of leaf picking, plant height is reduced and to cultivars, depending on photoperiodism, alti-
number of branches at the top and number of tude and cultivation practices (Wu et al. 2000).
leaves increased. The size of inflorescence Wild leafy vegetables are harvested from crop
increased, but the number of glomerules fields by rural people at different stages of plant
decreased. The leaf-plucked plants became growth. There may be a specific or preferred
stunted and stout and did not lodge which is the plant developmental stage when flavour and pal-
most desirable character. Late maturity by a week atability are most favourable for human con-
was observed in the case of leaf picking than the sumption. In the case of leafy vegetable, it is not
normal ones. Thus leaf picking at the vegetative likely that flavour and palatability are influenced
by environmental condition. Data on changes in eases have been identified as the serious diseases of
nutritional quality of leaf in response to plant age grain amaranth in India. Amaranths are generally
and environmental condition is scanty. Modi affected with some fungal diseases, specially
(2007) made an attempt to study the effect of damping-off disease of seedlings caused by
growth temperature on amaranth leaf yield and Pythium, Rhizoctonia and Aphanomyces spp. and
nutritional value at different stages of plant cankers caused by either Phoma or Rhizoctonia.
growth involving five species, viz. A. hybridus, Various root and stem rots can occur later in the
A. hybridus var. cruentus, A. hypochondriacus, season when the soils are wet, which contribute to
A. tricolor and A. thunbergii in South Africa. lodging problem. Alternaria leaf spot is the most
Vegetable amaranths are the most widely occur- serious foliar disease in amaranth. Amaranth is
ring leafy vegetable in South Africa and Africa in generally considered tolerant to nematodes and
general (Modi 2007). Nutritional quality of often has been recommended as a rotation crop to
Amaranthus leaves is significantly influenced by reduce nematode populations for subsequent
growth temperature and developmental stages at crops. The presence of root-knot nematodes in
which harvesting has been done. It was found amaranth roots has been reported. It is important
that cool environmental condition favours the to know whether or not amaranth can be used to
increase in total protein and amino acid content control nematodes and/or whether it can be culti-
in leaves. However the amount of mineral elevated where nematodes are a problem. A lot of
ment like calcium and iron increased in leaves in insects are known to feed on amaranth leaves that
response to increase in growth temperature. It may account for severe and sustaining yield loss.
was recommended that for greater nutritional A few insects may cause substantial damage.
value, amaranth leaves should be grown under Amaranth may succumb to caterpillars, web-
warm condition and leaves should be harvested at worms, blister beetles, lygus bugs and stem bor-
young stage (20 days after sowing). Warm condi- ers. The lygus bug, coffee bug or tarnished plant
tion was found to be associated with high yield bug (Lygus spp.) is a sucking insect that attacks
and improved germination capacity. flowers and seeds and causes severe damage both
On an average the three grain types have lon- by preventing flowers from producing seeds and
ger growth period than weedy species. Amaranth also by reducing seed weight. Solutions of
grains mature much earlier and the plant dries up pyrethrum or synthetic pyrethrins are helpful to
quite late. If the heads are allowed to remain till control lygus. Other insects that can injure the
the plant dry up, heavy shattering of grains is developing amaranth include fall armyworm
noticed which leads to heavy grain loss. The ter- (Spodoptera frugiperda), corn earworm (Heliothis
minal inflorescences or heads are cut when the zea) and the cowpea aphid (Aphis craccivora).
plant is still somewhat green and start weathering The amaranth weevil (Conotrachelus seniculus)
and kept for sundrying for 6–7 days in the threshing can damage roots, resulting in lodging or other
yard. Threshing is done by beating. The produce root diseases. The potato flea beetle (Epitrix cuc-
is threshed and winnowed like other cereals. umeris) can damage seedlings, and the beet leaf-
Unusually the harvesting is done early in the hopper (Circulifer tenellus) can transmit curly top
morning when the plants are somewhat wet due virus, but this has been seen only in areas nearing
to night dew to avoid grain shattering in cut large areas of sugar beet production.
heads. When the grains mature, the inflorescence Hymenia recurvalis and other caterpillars are
heads are cut dried under shade. serious pests of amaranth like Cletus sp., Asparia
sp. and Lygus lineolarus bug (cabbage looper),
Trichoplusiani (European cornborer), Ostrinia
10.4.6 Pests and Diseases nubilalis (corn earworm), Heliothis zea (cowpea
aphid), Aphis craccivora (striped blister beetle),
In general, soil fungus, damping-off, leaf blight, Epicauta vittata, a weevil, spinach flea beetle,
white rust and mycoplasma and virus-related dis- Disonycha xanthomelas, etc. Lygus bugs, more
specifically the tarnished plant bug, are recently damaging Amaranthus hybridus plant in
considered to be the most important insect pest of Mississippi, USA. The other species damaging the
grain amaranth. These insect feedings result crop are C. truncatum and C. capsici. Choanephora
in localised wilting and tissue necrosis followed cucurbitarum, causing wet rot, is the most trouble-
by abscission of fruits, morphologically deformed some disease of amaranth in Africa. Albugo bliti,
fruit and seed and altered vegetative growth. causing white rust, is a serious disease in Southeast
Serious damage of amaranths has been caused by Asia. Reddy et al. (1980) reported Xanthomonas
spider mites and stem weevil. Stem borer (Lixus amaranthicoia as a causal organism of bacterial
truncatucus) is another problem in Africa and Asia leaf spot disease of amaranths in India. Sharma
in early-sown plants which causes high degree of et al. (1981) and Naseema et al. (1983) reported
lodging in plants. A considerable damage is done that Aspergillus flavus, A. niger and Rhizopus
by the leaf rotters during rainy seasons. Seedbeds stolonifer were externally as well as internally
should be protected against ants and termites. The seed borne in most of the seeds of Amaranthus
other pests damaging the crops are Hypolixus gangeticus and A. caudatus and are the major
nubilaus (Egypt), Rhachi creagra (Costa Rica), storage fungi of amaranth crop in India. Sammons
Chrotogonus (Pakistan), Hyphurus (India), and Barnett (1987) reported about tobacco
Geocoris (California), Thysanoptera (Hawaii), ringspot virus damaging Amaranthus hybridus in
Diabnotica barberi and Spodoptera exigua. The California, USA. A severe mosaic disease caused
fungal population was higher in the pre-flowering by cucumber mosaic virus in Amaranthus cauda-
stage of the host. Pandy and Gupta (1985), while tus was reported in Himachal Pradesh by Sharma
studying the leaf surface mycoflora of Amaranthus and Chawla (1987) damaging the crop. The other
paniculatus grown in Almora hills, observed 24 viruses affecting the crop are ivy vein clearing
fungal species on leaves of this crop. McLean and virus, alfalfa mosaic virus and beet western
Roy (1988) reported Colletotrichum domatium yellow virus.
Future Prospects in Amaranth
Research 11
Amaranths especially the grain amaranths are other crops has gained a momentum in recent
considered as the golden crop of future. Though years. Breeding programme related to genetic
the cultivation of grain amaranths was initiated in improvement has escalated a lot. But still there
the prehistoric period with a promising note, later are few areas which are to be given emphasis.
it lagged behind conventional crop a lot, because While no fundamental obstacles to the crop’s
a large section of the people rejected that due to future development are apparent, many technical
its unpalatability. Later, a number of research details remain to be explored. Agencies funding
works on unique nutritive value of both vegetable agricultural research for developing countries
and grain amaranths rediscovered the plant and should consider supporting amaranth research
projected the group as useful super crop of future and testing. Some of the thrust areas for recom-
keeping in view its minimum agronomic demand mendations and research needs are listed here:
and food security of huge world population. The
available literatures on amaranth research reveal 1. Collection and screening of germplasm:
that this underutilised crop has not received much Amaranth offers more genetic diversity in its
attention as it deserves. Most of the research present undeveloped state than do many con-
works were concentrated on its nutritive value as ventional crops. The broad geographic
human and animal feed, utilisation of starch, spread of the genus has resulted in the evolu-
basic biology, genetics and breeding practices. A tion of many landraces in widely separated
little work has been done on taxonomy, phylog- areas. Several features of their highly vari-
eny and germplasm maintenance and screening able breeding system and overall reproduc-
and biotechnological approach in breeding tive biology provide ample choice of
programme. Much of the attentions was directed breeding methods. This huge gene pool will
towards grain species of amaranths; vegetable be very important to the future development
group received very little attention. Weed ama- of the crop. Further systematic collections of
ranths too are not evaluated for utilisation in amaranth germplasm should be made in
breeding programme. Thus, the area of amaranth Latin America, the Caribbean, India, Nepal,
research is vast and wide. A multidisciplinary China and the Pacific. These collections
research approach is very much needed to project should be coordinated with the International
amaranths as golden super crop of future. Board on Plant Genetic Resources (IBPGR),
Cultivation of amaranth has gradually crept which is starting germplasm collections in
into a number of countries. Research work on re- Southeast Asia and has recently completed
evaluation of its significance and importance and one in Peru. Proper screening of germplasm
contribution towards genetic manipulation of

168 11 Future Prospects in Amaranth Research
collection is necessary for the future devel- but will also assist people who traditionally
opment of the crop. cultivate the crop.
There is a need to develop an international 4. Role of weedy amaranths: From agronomic
cooperation network to enrich the germ- point of view, amaranth, specially the weedy
plasm collection with exotic elite lines or members, is interesting for their drought
cultivars also to test elite lines and germ- tolerance, low susceptibility to disease and
plasms of amaranths in as many locations as pest (Barba de la Rosa et al. 2009) and envi-
possible under various agroclimatic ronmental plasticity, i.e. ability to grow in
conditions. areas where traditional crops fail to adapt
2. Adaptability trials: Selection and standardi- (Brenner et al. 2000). Amaranths are leading
sation of uniform high-yielding lines for the modern weed science research and have
multilocation yield trials are essential to fish played as a model system for the study of
out types with desirable features like wide weedy nature of plants (Basu et al. 2004).
adaptability in the hill. Studies on inheri- Many amaranth species have developed
tance and combining ability, correlation and unique resistance to few herbicides, recom-
genetic divergence need due emphasis for mended for the control of annual dicot weed.
varietal improvement. The varieties selected Heap (2002) reported about six amaranth
for adaptability trials should include a range species among 25 worst resistant weeds in
of species and morphological types that the world. Amaranth weeds provide valuable
have agronomic potential. This would be an genomic resources that can be utilised for
excellent way to obtain indications of the improving important agronomic crop trait,
geographical areas where given types will like potentiality to generate herbicide resis-
grow best. It would be the first step towards tance. Genome analysis of resistant species
developing ‘zones’ for amaranth adaptabil- can be helpful to explore the evolution of
ity, similar to those used for soybeans. herbicide resistance in plants as well as the
Larger trials, involving perhaps 100–500 possibility to transfer the resistant trait from
varieties, should also be undertaken by weed to crop.
geneticists and screening nurseries. Seeds of the shattering, weedy amaranth
Demonstration trials under farm conditions species (e.g. Amaranthus hybridus,
are also recommended. Amaranthus palmeri, Amaranthus retro-
3. Ethnobotanical aspects of the crop: Studies flexus and Amaranthus spinosus) should not
of the ways people grow and use grain ama- be distributed for cultivation. Nevertheless,
ranths in Central and South America and these weeds can be useful to the amaranth
the Himalayas, as well as those methods breeder. Amaranthus hybridus is the wild
employed to grow and use vegetable ama- progenitor species of present-day cultivated
ranths elsewhere, could be very informative. Amaranthus hypochondriacus and easily
Data could be collected about the most exchanges genes with it. For introducing
favourable environments within which the desirable traits like faster maturation, disease
various species can be grown and much resistance and wider adaptability in the culti-
useful information could be collected by vated forms, the role of Amaranthus hybri-
documenting the rainfall, temperature, day dus could be invaluable. Amaranthus
length and soil conditions of these areas. spinosus (a diploid), sometimes a troublesome
This would enable to target new areas for weed, can be utilised in raising hybrids (F1
amaranth cultivation. It would also demon- triploids) with Amaranthus dubius (a tetra-
strate soil requirements, crop ecology and ploid) on a commercial scale for forage. This
social aspects of amaranth cultivation and is made possible by the peculiar distribution
use. Such studies will not only benefit those of male and female flowers in Amaranthus
wanting to grow amaranth for the first time spinosus. The hybrids are fast-growing and
11 Future Prospects in Amaranth Research 169
sterile and have very soft spines. Feeding 6. Biotechnological approach in genetic
trials and nutritional studies are, however, a improvement of amaranth: Previously the
prerequisite before using such hybrids for genetic improvement of amaranths has been
forage. done by conventional selection method and
5. Taxonomic delimitation in amaranths: hybridisation. But recently biotechnological
Taxonomic disputes in amaranths are still not procedures have been applied and found to
clearly resolved. The presence of a large num- be very effective. A novel protein AmA1 has
ber of morphotypes, landraces and overlap- been isolated from the seeds of Amaranthus
ping morphological features and frequent hypochondriacus and subsequently purified
misapplication of names and a large number and characterised; its cDNA was cloned, and
of synonyms have made taxonomic delimi- through transformation, transgenic potato
tation in amaranths much complicated. and wheat plant has been raised. This has
Subcategorisation in grain amaranth and its opened a new avenue to improve the nutritive
derivation from weed amaranths almost value of several other crops through introduc-
unequivocally resolved applying both mor- tion of AmA1 protein gene by Agrobacterium-
phological and molecular parameters. Weedy mediated transformation.
and grain amaranths are phylogenetically and Much of the research activities done on
historically linked inseparably. Disputes amaranths has focussed on its exceptional
regarding taxonomic segregation in vegetable nutritive value. The main reason could be the
amaranth are still awaiting proper attention. content of protein, fat and active substance
From Lower Gangetic Plain of West Bengal, with antidiabetic, anti-hyperlipidemic, sper-
India, few new species and varieties of vegeta- matogenic and anticholesterolemic effects
ble amaranths have been identified having (Sangameswaran and Jayakar 2008; Girija
specific morphological and reproductive iden- et al. 2011) and antioxidant and antimicro-
tity. The plains of India and other Southeast bial activities (Alvarez-Jubete et al. 2010;
Asian countries are rich source of vegetable Tironi and Aron 2010). Additional interest is
amaranth germplasm with unique diversity. generated by the oil and carbohydrate profile
This diversity should be explored to broaden of amaranth seeds which offers scope for
the vegetable list of nutritive potential. various industrial applications. Nutritive
It is assumed that both the grain and veg- value of vegetable amaranths is somewhat
etable amaranths have originated from their degraded by the presence of some anti-
respective weed progenitors at different nutrients like oxalic acid and nitrates. The
locale of the globe. But unlike grain ama- presence of saponin and phenolic com-
ranth, we don’t have any concrete idea about pounds in grains is responsible for its unpal-
the putative progenitor of vegetable ama- atability. Decarboxylative degradation of
ranth and phylogenetic linkage among them oxalic acid is catalysed by enzyme oxalate
as well. Information on genetic diversity and decarboxylase. A full-length cDNA for oxa-
relationship within and among crop species late decarboxylase was cloned and charac-
and their wild relatives is essential for the terised, and transgenic tomato plant was
fruitful utilisation of the Plant Genetic raised showing oxalate decarboxylase activity.
Resources. It is of common consensus that This approach is very promising for vegeta-
the monoecious grain and vegetable ama- ble amaranths to reduce oxalate problem and
ranths form two distinct groups. The subge- make it more acceptable. A protocol for
nus Acnida represents dioecious weedy Agrobacterium-mediated transformation of
amaranths. Its relation with grain amaranths Amaranthus tricolor has already been devel-
is yet to be explored conclusively which is oped and standardised. This could be utilised
very important for their utilisation in any to produce transgenic amaranth with oxalate
breeding programme. decarboxylase activity.
7. Some specific agronomic requirements: 3. Food quality such as white seeds, palatabil-
From agronomic point of view, though ama- ity and high levels of protein and essential
ranth requires little agronomic attention in amino acids
comparison with other conventional crops, 8. Processing of grain amaranths: Research is
still few steps need to be taken to improve needed on the physiology of postharvest
yield. Amaranth cultivation and harvesting handling, especially on the effect of moisture
practices require several types of research in on grain quality and storage. Studies are also
the following aspects: needed on some other agrotechnical aspects:
1. Selection of type best adapted to local 1. Grain cleaning
conditions 2. Removal of sand and weed seeds from the
2. Application of mixed cropping system in grain
amaranth cultivation and crop rotation 3. Applicability of existing machineries to
3. To learn about the most extreme rainfall, handle amaranth
evaporation and soil characteristics under 4. Drying and storage of harvested grains
which amaranths grow and produce a rea- 5. Processing of the whole grain, such as by
sonable yield extrusion cooking, milling whole and
4. Knowledge about soil requirements, i.e. popped seed and toasting, rolling, sprout-
fertility, tolerance to salinity, need for ing and popping, to assess any changes in
organic matter and maximum and mini- nutritional value or chemical compounds
mum moisture from such processing
5. Knowledge about the effects of growth 6. Commercial development requirements,
conditions on chemical composition and such as dry and wet milling and
nutritive value derivatives
6. Determination of best planting dates, 7. Storage and shelf life of products
plant density and weed and pest 8. Value of the grain and crop residues for
management ensilage and for direct feeding to livestock
7. Development or adaptation of machinery, 9. The use of grain amaranths as food:
planting implements, thresher, winnower Regarding the use of grain amaranths,
and grain cleaner research is needed in the following aspects:
8. Control of diseases and pests causing sig- • Basic characteristics of seed starch, pro-
nificant damage of the crop tein, bran, germ and oil
9. Proper strategies of seed storage to sus- • Uses in products, including breakfast
tain seed viability for longer period of foods and weaning mixtures, as well as
time recipe development
Future research should be directed towards • Nutritional testing in humans
achieving few breeding objectives which • Amaranth’s value as a wheat extender or a
are yet to be achieved. Presently, breeding supplement for added nutritional value in
objectives of grain amaranths prescribe traditional foods such as chapattis, torti-
selection for the following traits: llas, weaning foods, chicha and arepas
1. Desirable growth characteristics such as • Its use in infant foods
reduced plant size, reduced sensitivity to • Nutritional availability of minerals, vita-
photoperiod, synchronous flowering, early mins, proteins and starch
maturity, reduced lodging and uniform • Amaranth’s functional characteristics (vis-
drydown cosity, density, freeze-thaw stability, heat
2. Environmental adaptations such as drought stability, emulsifying properties) when
tolerance, pest and disease resistance, her- used in foods, and how grain types differ
bicide tolerance and efficient fertiliser from one another in those characteristics.
utilisation • Anti-nutritional constituents
11 Future Prospects in Amaranth Research 171
10. Vegetable amaranths: Vegetable amaranths manner. The nutraceutical activities of ama-
have been more thoroughly investigated than ranths are yet to be explored adequately.
the grain amaranths in Asia and more There is a need to evaluate the impact of pro-
specifically in Southeast Asia. Several cessing and cooking on the nutritional prop-
important aspects are still left for improve- erties of the species. Amaranth can also be a
ment. Selections have been made by Asian potent source of natural dyes and
growers for many years; varieties have been pharmaceuticals.
identified suitable for widespread culture. 12. Environmental impact: It is important to
Nevertheless, further improvement of the study the weediness of the most problematic
crop could be achieved by studies of the Amaranthus species and the likelihood of
following: their becoming pests. Amaranth pollen and
1. Pest and disease resistance grain may cause allergic reactions in some
2. Nutrient uptake and nutrient content at people, and this needs to be addressed. Both
different stages of harvest or crop growth the grain- and vegetable-type amaranths
3. Leaf yield could provide many nutritious foods for the
4. Food quality, including tenderness and world. The small seed size is a limitation in
storage methods to prolong the life of planting as well as in harvesting, threshing
the harvested produce and cleaning the grain. But modern experi-
5. The use of amaranth leaves as a remedy ence in the Northern Indian plains shows
for vitamin A deficiency that they have a good chance of adapting
6. Anti-nutritional factors and heavy-metal successfully. They might complement other
accumulation in response to type and cereals such as sorghum, millets or barley,
quantity of fertilisers used and type of thus helping countries that import large
soil amounts of wheat. In addition, they would
7. Production of leaf-nutrient concentrate provide a local source of feed grain for the
8. Regrowth after harvest poultry industries of developing nations.
9. Comparison of yield from clipping ver- Grain amaranths are the new promising crop
sus successive planting for drylands (areas with 600–800 mm of
10. Seed production and farmer-selection rainfall per year) and for tropical highlands
techniques up to extreme elevations (3500 m and above)
11. Leaf/stem ratio and as a quick-maturing, dry-season crop
12. Late emergence of inflorescences for monsoon areas. It would be a daydream-
13. Planting and cultural practices for effi- ing and too much optimism to expect ama-
cient use of land, water and fertiliser ranth to be on dinner plates next year; it took
14. Crop rotation to avoid soil-borne a century for the American people and the
diseases farmers to accept the soybean, and it took
15. Proper timing of harvest two centuries for Europeans to give recogni-
16. Benefits and possible toxic problems of tion to potato. Comparing with such now-
vegetable amaranth as a forage established crops, amaranth has attracted the
11. New uses: The germs and brans of grain attention of scientific research or testing
amaranths contain about 20 % oil. There is a though in small scale. With the help of
need to study and screen the germplasm for today’s communications and technology, the
edible oil. The amaranth oil is the rich source day is not far away when amaranth would
of squalene (a high-priced material found in find its niche. Within a few years, it seems
amaranth seed but normally obtained from likely that this ancient grain of the Americas
shark livers and used in cosmetics). The will return to grace in the modem age.
industrial application of amaranths is an area Eventually, it may prove to be as a rich leg-
which is to be explored in much extensive
acy of the American Indian as maize and fast cereal is available. Research has mainly
beans. emphasised grain amaranths so far, but in
Despite the growing evidence in ama- 1967 FAO started investigation on vegeta-
ranth’s favour, much research needs to be ble amaranth. The following year it began
done before the crop be commercially pro- field experiments in home garden projects in
duced on large scale like conventional cere- Nigeria and Benin. Later it commissioned
als and widely accepted. Nevertheless, the germplasm collections. As a result, the veg-
researchers are studying the crop’s responses etable branch of the amaranth family is
to climate, soil conditions, pests and dis- beginning to attract recognition, and FAO
eases. Also, they are engaged in breeding of has published a report on these species
short-statured plants of uniform height with (Grubben and Van Sloten 1981).
sturdy, wind-resistant stalks and high-yield- In any country among the crops under
ing seedheads that hold their seeds until they cultivation, some may be native, and some
are harvested. Much of the amaranth’s devel- may be of non-native origin from other
opment has been done in the Rodale Research regions. The Plant Genetic Resources for
Centre near Emmaus, Pennsylvania, where Food and Agriculture (PGRFA) formulated
more than a thousand different accessions the basis for the establishment of a multilat-
collected from all parts of the world were eral system of access and benefit sharing
bred, grown and evaluated. Further collabo- which is applicable to a list of crops under a
ration has been initiated with scientists in Standard Material Transfer Agreement
Africa, Asia and Latin America; as a result, (SMTA) for food security and interdepen-
plant lines have been selected to overcome dence irrespective of the origin of the crop.
tendencies towards lodging, seed shattering, Enforcement of the Convention on
indeterminate growth, succulence at harvest- Biological Diversity (CBD) from 1993 and
ing time and day-length dependence. This provisions under trade-related aspects of
research effort has produced grains with Intellectual Property Rights (TRIPS) led to
improved baking, milling, popping and taste the apprehension that exchange of germ-
qualities, as well as machinery adapted to plasm would get restricted. To increase the
planting, cultivating, harvesting and thresh- food production at global level on sustain-
ing the crop. Lines of uniform colour and able basis, dependence on crop genetic
height that bear their seedheads above the resources that originated from different geo-
leaves, thus making them suitable for graphical locations through introduction and
mechanical harvest, are now available. The exchange is inevitable. This holds good
crop can be said to be on the threshold of especially in the case of underutilised crop
limited commercial production in the USA. species which have same centre of origin
Several companies are testing the grain in and centre of domestication because domes-
their products, and an amaranth-based break- tication process is still in evolutionary phase.
Appendices
Appendix I IRRI International Rice Research

Institute
Abbreviation ISTA International Seed Testing
Association
AICRP All India Coordinated Research LIFDC’s Low-Income Food-Deficit
Project Countries
ALS Acetolactate Synthase MAF Major Allele Frequency
AVRDC Asian Vegetable Research and NAS National Academy of Sciences
Development Center NBPGR National Bureau of Plant Genetic
CBD Convention on Biological Resources
Diversity NCRPIS North Central Regional Plant
CGIAR Consultative Group on International Introduction Station
Agricultural Research NPU Net Protein Utilization
CIAT International Center for Tropical PER Protein Efficiency Ratio
Agriculture PGR Plant Genetic Resources
CSIRO Commonwealth Scientific PGRFA Plant Genetic Resources for Food
and Industrial Research and Agriculture
Organisation PIC Polymorphic Information Content
EPSPS 5’-enolpyruvyl-shikimate-3- PPO Protoporphyrinogen Oxidase
phosphate synthase ROS Reactive Oxygen Species
FAO Food and Agriculture Organization RRC- Rodale Research Center
GCA General Combining Ability SCA Special Combining Ability
IBPGR International Board of Plant SMTA Standard Material Transfer
genetic Resources Agreement
ICAR Indian Council of Agricultural SNPs Single Nucleotide Polymorphisms
Research TILLING Targeting Induced Local Lesion
ICARDA International Center for IN Genomes
Agricultural Research in the Dry TRIPS Trade-Related Aspects of
Areas Intellectual Property Rights
IPGRI International Plant Genetic UPGMA Unweighted Pair Group Method
Resources Institute with Arithmetic mean
IITA International Institute of Tropical VPKAS Vivekananda Parvatiya krishi
Agriculture Anusandhn Shala

S. Das, Amaranthus: A Promising Crop of Future, DOI 10.1007/978-981-10-1469-7
174 Appendices
USDA-ARS United States Department of Appendix II

Agriculture-Agricultural Research
Service Compiled list of Amaranthus species (including
USDA-GRIN United States Department of synonyms) based on Australian Plant Name
Agriculture-Germplasm Index (APNI), International Plant Names Index
Resources Information Network (IPNI), Index Kewensis (IK) and Gray Card
UUC Underutilized Crop Index (GCI)
Species Synonyms References

Amaranthus abyssinicus hort. ex Man. Cult. Pl. 252(1924) in Syn
L.H. Bailey
Amaranthus acanthobracteatus Sida 21(1): 12. 2004
Henrickson
Amaranthus acanthocarpa Wohlpart & Mabry 1968
Amaranthus acanthochiton J.D. Sauer = Acanthochiton wrightii Torrey in Madrono 13:44, 1955
L. Sitgreaves
Amaranthus acroglochin Spreng Syst.Veg. (ed 16) [Sprengel] 1:
927.1824
Amaranthus acutilobus Uline & Bot. Gaz. 19: 320.1894
W.L. Bray
Amaranthus adulterinus Thell. Repert. Spec. Nov. Regni Veg.
24: 301, hybr. 1928
Amaranthus aeneus Besser Cat. Hort. Volhyn. (1816) 8
Amaranthus affinis Thell. Repert. Spec. Nov. Regni Veg.
21: 324. 1925
Amaranthus albiformis Moq. Prodr. [A.P. de Candolle] 13(2):
263.1849
Amaranthus albomarginatus Uline & Bot. Gaz. 19: 318.1894
W.L. Bray.
Amaranthus albus L. = A. albus var. pubescens (Uline & Fl. Cap. (Thunberg, ed.2) 215.
Bray) Fernald 1823
= A. pubescens (Uline & Bray)
Rydberg
Amaranthus alius E.H.L.Krause Beih. Bot. Centralbl., Abt. 2.
33(2): 481, sp. aggreg. 1915
Amaranthus alopecurus Hochst ex Index Seminum [Berlin] 1:
A. Br. & C.D. Bouche 1872. Sauer (1950)
Amaranthus altissimus Riddell Acnida altissimus (Riddell) Moq. Syn. Fl. West States 41: 1835
Amaranthus amboinicus Buch.-Ham. Numer. List [Wallich] n.
6987.1832
Amaranthus ambigens Standley N. Amer. Fl. 21(2) : 106.1917
Amaranthus anacardana Hook.f. Fl. Brit. India [J.D. Hooker]
4(12): 719. 1885
Amaranthus anardana Buch.-Ham. ex = A. hypochondriacus L. Numer. List. [Wallich] n. 6903.
Moq. 1832
Amaranthus andersonii J.T. Howell = Scleropus urceolatus Andersson Proc. Calif. Acad. Sci. Ser. 4,
21: 95. 1933
Amaranthus angustifolius Lam. = A. graecizans ssp. graecizans Encyclopedie Methodique
= A. blitum var. graecizans Botanique 1: 115. 1783
= A. graecizans L.
Amaranthus annectens S.F. Blake J. Bot. 53: 103, 1915
(continued)
Appendices 175

Amaranthus aragonensis Sennen Bull. Geogr. Bot. 1911, xxi.123
Amaranthus arardhanus Sweet Hort. Brit. [Sweet] ed. 3.
569.1839.
Amaranthus arctioideus Perr. ex Moq. Prodr. [A.P. de Candolle] 13(2):
332. 1849
Amaranthus arenicola I.M. Johnston J. Arnold Arbor. 29: 193. 1948
Amaranthus artineanus Muschl. Man. Fl. Egypt i 311 (1912).
hybr.
Amaranthus arvensis E.H.L.Krause Deutschl. Fl. (Sturm) ed. 2. 5:
137. 1901
Amaranthus ascendens Loiseleur = A. ascendens sensu auct Japon = A. Not. Fl. France 141. 1810.
lividus L.
= Euxolus viridis var. ascendens
(Lois) Moq.
Amaranthus aschersonianusThell. Graebn. Syn. Mitteleur. Fl. V.
309 (1914)
Amaranthus asplundii Thell. Repert. Spec. Nov. Regni Veg.
21: 322. 1925
Amaranthus ataco Thell. Repert. Spec. Nov. Regni Veg.
16: 23. 1919.
Amaranthus atropurpureus Roxb. = A. oleraceus Fl. Brit. Ind. [J.D. Hooker] 3.
608. 1885
Amaranthus atrosanguineus Hort. (18380 ex Moq. In DC. Prod.
Lugd. Xiii.II 266
Amaranthus aureus Hort. ex Moq. = A. hybridus L. Prodr. [A.P. de Candolle] 13(2):
259. 1849. Kirpicznikov (1969)
Amaranthus australis (A. Gray) = Acnida australis A. Gray Madrono 13: 15. 1955.
J.D. Sauer = A. alabamensis Standley Sauer (1955)
= A. cannabina L. var. australis
(A. Gray) Uline & W.L. Bray
= A. cuspidata Bertero ex Sprengel
Amaranthus bahiensis Mart. Herb. Fl. Bras.n. 969
Amaranthus batalleri Sennen Butl. Inst.Catalana Hist.
Nat.1932 xxxii.iii. in syn
Amaranthus bellardii Hort. ex Moq. Prodr. [A.P. deCandolle] 13(2):
259. 1849
Amaranthus bengalensis Saubhik Das Phytotaxa 181(5): 297 2014
& Iamonico
Amaranthus berlandieri (Moq.) Uline Bot. Gaz. 19: 268. 1894
& W.L. Bray
Amaranthus berchtholdi Hort. ex Prodr. [A.P.deCandolle] 13(2):
Moq. 259. 1849
Amaranthus berchtoldii Seidl. ex Boehm. Gen 164. Natural 1.
Opiz. 1823
Amaranthus bicolor Nocca ex Willd. Sp.Pl. ed. 4. [Willdenow] 4(1):
384. 1805
Amaranthus bernhardi hort ex Moq. Prodr. [A.P. deCandolle] 13(2):
258. 1849
Amaranthus bigelovii Uline & Bot. Gaz. 19: 271. 1894
W.L. Bray
Amaranthus blitoides S. Watson Proc. Amer. Acad. Arts. 12: 273.
1877
(continued)
176 Appendices

Amaranthus blitonius St. Lag Ann. Soc. Bot. Lyon vii: 119
(1880)
Amaranthus blitum L. = A. lividus L. Sp.Pl. 2: 990. 1753
= A. ascendens Loiseleur Feine (1981); Robertson (1981);
= A. lividus ssp. lividus Sreelathakumary & Peter
(1993);
= A. blitum var. blitum
= A. blitum auct. non L.
= A. graecizans L.
Amaranthus blitum var. oleraceus (L.) = Cultivated form of A.lividus L. Fl. Brit. India 4: 721. 1885
Hook.F.
Amaranthus blitum var. ascendens Cat. Pl. Horti Monsp. 4. 1813
(Loisel.) DC
Amaranthus bouchonii Thell. = Aberrant form of A.powellii Le Monde des Plantes 27(160):
= Amaranthus bouchonii Thell. 4. 1926
= A. powellii
Amaranthus parganensis Saubhik Das Novon 23(4): 406. 2015
Amaranthus bengalense Saubhik Das Phytotaxa 181(5): 297. 2014
& Iamonico
Amaranthus brandegeei Standl. N. Amer. Fl. 2: 109. 1917. Sauer
(1978)
Amaranthus brownie m Christoph. & Bull. Bernice P. Bishop Mus. 81:
Caum 25. 1931
Amaranthus buchtienianus Thell. Repert. Spec. Nov. Regni Veg.
21: 323 91925)
Amaranthus cacciatoi (Aellen ex Willdenowia 43(2): 239 (2013)
Cacciato) Iamonico
Amaranthus californicus (Moquin- = Mengea californica Moq. Bot. California [W.h. Brewer] 2:
Tandon) S. Watson 42.1880
Amaranthus campestris Willd. Sp. Pl. ed. 4 [Willdenow] 4(1):
382. 1805
Hooker (1885); Tanaka (1976)
Amaranthus canariensis Besser Cat. Hort. Volhyn. (1816) 8.
Amaranthus cannabinus (L.) = Acnida cannabina L. Madrono 13: 11.1955
J.D. Sauer
Amaranthus capensis Thell. Syn. Mitteleur. Fl. [Ascherson &
Graebner]. 5(1): 293. 1914
Amaranthus capitatus Cat. Hort. Turic. (1827–28)
Amaranthus caracam Besser Cat. Hort. Volhyn (1816)g
Amaranthus caracasanus Kunth Nov. Gen. Sp. [H.B.K] 2:195
(1818)
Amaranthus caracu Zucc. ex Steud. Nomencl. Bot. [Steudel] ed. 2.
1:69. 1840.
Amaranthus cararia Besser Cat. Hort. Volhyn (1816) 8
Amaranthus cararu Jacq ex Zuccagni Cent. Observ. Bot. [p.46] No.
94. 1806
Amaranthus cardenasianus Hunz. Bol. Soc. Argent. Bot. Iv. 136
(1951)
Amaranthus carneus Greene Pittonia 2(8): 105. (1890)
Amaranthus carolinae Kov. Nauch. Trud. Vissh Selskostop.
Inst. Plovdiv 23(1): 51. 1978
(continued)
Appendices 177

Amaranthus cathecu Hort. ex Moq. Prodr. [A.P. deCandolle] 13(2):
259. (1849)
Amaranthus caturus B. Heyne ex Fl. Brit. India [J.D. Hooker]
Hook.f. 4(12): 720. 1885
Amaranthus caudatus L. Sp. Pl. 2: 990 1753
Amaranthus cauliflorus Link. Enum. Hort. Berol. Alt. 2:389.
1822
Amaranthus celosioides Kunth Nov. Gen. Sp [H.B.K.] 2: 194.
1818
Amaranthus centralis J.Palmer & Nuytsia 19(1); 111. (2009)
Mowatt
Amaranthus cernuus Besser In Cat. Hort. Turic. (1827–8)
“An Celosia”?
Amaranthus chihuahensis S. Watson Proc. Amer. Acad. Arts 21:436.
(1886)
Amaranthus chipendalei Kov. Nauch. Trud. Vissh Selskostop.
Inst. Plovdiv 23(1): 52. (1978)
Amaranthus chlorostachys Willd. = A. hybridus ssp. Hybridus Hist. Amaranth. 34 (t.10,fig.19).
var. hybridus 1790
= A. hybridus
= A. chlorostachys auct.
= A. powellii
= A. hybridus var. hybridus
Amaranthus circinnatus Hort. Parisex Encycl. [J. Lamarck & al] Suppl.
Poir. 1.311. 1810
Amaranthus clementii Domin Bibliotheca Botanica 89(4) 1928
Amaranthus coesius F. Dietr. Ex Moq. Prodr. [A.P. de Candolle] 13(2):
266. 1849.
Amaranthus communicates Kerner Oester. Bot. Z. 25: 194.1875
Amaranthus coracanus Mart. Hort. Erlang. 197
Amaranthus congestus C.C. Townsend Kew Bull. 43(1): 103. (1988)
Amaranthus crassipes Schlechtendal = A. crassipes var. warnockii Linnaea 6: 757. (1831)
(I.M. Johnston) Hendrickson
= A. warnockii I.M. Johnston
Amaranthus crispus Terrac. Atti. Accad. Sc. Napoli Ser. 2 iv.
(1890) App. 2
Amaranthus cristatus Noronha Verh. Batav. Genootsch. Kunst.
5(Art. 4) : 7. 1790
Amaranthus cristulatus Speg. Comun. Mus. Nac. B. Aires 1:
345. 1901
Amaranthus crocatus Besser Cat. Hort. Volhyn. (1816) g
Amaranthus cruentus L. = A. hybridus (L.) ssp. cruentus Syst. Nat. Ed. 10. 2:1269. 1759
L. Thellung
Amaranthus cruentus L. var. patulus Bull. Jard. Bot. Natl. Belg.
(Bertol) Lambinon 47(1–2): 247. 1977.
Amaranthus cruentus L. var. albus Nordic J. Bot. 30(4): 418. 2012
Saubhik Das
Amaranthus curvifolius Spreng. Syst. Veg. ed. 16 [Sprengel] 1:
928. 1824
Amaranthus cuspidifolius Domin Bibliotheca Botanica Ixxxix 78.
1921
(continued)
178 Appendices

Amaranthus deflexus L. Mantissa Plantarum Altera 295.
1771
Amaranthus delilei Richter & Loret Bull. Soc. Bot. France 13: 316.
1868.
Amaranthus desfontanii Kov. Nauch. Trud. Vissh Selskostop.
Inst. Plovdiv. 23(1): 51. 1978
Amaranthus diacanthus Raf. Fl. Ludov. 31. 1817
Amaranthus diandrus Spreng. Neue Entdeck. Pflanzenk. 3:20.
1822.
Amaranthus diffusum Dulac Fl. Hautes-Pyrences 174. 1867
Amaranthus dioicus Michx ex Moq. Prodr. [A.P. de Candolle] 13(2):
277. 1849
Amaranthus dinteri Schinz Mem. Herb. Boiss. No. 29. 15
Amaranthus divaricatus Andrz. ex Prod. Fl. Cherson. 185.
Lindem.
Amaranthus dubius Mart ex Thell. = Tetraploid Fl. Adv. Montpellier 38: 203.
1912. Thell. in Aschers &
Graebn. Syn. Mitteleur Fl. 591):
265. 1914
Amaranthus dussil Sprenger Bull. Reale Soc. Tosc. Ortic. 21:
178. 1896.
Amaranthus edulis Speg. = A. caudatus L. Physis (Buenos Aires) 3: 163.
=A. caudatus ssp. Mantegazzianus 1917
Amaranthus edulis var. spadiceus Revista Argent. Agron. 10: 330.
Hunz. 1943
Amaranthus edulis var. typicus Hunz. Revista Argent. Agron. 10: 330.
1943
Amaranthus emarginatus Salzm. ex Bot. Gaz. 19: 319. 1894
Uline & W.L. Bray
Amaranthus enervis (F. Muell.) Kov. Nauch. Trud. Vissh Selskostop
Inst. Plovdiv 23(1): 50. 1978
Amaranthus esculentus Besser Cat. Hort. Volhyn. (1816) g. Ex
Moq. in DC. Prod. Xiii. II.266
Amaranthus eugenii Sennen Butl. Inst. Catalana Hist. Nat.
Xxxii. 111. 1932.
Amaranthus farinaceous Roxb. ex Prodr. [A.P. de Candolle] 13(2):
Moq. 266. 1849; Sauer (1950)
Amaranthus fimbriatus (Torr.) = A. fimbriatus var. denticulatus Bot. California [W.H. Brewer]
Bentham ex S. Watson (Torrey) Uline & W.L. Bray 2:42. 1880
Amaranthus flavus L. = A. hypochondriacus L. Syst. Nat. Ed. 10. 2: 1269. 1759;
Feine (1981)
Amaranthus floridanus (S. Watson) = Acnida floridana S. Watson Madrono 13: 25. 1955; Sauer
J.D. Sauer (1955)
Amaranthus frumentaceous = A. paniculatus Buch.-Ham. in Roxb. Fl. Ind. Iii.
Buch.-Ham. var. frumentaceous 699
= A. hypochondriacus L.
= A. hybridus ssp. hybridus
Amaranthus filicaulis Sennen Cavanillesia ii 34. 1929. Hybr. ?
Amaranthus flexuosus Ambrosi Fl. Tirolo Mer. 2: 187. 1857
Amaranthus floridus Benth. Bot. Voy. Sulphur [Bentham]
158, t. 51. 1846
(continued)
Appendices 179

Amaranthus frutescens Hort. ex Moq. Prodr. [A.P. de Candolle] 13(2):
348. 1849
Amaranthus furcatus J.T. Howell Proc. Calif. Acad. Sci. Ser. 4,
21: 94. 1933
Amaranthus galii Sennen & Gonzalo Cavanillesia ii. 34. (1929) hybr.
Amaranthus gangeticus L. Syst. Nat. Ed. 10. 2: 1268. 1759
Amaranthus giganteus Besser Cat. Hort. Volhyn. (1816)
91 F. G. Dietr. Lexik. Gaertn. i.
313.
Amaranthus glaucus Biv. Nuove Piante II, footnote, 1838.
Amaranthus glomeratus Posp. Fl. Oesterr. Kustent. i. 375. 1897
Amaranthus gracilis Desf. = A. viridis L. Tabl. Ecole Bot. 43. 1804
Amaranthus graecizans L. = A. albus L. Sp. Pl. 2: 990. 1753
= A. graecizans var. graecizans
Amaranthus grandiflorus (J.M. Black) Transactions and Proceedings of
J.M. Black the Royal Society of South
Australia 60. 1936
Amaranthus greggii S. Watson Proc. Amer. Acad. Arts 12:
274.1877
Amaranthus guadeloupensis Hort. ex Prodr. [A.P. de candolle] 13(2):
Moq. 257. 1849
Amaranthus hierichuntinus Vis. In Att. Ist. Ven. Sc. Ser. III, iv.
(1858–59) 139
Amaranthus haughtii Standl. Publ. Field Mus. Nat. Hist. Bot.
Ser. 11: 149. 1936
Amaranthus hungaricus Soo. Repert. Spec. Nov. Regni Veg.
22: 318, hybr.1926
Amaranthus hunzikeri N. Bayon Novon 17(3): 294. 2007
Amaranthus huttonii Hort. Veitch. Gard. Chron. 215 (1872)
Amaranthus hybridus L. Sp. Pl. 2: 990. 1753
A. bybridus var. bouchonii (Thell.) Candollea 52(2): 273. 1997
lambinon
A. hybridus L. var. chlorostachya = A. chlorostachya Willd. Fl. Advent. Montpellier 205.
Thell. 1912
A. hybridus L. var. densus Farw = A. paniculata L. var. densus Regel. Rep. (Annual) Michigan Acad.
Sci. 20; 175. 1918
A. hybridus L. var. erythrostachys Prodr. [A.P. de Candolle] 13(2):
Moq. 259. 1849
A. hybridus L. var. hecticus (Willd.) = A. hecticus Willd. Prodr. [A.P. de Candolle] 13(2):
Moq. 260. 1849
A. hybridus L. var. hypochondriacus = A. hypochondriacus Rhodera 10: 32. 1908
(L.) B.L. Rob.
A. hybridus L. var. paniculatus Uline = A. paniculatus Mem. Torry Bot. Club 5: 145.
& W.L. Bray 1894
A. hybridus L. var. batalieri (Sennen) Collect. Bot. (Barcelona) 11:
Carretero 129.1979
A. hybridus L. var. bellardii Moq. Prodr. [A. P.de Candolle] 13(2):
259. 1849
A. hybridus L. var. laetus (Willd.) = A. laetus Willd. Prodr. [A. P.de Candolle] 13(2):
Moq. 259. 1849
(continued)
180 Appendices

A. hybridus L. var. leucocarpus = A. leucocarpus Revista Argent. Agron. 10:
(S. Watson) Hunz. 340.1943
A. hybridus L. var. patulus Thell. = A. patulus Bertol Fl. Advent. Montpellier 206.
1912
A. hybridus L. var. pergaminensis Darwiniana 5: 336. 1941
Covas
A. hybridus L. var. prostrate Moq. Prodr. [A. P.de Candolle] 13(2):
260. 1849
A. hybridus L. var. pseudoretroflexus Collect. Bot. (Barcelona) 11:
(Thell.) Carretero 125. 1979
A. hybridus L. var. quitensis (Kunth) = A. quitensis Kunth Darwiniana 5: 336. 1941
Covas
A. hybridus L. var. rubricaulis Moq. Prodr. [A. P.de Candolle] 13(2):
259. 1849
A. hybridus L. var. sanguineus Farw. = A. paniculatus L. var. sanguineus Rep (Annual) Michigan Acad.
Regel. Sci. 20; 175. 1918
Amaranthus hypochondriacus L. = A. hybridus ssp. hybridus var. Sp. Pl. 2: 991. 1753
erythrostachys
= A. hybridus var. erythrostachys
Amaranthus hypochondriacus L. var. Prodr. [A. P.de Candolle] 13(2):
macrostachys Moq. 256. 1849
monstrosus Moq. 256. 1849
Amaranthus hypochondriacus L. var. = A. powellii S. Watson Monogr. Syst. Bot. Missouri
powellii (S. Watson) Pedersen Bot. Gard. 74(2): 1245. 1999
racemosus Moq. 256. 1849
tortuosus Moq. 256. 1849
Amaranthus inamoenus Willd. = A. mangostanus Sp. Pl. Ed. 4 [Willdenow] 491):
386. 1805
Amaranthus incarnates Hort. Ex Moq. Prodr. [A. P.de Candolle] 13(2):
257. 1849
Amaranthus incomptus Willd. Enum. Hort. Berol. 64
Amaranthus incurvatus Gren. & Godr. Prosp. Fl. Fr. 8
Amaranthus induratus C.A. Gardner Nuytsia 19(1):117. 2009
ex Palmer & Mowatt
Amaranthus intermedius Guss. ex Prodr. [A. P.de Candolle] 13(2):
Moq. 259. 1849
Amaranthus interruptus R. Br. Prodr. Fl. Nov. Holland.
414.1810
Amaranthus lecocarpus S. Watson = A.leucospermum S. Watson. Proc. Amer. Acad. Arts 10:
= A. hybridus L. 347.1875
= A. hypochondriacus
= A. paniculatus
Amaranthus jansen-wachterianus In Graebn Syn. Mitteleur. Fl. v.
Thell. 347 (1914)
Amaranthus japonicas Hoult. ex Sp. Pl. Ed. 4 [Willdenow] 4(1);
Willd. 386. In syn. 1805
Amaranthus johnstonii Kov. Nauch. Trud. Vissh Selskostop.
Inst. Plovdiv 23(1): 50. 1978.
nom.nov
(continued)
Appendices 181

Amaranthus jonesii (Uline & Nauch. Trud. Vissh. Selskostop.
W.L. Bray) Kov. Inst. Plovdiv 23(1); 49.1978
Amaranthus kloosianus Hunz. Bol. Soc. Argent. Bot.
4:138.1951
Amaranthus laetus Willd. Amaranth 28; Sp. Pl. Iv. 391
(1805)
Amaranthus lanatus Dum. Cours. Bot. Cult. 1: 640. 1802
Amaranthus lancifolius Roxb. Hort. Bengal 67
Amaranthus lancifolius Del. Ex Moq. Prodr. [A.P. de Candolle] 13(2):
276. 1849
Amaranthus lanceolatus Roxb. Fl. Ind. III. 607
Amaranthus laxiflorus Comell ex Poll. Fl. Veron. Iii. 114. 1824
Amaranthus lecocarpus S. Watson = A. leucospermus S. Watson Proc. Amer. Acad. Arts 10: 347.
1875
Amaranthus lineatus R. Br. Prodr. Fl. Nov. Holland.
414.1810
Amaranthus littoralis Bernh. In Hort. Tur. (1813) ex Moq. in
DC. Prod. Xiii. II.274
Amaranthus lividus L. Sp. Pl. 2: 990.1753
Amaranthus lividus L. var. ascendens Adent. Fl. Tweedside 177.1919
(Loisel.) Hayw. & Druce
Amaranthus lividus L. var. = Euxolus viridis var. polygonoides Rep. Bot. Soc. Exch. Club Brit.
polygonoides (Moq.) Thell. ex Druce Moq. Isles 5: 574. 1920
Amaranthus lombardoi Hunz. Bol. Soc. Argent.
Bot.4:141.1951
Amaranthus looseri Suess. Littoa 4:128. 1939
Amaranthus macrocarpus Benth. Fl. Austral. 5: 216. 1870
Amaranthus macrocaulos Poir. Encycl. [J. Lamarck & al.]
Suppl. 1: 314. 1810
Amaranthus macrostachyus Merat ex Prodr. [A.P. de Candolle] 13(2):
Moq. 256. 1849
Amaranthus major Salzm.ex Moq. Prodr. [A.P. de Candolle] 13(2):
274. 1849
Amaranthus mangostanus L. = A. tricolor L. Cent. Pl. I. 32. 1755
Amaranthus mantegazzianus Passer. =A. caudatus (differs significantly in In Hort. Parm. 4. 1864;
DNA content as in A. caudatus) Greizerstein & Poggio (1994)
= A. caudatus var. edulis
= A. caudatus ssp. edulis
Amaranthus margaritae Dam. Wiener III. Gart.-Zeitung (1887)
433-35
Amaranthus melancholicus L. = A. tricolor L. Sp. Pl. 2: 989. 1753
Amaranthus miamiensis Riddell Syn. Fl. West St. 41. 1835
Amaranthus microphyllus Shinners Sida 1: 248. 1964
Amaranthus miniatus Hort. Avign. Ex Fl. Brit. India [J.D. Hooker]
Hook.f. 4(12): 721. 1885
Amaranthus minimus Standl. N. Amer. Fl. 21(2): 119. 1917
Amaranthus minor Gray. Nat. Arr. Brit. Pl. 2: 289. 1821
Amaranthus mitchellii Benth. Fl. Austral. 5: 214. 1870
Amaranthus monstrosus Cal. Hort. Div. (1837); Hort. Tonelle
ex Moq. in DC Prod.
(continued)
182 Appendices

Amaranthus moquinii Sennen. Cavanillesia ii. 34. 1929
Amaranthus morosus Rchb. Fl. Germ. Excurs 585
Amaranthus mucronatus Poir. Encycl. [J. Lamarck & al.]
Suppl. 1. 311. 1810.
Amaranthus mucronatus Hort. Petrop. Fl. Brit. India [J.D. Hooker]
ex Hook.f. 4(2): 720. 1885.
Amaranthus muelleri (Uline & Nauch. Trud. Vissh. Selskostop
W.L. Bray) Kov. Inst. Plovdiv 23(1): 50. 1978
Amaranthus muricatus Gillies ex = Euloxux muricatus Gillies ex Moq. Prodr. [A.P. de Candolle] 13(2):
Moq. 276. 1849.
Amaranthus myrianthus Standl. Bull. Torry Bot. Club 41:
506.1914.
Amaranthus necticus Willd.
Amaranthus neglectus Hort. ex Moq. Prodr. [A.P. de Candolle] 13(2):
259. 1849.
Amaranthus nepalensis Cal. Hort. Lugd. (1836); ex Moq. in
DC Prod. Xii. II. 259
Amaranthus nettii Kov. Nauch. Trud. Vissh. Selskostop
Inst. Plovdiv 23(1): 50. 1978
Amaranthus obcordatus Standl. N. Amer. Fl. 21(2): 107. 1917
Amaranthus obovatus S. Watson Proc. Amer. Acad. Arts xii. 275.
1877
Amaranthus oleraceus L. = A. blitum var. oleraceus Sp.Pl. ed. 2. 2: 1403. 1763
= A. lividus ssp. lividus
Amaranthus obtusiflorus (Mart.) Kov. Nauch. Trud. Vissh. Selskostop
Inst. Plovdiv 23(1): 50. 1978
Amaranthus officinalis Gromov ex = A. blitum subsp. oleraceus (L.) Trudy Imp. S.-Peterburgsk. Bot.
Trautv. Costea Sada 9. 139. 1884.
Amaranthus olitorius Besser Cat. Hort. Volhyn. (1816) g
Amaranthus pachystachys Rchb. ex Prodr.[A.P. de Candolle] 13(2):
Moq. 265. 1849.
Amaranthus pallidiflorus F. Muell Fragm. (Mueller) 1(5): 140.
1859
Amaranthus pallidus M. Bieb. Fl. Taur.–Caucas. 2: 399.1808
Amaranthus palmeri S. Watson Proc. Amer. Acad. Arts 12:
274.1877.
Amaranthus paniculatus L. = A. hybridus L. var. paniculatus Sp. Pl. ed. 2. 2: 1406. 1763
(L.) Thell.
Amaranthus paniculatus L. var. = A. cruentus L. Prodr. [A.P. de Candolle] 13(2):
cruentus (L.) Moq. 257. 1849.
Amaranthus paniculatus L. var. Fl. Bras. (Maritius) 5(1):
purpurascens Seub. 238.1875
Amaranthus paniculatus L. var. Prodr. [A.P. de Candolle] 13(2):
sanguineus (L.) Moq. 238. 1849.
Amaranthus paniculatus L. var. = A. speciosus Sims. Stand. Cycl. Hort. 1: 270. 1914
speciosus L.H. Bailey
Amaranthus paolii Chiov. Nuovo Giorn. Bot. Ital. Ser. 2,
34: 845. 1927
Amaranthus parganensis Saubhik Das Novon 23(4): 406. 2015
Amaranthus paraguayensis D. Parodi Anales Soc. Cl. Argent. V. 273.
1878
(continued)
Appendices 183

Amaranthus parisiensis Schkuhr. Bot. Handb. [C. Schkuhr] 3:
249. 1802
Amaranthus parodii Standl. Publ. Field Mus. Nat. Hist., Bot.
Ser. 17: 240. 1937
Amaranthus parvulus Peter Repert. Spec. Nov. Regni Veg.
Beih. 40: 25. 1932
Amaranthus patulus Bertol = A. hybridus L. Comment Itin. Neapol. 171,t.
12. 1837
= A. patulus auct. non L.
= A. cruentus L.
Amaranthus pedersenianus N. Bayon Novon 22(2): 133. 2012
& C. Pelaez
Amaranthus pendulus Hort. ex Moq. Prodr. [A.P. de Candolle] 13(2):
255. 1849.
Amaranthus perennis Bellardi Ex Colla, Herb. Pedem. Iv. 578.
1835
Amaranthus persicarioides Hort. ex Encycl. [J. Lamarck & al] Suppl.
Poir. 1. 311. 1810
Amaranthus persimilis Hunz. Bol. Soc. Argent. Bot. Iv. 133.
1951
Amaranthus peruvianus (Schauer) Publ. Field Mus. Nat. Hist., Bot.
Standl. Ser. 13(2): 487. 1937
Amaranthus polychroa Raeusch. Nomencl. Bot. [Raeusch] ed. 3.
275. 1797
Amaranthus polyflagellus Spreng. In Hort. Argent. (1834). ex Moq.
in DC. Prod. Xiii. II. 266
Amaranthus polygamus L. = A. tricolor L. Cent. Pl. I. 32. 1755
Amaranthus polygonoides L. = A. lividus L. Pl. Jamaic. Pug. 27. 1759
= A. berlandieri (Moquin-Tandon)
Uline & W.L. Bray
Amaranthus polystachyus Willd. = A gracilis L. Sp. Pl. ed. 4 [Willdenow] 4(1):
385. 1805
Amaranthus powellii S. Watson = A. hypochondriacus L. var. powellii Proc. Amer. Acad. Arts 10: 347.
(S. Watson) Pedersen 1875
= A. chlorostachys Willd. var. powellii
(S. Watson) Priszter
= A. bracteotus Uline & W.L. Bray
= A. retroflexus L. var. powellii
(S. Watson) B. Boivin
Amaranthus powellii S. Watson subsp. Sida 19(4); 964. 2001
bouchonii (Thell.) Costea & Carretero
Amaranthus powellii S. Watson subsp. = A. bouchonii Thell. Nordic J Bot. 30(1): 13. 2012
cacciatoi (Aellen ex Cacciato) = A. bouchonii Thell. var. cacciatoi
Iamonico
Amaranthus praetermissus Brenan J.S. African Bot. 47(3): 478.
1981
Amaranthus pringlei S. Watson Proc. Amer. Acad. Arts 22: 476.
1887
Amaranthus probstii Thell. Repert. Spec. Nov. Regni Veg.
23: 271. 1926
(continued)
184 Appendices

Amaranthus prostratus T. Bastard = A. blitum L. Essai Fl. Maine-et-Loire 344.
1809
Amaranthus pseudogracilis (Thell.) Jahrb. Bochum. Bot. Vereins 1:
G.H. Loos. 117. 2010
Amaranthus pubescens Rydb. Bull. Torry Bot. Club 1912
xxxix. 313
Amaranthus pumilus Rafinesque Med. Repos. New York v. 360.
1808
Amaranthus purgans Hort. ex Moq. Prodr. [A.P. de Candolle] 13(2):
257. 1849
Amaranthus pycnostachys St.-Lag. Etude Fl. Ed. 8 [A. Cariot] 2:
697. 1889
Amaranthus pyramidalis Noronha Verh. Batav. Genootsch. Kunst.
5 (Art. 4): 7. 1790
Amaranthus quitensis Kunth Nov. Gen. Sp. [H.B.K.] 2: 194.
1818
Amaranthus recurvatus Desf. Tabl. Ecole. Bot. Ed. 3. (Cat. Pl.
Horti Paris) 39. 1829
Amaranthus retroflexus L. = A. powellii S. Watson Sp. Pl. 2: 991. 1753
= A. retroflexus var. salicifolius
I.M. Johnston
= A. bulgaricus Kov.
Amaranthus retroflexus var. powellii Naturaliste Carad. 93: 641. 1966
(S. Watson) B. Boivin
Amaranthus retroflexus var. = A. chlorostachys Willd. var. Phytologia 17: 70. 1968
pseudoretroflexus (Thell.) Boivin pseudoretroflexus
Amaranthus retroflexus var. rubricaulis = A. retroflexus L. f. rubricaulis Thell. In Asch & Graebn-Syn.
Thell. ex Probst. Mitteleur. Fl. [Ascherson &
Graebner] 5, Abt. 1: 260. 1914
A. retroflexus var. salicifolius J. Arnold Arbor. 25:157. 1944
I.M. Johnston
Amaranthus reverchonii (Uline & = A. blitoides S. Watson Nauchni Trudove Selskost Inst.
W.L. Bray) Kov. “Vasil Kolarov” 23: 49. 1978
Amaranthus rhombeus R. Br. Prodr. Fl. Nov. Holland. 414.
1810
Amaranthus rigidus Schult. ex Steud. Nomencl. Bot. [Steudel] ed.2.1:
70. 1840
Amaranthus rosengurttii Hunz. Kurtziana iii. 201. 1966
Amaranthus rotundifolius Herb. Par. Prodr. [A.P. de Candolle] 13(2):
ex Moq. 261. 1849
Amaranthus roxburghianus Nevski Trudy Bot. Inst. Akad. Nauk
S.S.S.R., Ser. 1, Fl. Sist. Vyssh.
Rast. 4: 311. 1937
Amaranthus roxburghianus var. J Bombay Nat. Hist. Soc. 73(1):
angustifolius (Moq.) N.C. Nair 61. 1976
Amaranthus roxburghianus var. J Bombay Nat. Hist. Soc. 73(1):
aschersonianus (Thell.) N.C. Nair 61. 1976
Amaranthus ruber E.H.L.Krause Beih. Bot. Centralbl., Abt. 2.
33(2): 479. 1915
Amaranthus rubescens Hort. ex Moq. Prodr. [A.P. de Candolle] 13(2):
257. 1849
(continued)
Appendices 185

Amaranthus rubricaulis Page & Hort. Prodr. [A.P. de Candolle] 13(2):
Angl.ex Moq. 267. nomen. 1849
Amaranthus ruderalis Koch ex moq. Prodr. [A.P. de Candolle] 13(2):
274. 1849
Amaranthus rudis J.D. Sauer Madrono 21(6): 1972
Amaranthus ruebelii Thell. Repert. Spec. Nov. Regni Veg.
24: 299, hybr. 1928
Amaranthus salicifolius Hort. Veitch Sauer (1950); Kirpicznikov
(1969); Wohlport & Mabry
(1968)
Amaranthus sanguineus L. Sp. Pl. ed. 2. 2: 1407. 1763
Amaranthus sanguinolentus Schrad. Prodr. [A.P. de Candolle] 13(2):
ex Moq. 267. 1849
Amaranthus scariosus Benth. Bot. Voy. Sulphur [Bentham]
158, t.51. 1846
Amaranthus scandens L.f. Suppl. Pl. 419. 1782
Amaranthus scleropoides Uline & Bot. Gaz. 19: 316. 1894
W.L. Bray
Amaranthus schinzianus Thell. Vierteljahrsschr. Naturf. Ges.
Zurich Ivii. 535. 1912
Amaranthus sclerantoides (Anderson) Kongl. Svenska Freg. Eugenix
Anderson Resa, Bot.2: 59. 1861
Amaranthus sylvestris Vill. Cat. Jard. Pl. Strasbourg iii.
1807
Amaranthus sparganiocephalum In Graebn. Syn. Mitteleur. Fl. V.
Thell. 312 (1914), in obs.
Amaranthus spathulatus Desf. ex Moq. Prodr. [A.P. de Candolle] 13(2):
276. 1849
Amaranthus speciosus Sims Sauer (1950)
Amaranthus spinosus L. Sp. Pl. 2: 991. 1753
Amaranthus spiratus Zipp. ex Span. Linnaea 15: 345. Nomen. 1841
Amaranthus splendens Hort. Vilm. Blumengartn., ed. 3. 1:
868, in Syn. 1895
Amaranthus squamulatus B.L. Rob. Proc. Amer. Acad. Arts xiiii. 32
(1907)
Amaranthus squarrulosus Uline & = Amblogyna squarrulosa A. Gray Bot. Gaz. 19: 270. 1894
W.L. Bray
Amaranthus standleyanus Parodi ex Darwiniana v. 339. 1941
Covas
Amaranthus strictus Ten. Syll. Pl. Fl. Neapol. 127. 1831
Amaranthus sylvestris Desf. = A. graecizans ssp. sylvestris Tabl. Ecole. Bot. 44, nom. Nud.
= A. blitum var. sylvestris 1804
= A. sylvestris Vill.
= A. graecizans
Amaranthus tamaulipensis Henrickson Sida 18(3): 808. 1999
Amaranthus taishanensis F. Z. Li. & Acta Phytotax. Sin. 19(1): 116.
C.K. Ni 1981
Amaranthus tamariscinus Nutt. = A. rudis J.D. Sauer Trans. Amer. Philos. Soc. Ser. 2,
5: 165. 1835
Amaranthus tarraconensis Sennen & Bull. Geogr. Bot. xxi. 124. 1911,
Pau
(continued)
186 Appendices

Amaranthus tenuiflorus Fisch. Hort. Prodr. [A.P. de Candolle] 13(2):
Hal. ex Moq. 267. 1849
Amaranthus tenuifolius Willd. = Mengea tenuifolia Sp. Pl. ed. 4 [Willdenow] 4(1):
381. 1805
Amaranthus tenuis Benth. Fl. Austral. 5: 216. 1870
Amaranthus thellungianus Nevski ex = A. graecizans ssp. thellungianus/ Fl. USSR, ed. Komarov vi. 365
Vassilcz = A. polygamus/A. blitum 1936; Townsend (1980)
var.polygonoides /A. polygonoides
Amaranthus thevenoei Degen & Thell. In Graebn. Syn. Mitteleur. Fl. V.
346. 1914. hybr.
Amaranthus thunbergii Moq. Prodr. [A.P. de Candolle] 13(2):
262. 1849
Amaranthus timeroyi Jord. ex Moq. Prodr. [A.P. de Candolle] 13(2):
259. 1849
Amaranthus torreyi Benth. ex Bot. California [W.H. Brewer]
S. Watson ii. 42. 1880
Amaranthus tortuosus Hornem. Hort. Hafn. Suppl. 107
Amaranthus tricolor L. Sp. Pl. 2: 989. 1753
A. tricolor L. var. acutus Saubhik Das Phytotaxa 88(2): 27.2013
A. tricolor L.var. tristis (Willd.) Companion to Chopra’s Gloss.
Mehrotra, Aswal & B.S. Bisht Indian Medicin. Pl. 9. 1987
Amaranthus tricolor splendens Kirpicznikov (1969)
Amaranthus tristis L. = A. tricolor L. Sp. Pl. 2: 989. 1753
= A. dubius Mart ex Thell
= A. tristis Moq.
= A. dubius
Amaranthus trivialis Rota. Giorn. Bot. Ital. Ii. II. 287. 1846
Amaranthus tuberculatus (Moq.) = Acnida tuberculatus Moquin- Madrono 13; 18. 1955
J.D. Sauer Tandon
= A. rudis J.D. Sauer
= A. tamariscinus var. tuberculate
(Moquin-Tandon) Uline & W.L. Bray
Amaranthus tucsonensis Henrickson Sida 18(3): 804. 1999
Amaranthus turcomanicus Gand. Bull. Soc. Bot. France 66: 222.
1919
Amaranthus undulatus R. Br. Prodromus Florae Novae
Hollandiae. 414. 1810
Amaranthus urceolatus Benth. Bot. Voy. Sulph. 158
Amaranthus ulinei Kov. = A. bigelovii Uline & W.L. Bray var. Nauch. Trud. Vissh Selskostop.
emarginatus Inst. Plovdiv 23(1): 49. 1978
Amaranthus velutina Spruce ex Proc. Amer. Acad. Arts 22: 366.
K. Schum 188
Amaranthus venulosus S. Watson Proc. Amer. Acad. Arts 17: 376.
1882
Amaranthus vernus Opiz ex Moq. Prodr. [A.P. de Candolle] 13(2):
267. 1849
Amaranthus verticillatus Sesse & Fl. Mexic. ed. 2. 217. 1894
Moc.
Amaranthus violaceus Hort. ex Moq. Prodr. [A.P. de Candolle] 13(2):
270. 1849
(continued)
Appendices 187

Amaranthus viridis L. = A. gracilis Desfontaines ex Poiret Sp. Pl. ed. 2. 2: 1405. 1763
Amaranthus viscidulus Greene = A. bracteosus Pittonia 3(19): 344. 1898
******
Amaranthus vulgatissimus Speg. = A. vulgatissimus auct. Anales Soc. Ci. Argent. 53: 281.
= A. standleyanus 1902
Amaranthus warnockii I.M. Johnst. J. Arnold Arbor. 25: 153. 1944
Amaranthus watsonii Standley = A. torreyi var. suffruticosum Uline & Bull. Torrey Bot. Club 41:
W.L. Bray 505.1914
Amaranthus wrightii S.Watson Proc. Amer. Acad, Arts 12: 275.
1877
Amaranthus zanensis Horn. ex Moq. Prodr. [A.P. de Candolle] 13(2):
267. 1849
References
Abbott RJ (1992) Plant invasions, interspecific hybridiza- Arora RK, Nayar ER (1984) Wild relatives of crop plants
tion and the evolution of new plant taxa. Trends Ecol in India. NBPGR science monograph no. 7. New
Evol 7:401–405 Delhi, India, 90 p
Abdullah MA, Abdullah AA, Safwat OK et al (2011) Arora RK et al (1991) Plant diversity in the Indian gene
Influence of storage conditions on seed quality and centre. In: Plant genetic resources conservation and
longevity of four vegetable crops. Agric Environ Sci management – concepts and approaches. IBPGR
11(3):353–359 Regional Office for South and Southeast Asia, New
Adhikary D, Pratt DB (2015) Morphological and Delhi, pp 25–54
Taxonomic analysis of the weedy and cultivated Arus P, Shields CR, Orton TJ (1985) Application of iso-
Amaranthus hybridus species complex. Syst Bot zyme electrophoresis for purity testing and cultivar
40(2):604–610 identification of F1 hybrids of Brassica napus.
Aellen P (1959) Amaranthus L. In: Hegi G (ed) Illustrierte Euphytica 3493:651–657
Flora von Mitteleuropa, 2nd edn., vol 3, Part 2. Carl Aufhammer W, Czuczorova D, Kaul Kruse M (1998)
Hanser Verlag, Munchen, pp 465–516 Germination of grain amaranth (Amaranthus hypo-
Aellen P (1961) Die Amaranthaceen Mitteleleuropus. chondriacus × A. hybridus): effects of seed quality,
Carl Hanser Verlag, Munchen temperature, light, and pesticides. Eur J Agron
Ahrens WH, Stoller EW (1983) Competition, growth rate, 8(1–2):127–135
and CO2 fixation in triazine-susceptible and -resistant Avise C, Hamrick JL (eds) (1996) Conservation genetics:
smooth pigweed (Amaranthus hybridus). Weed Sci case studies fromnature. Chapman and Hall, New York
31:438–444 Awasthi AK, Nagaraja GM, Naik GV et al (2004) Genetic
Ahrens WH, Wax LM, Stoller EW (1981) Identification of diversity and relationships in mulberry (genus Morus)
triazine resistant Amaranthus spp. Weed Sci as revealed by RAPD and ISSR marker assays. BMC
29:345–348 Genet. doi:10.1186/1471-2156-5-1
Ali MA, Nawab NN, Rasool G et al (2008) Estimates of Ayiecho PO (1986) Quantitative studies in two grain ama-
variability and correlations for quantitative traits in ranth populations using two selection methods.
Cicer arietinum L. J Agric Soc Sci 4(4):177–179 Dissertation Abstracts, International, B (Sciences and
Alvarez-Jubete L, Wijngaard H, Arendt EK et al (2010) Engineering) 46(7):2189B
Polyphenol composition and in vitro antioxidant activ- Ayorinde FO, Ologunde MO, Nana EY et al (1989)
ity of amaranth, quinoa, buckwheat and wheat as Determination of fatty acid composition of Amaranthus
affected by sprouting and Baking. Food Chem species. J Am Oil Chem Soc 66:1812–1814
119:770–778 Azam-Ali S, Battcock M (2002) Fermented fruits and
Ames O (1939) Economic annuals and human cultures. vegetable. A global perspectives FAO Agricultural
Botanical Museum of Harvard University, Cambridge Services Bulletin
Anderson E, Hubricht I (1938) Hybridization in Babatola J, Awoderu JB (1986) Screening Amaranthus
Tradescantia. III. The evidences for introgresive germplasm against root-knot nematodes, Meloidogyne
hybridization. Am J Bot 25:396–402 incognita. Test Agrochem Cult 7:140–141
Anonymous (1994) Amaranth; modern prospects for an Badami S, Gupta MK, Suresh B (2003) Antioxidant activ-
ancient crop. National Research Council. National ity of the ethanol extract of Striga orobanchioides.
Academy Press, Washington, DC J Ethno Pharmacol 85:227–230
Arora RK (1985) Genetic resources of less known culti- Baker W, Ollis WD (1961) Recent developments in the
vated food plants. NBPGR science monograph 10 chemistry of natural phenolic compounds. Pergamon
Arora RK (1987) Ethbotany and its role in domestication Press, New York
and conservation of native plant resources. In: Jain SK Baltensperger DD, Weber LE, Nelson LA (1992)
(ed) Manual of ethnobotany. Scientific Publication, Registration of ‘Plainsman’ grain Amaranths. Crop Sci
Jodhpur, pp 94–102 32:1510–1511

S. Das, Amaranthus: A Promising Crop of Future, DOI 10.1007/978-981-10-1469-7
190 References
Bansal GL (1996) Physiological investigation on grain and Bennici A, Schiff S, Bovelli R (1992) In vitro culture of
green amaranths (Amaranthus spp.) in relation to produc- species and varieties of four Amaranthus species.
tivity under mid hill condition, Final report. Department Euphytica 62:181–186
of Plant Physiology, College of Basic Sciences, Himachal Berjack P, Villiers TA (1972) Ageing in plant embryos.
Prdesh Krishi Vishvavidyalaya, Palampur II. Age induced damage and its repair during early ger-
Barba de la Rosa AP, Fomsgaard IS, Larsen B et al (2009) mination. New Phytol 71:135–144
Amaranth (Amaranthus hypochondriacus) as an alter- Bernhardt S, Schlich E (2006) Impact of different cooking
native crop for sustainable food production: phenolic methods on food quality: retention of lipophilic vita-
acids and flavonoids with potentialimpact on its nutra- mins in fresh and frozen vegetables. J Food Eng
ceutical quality. J Cereal Sci 49:117–121 77:327–333
Bardini M, Lee D, Donini P et al (2004) Tubulinbased Bertin RI (1982) The evolution and maintenance of andro-
polymorphism (TBP): a new tool, based on function- monoecy. Evol Theory 6:25–32
ally relevant sequences, to assess genetic diversity in Betschart AA, Irving DW, Shepherd AD et al (1981)
plant species. Genome 47(2):281–291 Amaranthus cruentus: milling characteristics, distri-
Barker LA, Duarte PR (1998) Retrogradation of amaranth bution of nutrients within seed components and the
starch at different storage temperatures and the effects effects of temperature on Nutritional quality. J Food
of salt and sugars. Cereal Chem 75(3):308–314 Sci 46:1181–1187
Barralis G, Chadoeuf R, Lonchamp JP (1988) Longevity Bhatia AL, Jain M (2003) Amaranthus paniculatus
of annual weed seeds in cultivated soil. Weed Res (Linn.) improves learning after radiation stress.
28:407–418 J Ethnopharmacol 85:73–79
Barton NH (2001) The role of hybridization in evolution. Blatchley WS (1912) The Indiana weed book. Nature
Mol Ecol 10:551–568 Publishing Company, Indianapolis
Baskin JM, Baskin CC (1998) Seed dormancy and germi- Bonasora MG, Poggio L, Greizerstain EJ (2013)
nation in the rare plant species Amaranthus pumilus. Cytogenetic studies in four cultivated Amaranthus
Castanea 63:493–494 (Amaranthaceae) species. Com Cytogenet
Basu A, Ghosh M, Meyer R et al (2004) Analysis of 7(1):53–61
genetic diversity in cultivated jute determined by Bradova J, Matejova E (2008) Comparison of the results
means of SSR markers and AFLP profiling. Crop Sci of SDS PAGE and chip electrophoresis of wheat stor-
44:678–685 age proteins. Chromatographia 67:583–588
Bate-Smith EC, Lerner NH (1954) Leucoanthocyanins. Braglia L, Manca A, Mastromauro F et al (2010) cTBP: a
(2) Systematic distribution of Leucoanthocyanins in successful Intron Lenght Polymorphism (ILP)-based
leaves. Biochem J 58:126–132 genotyping method targeted to well defined experi-
Bawa KS, Beach JH (1981) Evolution of sexual systems mental need. Diversity 2:572–585
in flowering plants. Ann Mo Bot Gard 68:254–274 Brenan JPM (1961) Amaranthus in Britain. Watsonia
Beck G (1909) Reichenb. Icon. Fl. Germanicae et 4:261–280
Helveticae 24:182 Brenan JPM, Townsend CC (1980) Proposal to reject
Becker R, Wheeler EL, Lorenz K et al (1981) A composi- Amaranthus blitum L. under art.69 in favor of A. livi-
tional study of amaranth grain. J Food Sci 46:1175 dus L. Taxon 29:695–696
Becker HC, Damgaard C, Karlsson B (1992) Brenner DM (1990) Amaranth catalogue. USDA Regional
Environmental variation for out-crossing rate in rape- Plant Introduction Station, Ammes
seed. Theor Appl Genet 84:303–306 Brenner DM (1993) Hybrid seeds for increased amaranth
Been MM, Fellers DA (1982) Composite breads in grain yield. Legacy 6:9–11
Bolivia; technical aspects. In: Proceedings 7th World Brenner DM, Hauptli H (1990) Seed shattering control
Cereals and Bread Congress, Prague with indehiscent utricles in grain amaranths. Legacy
Behera B, Patnaik SN (1974) Chemotaxonomic studies in 3:2–3
the family Amaranthaceae. Cytologia 39:121–131 Brenner DM, Widrlechner MP (1998) Amaranthus seed
Behera B, Patnaik SN (1975) Induced polyploidy in germination in plastic tents in green houses. FAO/
Amaranthus hypochondriacus and Amaranthus dubius IPGRI Plant Gen Resour Newsl 116:1–4
Mart ex Thell. Cytologia 40:157–168 Brenner DM, Baltensperger DD, Kulakow PA et al (2000)
Behera B, Patnaik SN (1982) Genome analysis of Genetic resources and breeding in Amaranthus. In:
Amaranthus dubius Mart. ex Thell.through the study Janick J (ed) Plant breeding reviews, vol 19. Wiley,
of Amaranthus spinosus X A. dubius hybrids. New York, pp 227–285
Cytologia 47:379–389 Brenner DM, Johnson WG, Sprague C et al (2013) Crop
Behera B, Tripathy A, Patnaik SN (1974) Histological weed hybrids are more frequent for the grain amaranth
analysis of colchicines-induced deformities and cyto- ‘Plainsman’ than for D136-1. Genet Resour Crop Evol
chimeras in Amaranthus caudatus and A. dubius. 60:2201–2205
J Heridity 65:179–184 Bressani R (1993) Amaranth. In: Macrae R, Robinson
Bejosano FP, Corke H (1998) Protein quality evaluation RK, Sadler MJ (eds) Encyclopaedia of food science.
of Amaranths whole meal flours and protein concen- Food technology and nutrition, vol 1. Academic,
trate. J Sci Food Agric 76:100–106 London, pp 135–140
References 191
Bressani R, Garcia-Vela LA (1990) Protein fractions in in Australasian Wurmbea (Colchicaceae). Int J Plant
Amaranth grain and their chemical characterization. Sci 169:141–156
J Agric Food Chem 38(5):1205–1209 Ceccarelli S, Grando S, Baum M et al (2004) Breeding for
Bressani R, Gorizleg JM, Elias LG et al (1987) Effect of drought resistance in a changing climate. In: Rao SC,
fertilizer application on yield, protein and fat content, Ryan J (eds) Challanges and strategies for dryland
and protein quality of raw and cooked grain of three agriculture. Madison, pp 167–190
amaranth species. Qual Plantarum Plant Foods Hum Cecilia TNE (2009) Transgenic approach to improve the
Nutr 37(1):59–67 nutritional quality of wheat (Triticum aestivum L.). Ph.
Bretting PK, Widrlechner MP (1995) Genetic markers and D thesis, Agricultural Research Institute of HAS,
plant genetic resource management. Plant Breed Rev Mastonvasar
13:11–86 Chakraborty AG (1977) Effects of temperature shift on
Breviario D, Baird WV, Sangoi S et al (2007) High poly- weed seed germination. Castanea 42:279–285
morphism and resolution in targeted fingerprinting Chakraborty S, Chakrborty N, Datta S (2000) Increased
with combined β-tubulin introns. Mol Breed nutritive value of transgenic potato by expressing non-
20(3):249–259 allerganic seed albumin gene from Amaranthus hypo-
Breviario D, Gianì S, Ponzoni E et al (2008) Plant tubulin chondriacus. Proc Natl Acad Sci U S A
intronics. Cell Biol Int 32(5):571–573 97:3724–3729
Britton NL, Brown A (1896) An illustrated flora of the Chakraborty S, Chakraborty N, Agrawal L et al (2010)
Northern United States, Canada and the British Next-generation protein-rich potato expressing the
Possessions, vol 2. Charles Scribner’s Sons, seed protein gene AmA1 is a result of proteome rebal-
New York ancing in transgenic tuber. Proc Natl Acad Sci
Britton NL, Brown A (1913) An illustrated flora of the 107(41):17533–17538
Northern United States, Canada and the British pos- Chakraborty N, Ghosh R, Ghosh S et al (2013) Reduction
sessions. Scribner, New York of Oxalate levels in Tomato fruit and consequent meta-
Brumitt R (1984) Report of the committee for spermato- bolic remodelling following over expression of a fun-
phytes 27. Taxon 33:297–301 gal Oxalate Decarboxylase. Plant Physiol
Buhler DD, Hartzler RG (2001) Emergence and persis- 162(1):364–378
tence of seed of Velvetleaf, Common Waterhemp, Chamberlain JR (1998) Isozyme variation in Calliandra
Woolly Cupgrass, and Giant Foxtail. Weed Sci calothyrsus (Legumonosae): it’s implication for spe-
49:230–235 cies delimitation and conservation. Am J Bot
Busi R, Yu Q, Barrett-Lennard R et al (2008) Long dis- 85:37–47
tance pollen-mediated flow of herbicide resistance Chan KF (1996) M.Phil. thesis. University of HongKong,
genes in Lolium rigidum. Theor Appl Genet Hong Kong
117:1281–1290 Chan KF, Sun M (1997) Genetic diversity and relation-
CAD (2009) Committee for Agricultural Development. ships detected by isozyme and RAPD analysis of crop
Iowa State University (online), Ames. http://www.ag. and wild species of Amaranthus. Theor Appl Genet
iastate.edu/centers/cad/cadamaranth.html 95:865–873
Cai Y, Sun M, Wu H et al (1998) Characterization and Chao WS, Horvath DP, Anderson JV et al (2005) Potential
quantification of betacyanin pigments from diverse model weed to study genomics, ecology and physiol-
Amaranthus species. J Agric Food Chem ogy in the 21st century. Weed Sci 53:929–937
46:2063–2070 Chaturvedi M, Dutta K, Pal M (1997) Cytopalynology of
Campbell JY (1987) Does saving anticipate declining Amaranthus L. and Chenopodium L. – the two panto-
labor income? An alternative test of the permanent porate taxa. Feddes Repertorium 108(5–6):325–333
income hypothesis, Econometrica. Economet Soc Chauhan GS, Eskin NAM, Tkachuk R (1992) Nutrients
55(6):1249–1273 and antinutrients in quinoa seed. Cereal Chem
Caperuto LC, Amaya-Farfan J, Camargo CRO (2001) 69:85–88
Performance of Quinia (Chenopodium quinoa Willd.) Chawla B, Bernatzky R, Liang W et al (1997) Brekdown
flour in the manufacture of gluten-free spaghetti. J Sci of self-incompatibility in tetraploid Lycopersicon
Food Agric 81:95–101 peruvianum: inheritance and expression of S-related
Cardina J, Norquay JM, Stinner BA et al (1996) protein. Theor Appl Genet 95:992–996
Postdispersal predation of Velvetleaf (Abutilon theo- Cheeke PR, Carlsson R, Kohler GO (1981) Nutritive
phrasti) seeds. Weed Sci 44:534–539 value of leaf protein concentrates prepared from
Carlsson R (1982) Leaf protein concentrates from plant Amaranthus species. Can J Anim Sci 61(1):199–204
sources in temperate regions: In: Telek L, Graham HD Chepil WS (1946) Germination of weed seeds II. The
(eds) Leaf protein concentrates. AVI Technical Book, influence of tillage treatment on germination. Sci
Inc, West Port, pp 52–80 Agric 26(8):347–357
Carretero JL (1985) Consideraciones sobre las amaranta- Cheung AY, Bogord L, Montagu MV et al (1988)
ceas ibericas. An Jard Bot Madr 41(2):271–286 Relocating a gene for herbside tolerance: a Chloroplast
Case AI, Graham SW, Macfarlane TD (2008) A phyloge- gene is converted into a nuclear gene. Proc Natl Acad
netic study of evolutionary transition in sexual system Sci U S A 85:391–395
192 References
Coetzer E, Al-Khatib K, Peterson DE (2002) Glufosinate Covas G (1992) Clove para la identification de los ama-
efficacy on Amaranthus species in glufosinate- resis- rantes cultivados y cspecies silvestres utilizable como
tant Soybeans (Glycine max). Weed Technol hortalizas o forrajeras: 9–12. – Amarantos, novedades
16:326–331 e informaciones no. 12. Fac de Agronomia, INTA
Colmenarez de Ruiz AS, Bressani R (1990) Effect of ger- Anguil. Santa Rosa, La Pampa. –Argentina
mination on the chemical composition and nutritive Cramer C (ed) (1988) Montana releases new amaranth
value of amaranth grain. Cereal Chem 67(6):519–522 line. Amaranth Today 4(2–3):6
Coons MP (1975) The genus Amaranthus in Ecuador. Crawford DJ (1983) Phylogenetic and systematic infer-
Ph.D thesis, Indiana Universitiy, Bloomington, USA ences from electrophoretic studies, part A. In: Tanksley
Coons MP (1978) The status of Amaranthus hybridus L. S, Orton T (eds) Isozyme in plant genetics and breed-
in South America, part 2. The taxonomic problem. ing. Elsevier Publication, Amstardam, pp 257–287
Ciencia y Naturaleza (Quito) 19:66–71 Crocker W (1916) Mechanics of dormancy in seeds. Am
Coons MP (1982) Relationships of Amaranthus caudatus. J Bot 3:99–120
Econ Bot 36(2):129–146 Cross H (1930–1933) Laboratory germination of weed
Copeland LO, McDonald MB (1995) Principles of seed seed. In: Proceedings of the association of official seed
science and technology. Mc Millan Publishing analyst of North −128
Company, New York Cuenoud P, Savolainen V, Chatrou LW et al (2002)
Corke H, Wu H, Yue S et al (1997) Developing specially Molecular phylogenetics of Caryophyllales based on
starches from new crops. In: Campbell GM, Webb C, nuclear 18S rDNA and Plastid rbcL, atpB and matK
McKee SL (eds) Cereals: novel use and processes. DMA sequences. Am J Bot 89(1):132–144
Plenum Press, New York, pp 90–102 Culpepper AS, Grey TL, Vencill WK et al (2006)
Cornille A, Gladieux P, Smulders MJM et al (2012) New Glyphosate-resistant Palmer amaranth (Amaranthus
insight into the history of domesticated apple: second- palmeri) confirmed in Georgia. Weed Sci 54:620–626
ary contribution of the European wild apple to the Dangi R, Lagu M, Choudhary L et al (2004) Assessment
genome of cultivated varieties. PLoS Genet 8, e1002703 of genetic diversity in Trigonella foenum-graecum and
Costea M (1997) Morphology of seed in some species of Trogonella caerulea using ISSR and RAPD markers.
the genus Amaranthus L. Acta Horti Botanici BMC Plant Biol 4:13
Bucurestiensis (University of Bucharest) 53:24–37 Darlington CD, Janaki Ammal EK (1945) Chromosome
Costea M (1998a) Amaranthus L. subgenus Albersia atlas of cultivated plants. George Allen and Unwin
(Kunth) Gren. & Godr.in Romania. Revue Roumaine Ltd, London, p 397
de Biologie (Romanian Academy of Science) Das S (2012a) Systematics and taxonomic delimitation of
43:95–112 vegetatable, grain and weed amaranths: a morphologi-
Costea M (1998b) Monograph of the genus Amaranthus cal and biochemical approach. Genet Resour Crop
L. in Romania. Ph.D thesis, University of Bucharest, Evol 59:289–303
College of Biology, Bucharest, Romania Das S (2012b) Taxonomical observation on the grain ama-
Costea M, DeMason DA (2001) Stem morphology and ranths and new varieties of Amaranthus cruentus
anatomy in Amaranthus L. (Amaranthaceae) – taxo- (Amaranthaceae). Nor J Bot 30:412–420
nomic significance. J Torrey Bot Soc 128(3):254–281 Das S (2013) Infraspecific variability of Amaranthus tri-
Costea M, Tardif FJ (2003a) Conspectus and notes on the color (Amaranthaceae) in India with a new variety.
genus Amaranthus in Canada. Rhodora 105:260–281 Phytotaxa 88(2):25–30
Costea M, Tardif FJ (2003b) The bracteoles in Amaranthus Das S (2015) Amaranthus parganensis (Amaranthaceae),
(Amaranthaceae): their morphology, structure, func- a new species from West Bengal, India. Novon
tion, and taxonomic significance. Sida 20:969–985 23:406–410
Costea M, Tardif FJ (2003c) The biology of Canadian Das S, Iamonico D (2014) Amaranthus bengalense
weeds 126. Amaranthus albus L., A. blitoides (Amaranthaceae) a new species from India with taxo-
S. Watson and A. blitum L. Can J Plant Sci nomic notes on A. blitum aggregate. Phytotaxa
83:1039–1066 181(5):293–300
Costea M, Sanders A, Waines G (2001a) Preliminary Das S, Mukherjee KK (1995) Biochemical studies on
results towards revision of the Amaranthus hybridus Ipomoea pollen to understand species homology.
species complex (Amaranthaceae). Sida 19:931–974 Grana 34:332–337
Costea M, Waines G, Sanders A (2001b) Notes on some Davis L, Kuttan G (2001) Effects of Withania somnifera
little known Amaranthus taxa (Amaranthaceae) in the on DMBA induced carcinogenesis. J Ethnopharmacol
United States. Sida 19:975–992 75:165–168
Costea M, Brenner DM, Tardif FJ et al (2006) Delimitation de Candolle AP (1849) Prodromus Systematis Naturalis
of Amaranthus cruentus L. and Amaranthus caudatus Regin Vegetabilis 13(2):262
L. using micromorphology and AFLP analysis: an de Candolle A (1883) Origine des plantes cultivees.
application in germplasm identification. Genet Resour G. Bailliere et cie, Paris
Crop Evol 53(8):1625–1633 De Jong TJ, Klinkhamer PGL (1994) Plant size and repro-
Covas G (1991) Nuevo cultivar de amaranto granifero. ductive success through female and male function. J
Amarantos Novedades e informaciones 7:4–5 Ecol 82:399–402
References 193
Delano-frier JP, Aviles-Arhaut H, Casarrubias-Castillo K the third Amaranth Conference. Rodale Press, Inc,
et al (2011) Trancriptomic analysis of grain amaranth Emmaus, PA
(Amaranthus hypochondriacus) using 454 pyrose- Dziwulska-Hunek A, Kornarzyński K (2009) Kiełkowanie
quencing: comparison with A. tuberculatus, expresión nasion amarantusa odmian Aztek i Rawa w różnych
profiling in stem and in response to biotic and abiotic temperaturach. Acta Sci Pol Technica Agraria
stress. BMC Genomics 12:363 8(1–2):3–10
Deno N (1993) Seed germination, theory and practice, Dzunkova M, Janovska D, Hlasna-Cepkova P et al (2011)
2nd edn. 139 Lenor Drive, State College, PA Glutelin protein fraction as a tool for clear identifica-
Deutsch J (1977) Genetic variation of yield and nutrition of Amaranth accessions. J Cereal Sci
tional value in several Amaranthus species used as a 53(2):198–205
leafy vegetable. Ph.D. disscussion, Cornell University, Edelstein S, Sharlin J (2009) Life cycle nutrition. Jones
Ithaca, NY, USA and Bartlett Publishers, Sudbury
Devadas VS, Gopalakrishnan PK, Peter KV (1984) Egley GH, Chandler JM (1978) Germination and viability
Breeding for low antinutrient factor in vegetable ama- of weed seeds after 2.5 years in a 50-year buried seed
ranths. Amaranth Newsl 2:2–3 study. Weed Sci 26(3):230–239
Devadas VS, Gopaiakrishnan PK, Peter KV (1992) Egley GH, Chandler JM (1983) Longevity of weed seeds
Genetic divergence in vegetable amaranths. South after 50 years in the Stoneville 50-year buried-seed
Indian Hortic 40(1):16–20 study. Weed Sci 31:264–270
Dieleman A, Hamill AS, Fox GC et al (1996) Decision Egley GH, Williams RD (1990) Decline of weed seeds
rules for postemergency control of pigweed and seedling emergence over five years as affected by
(Amaranthus spp) in Soybean (Glycine max). Weed soil disturbances. Weed Sci 38:504–510
Sci 44:126–132 El-Aydam M, Burki HM (1997) Biological control of
Dodak L, Modhir AA, Halasova G et al (1994) Importance noxious pigweed in Europe: a literature review of the
and utilization of amaranth in food industry. Part 1. insect species associated with Amaranthus spp. world-
Characteristic of grain and average chemical constitu- wide. Biocontrol News Inf 8:11–20
tion of whole amaranth flour. Nahrung Food El-Esawi M (2008) M.Sc. thesis. Tanta University, Faculty
38:378–381 of Science, Botany Department
Dodak L, Modhir AA, Buchtova V et al (1997) Importance Eliasson U (1988) Floral morphology and taxonomic
and utilization of amaranth in food industry. 2. relation among genera of Amaranthaceae in the new
Composition of amino acid and fatty acids. Nahrung world and the Hawaii islands. J Linn Soc Bot
41:108–110 96:235–283
Dorken ME, Friedman JE, Barrett SCH (2002) The evolu- Ellis RH, Roberts EH (1980) Improved: equation for pre-
tion and maintenance of monoecy and dioecy in diction of seed longevity. Ann Bot 45:13–80
Sagittaria latifolia. Evolution 56:31–41 Ellstrand NC, Prentice HC, Hancock JF (1999) Gene flow
Downey SR, Katz-Downey DS, Cho KJ (1997) and introgression from domesticated plants into their
Relationship in the Caryophyllales as suggested by wild relatives. Annu Rev Ecol Syst 30:539–563
phylogenetic analysis of partial chloroplast ORF 2280 El-Sharkawy MA, Loom’s RS, Williams WA (1968)
homolog sequence. Am J Bot 84:253–273 Photosysthetic and respiratory exchange of carbón
Downie SR, Palmer JD (1994) A chloroplast DNA phy- dioxide by leaves of the grain grain amaranths. J Appl
logeny of the Caryophyllales based on structural and Ecol 5(1):243–251
inverted repeat restriction sites variation. Syst Bot Endress PK (2001) The flowers in extant basal angio-
19:236–252 sperms and inferences on ancestral flowers. Int J Plant
Doyle JA (1998) Phylogeny of vascular plants. Annu Rev Sci 162:1111–1140
Ecol Syst 29:567–599 Engel KH, Frenzel TH, Miller A (2002) Current and
Drzewiecki J (2001) Similarities and differences between future benefits from the use of GM technology in food
Amaranthus species and cultivars and estimation of production. Toxicol Lett 127:329–336
outcrossing rate on the basis of electrophoretic separa- Erdtman G (1952) Pollen morphology and plant
tions of urea-soluble seed proteins. Euphytica taxonomy, Angiosperms. Almquist Wiksell,
119:279–287 Stockholm
Drzewiecki J, Delgado-Licon E, Haruenkit R et al (2003) Erdtman G (1966) Pollen morphology and plant taxon-
Identification and differences of total proteins and omy, Angiosperms. Alquimist & Wiksell, Stokholm
their soluble fractions in some Pseudocereals based on Espitia RE (1986) Caracterizacion y evaluacion prelimi-
electrophoretic patterns. J Agric Food Chem nar de germoplasma de Amaranthus spp. Thesis
51(26):7798–7804 (Ingeniero Agronomo) Universisdad Autonoma
Duggleby RG, Pang SS (2000) Acetohydroxy acid syn- Agraria “Antonio Narro”, Chapingo, Mexico
thase. J Biochem Mol Biol 33:1–36 Espitia RE (1994) Breeding of grain amaranth. In:
Dumortier B-CJ (1827) Florula Belgica, operis majoris Peredes-Lopez O (ed) Amaranth biology, chemistry
prodromus auctore. Casterman, Tourney and technology. CRC Press, Boca Raton, pp 23–38
Duriyaprapam S (1986) Study on grain amaranth Fang X, Jin W, Liang Z et al (1996) Electron- microscopic
production in northeast Thailand. In: Proceedings of study of microsporogenesis in male sterile and male
194 References
fertile grain Amaranths (Amaranthus hypochondria- resistant wheat hybridization under field conditions.
cus L.). Dev Reprod Biol 5:55–61 Weed Sci 56:32–36
FAO (1988) Traditional food plants, FAO food and nutri- Gaines TA, Zhang W, Wang D et al (2010) Gene amplifi-
tion paper 62. FAO, Rome cation confers glyphosate resistance in Amaranthus
FAO (1996a) Global plan of action for the conservation palmeri. Proc Natl Acad Sci U S A 107:1029–1034
and sustainable use of plant genetic resources for food Gaines TA, Ward SH, Bukun B et al (2012) Interspecific
and agriculture. FAO, Rome hybridization transfers a previously unknown glypho-
FAO (1996b) Report on the State of the World’s Plant sate resistance mechanism in Amaranthus species.
Genetic Resources for Food and Agriculture. Prepared Evol Appl 5(1):29–38
for the international technical conference on plant Gajdošová A, Libiaková G, Ostrolucká MG et al (2008)
genetic resources, Leipzig, Germany, 17–23 June Mutation breeding in selected Amaranthus spp. In:
1996. FAO, Rome, Italy. Available at http://www.fao. Libiakova G, Gajdosova, A (eds) Absatracte of the 5th
org/ag/AGP/AGPS/PGRFA/pdf/swrfull.pdf. Accessed International symposium of the European Amaranth
26 Dec 2008 Association on Amaranth – Plant for the Future,
FAO (1996c) Global plan of action for the conservation Instutute of Plant Genetics and Biotechnology SAS,
and sustainable utilization of plant genetic resources Nitra, Slovak Republic, 9–14 November 2008,
and the Leipzig methodologies for generating variabil- pp 93–94
ity. Part 1: use of genetic resources in plant breeding Ghafoor A, Ahmadl Z (2003) Exploitation of (vigna
125 Declaration, adopted by the International mungo (L.) Heeper) germplasm using multivariate
Technical Conference on Plant Genetic Resources, analysis based on agronomic trait. Pak J Bot
Leipzig, Germany, 17–23 June 1996. FAO, Rome. 35:187–196
Available at: ftp://ftp.fao.org/ag/cgrfa/GS/gpaE.pdf. Ghorbani R, Seel W, Leifert C (1999) Effects of environ-
Accessed 26 Dec 2008 mental factors on germination and emergency of
FAO (2005) The state of food insecurity in the world. Amaranthus retroflexus. Weed Sci 48:505–510
FAO, Rome Girija K, Lakshman K, Chandrika U et al (2011) Anti-
Feine-Dudley LB (1980) A provisional key some edible diabetic and anti-cholesterolemic activity of methanol
species of the family Amaranthaceae. In: Grubben extracts of three species of Amaranthus. Asian Pac
GJH, van Sloten DH (eds) Genetic resources of J Trop Biomed 1(2):133–138
Amaranths. The International Board for Plant Genetic Gleiser G, Verdu M (2005) Repeated evolution of dioecy
Resources, Rome, pp 35–47 from androdioecy in Acer. New Phytol 165:633–640
Filias FR, Gaulliez A, Guedes M (1980) Amaranthus bli- Gomez-Pando L, Eguiluz A, Jimenez J et al (2009) Barley
tum vs. A. lividus (Amaranthaceae). Taxon (Hordeum vulgare) and Kiwicha (Amaranthus cauda-
29:149–150 tus) improvement by mutation induction in Peru. In:
Fitterer SA, Johnson BL, Schneiter AA (1996) Grain ama- Shu QY (ed) Induced plant mutation in the genomics
ranth harvest timeliness in Eastern North Dakota. In: Era. Food and Agriculture Organization of the United
Janick J (ed) Progress in new crops. ASHS Press, Nations, Rome, pp 330–332
Alexandria, pp 220–223 Gorinstein S, Moshe R, Greene LJ (1991) Evaluation of
Flores HE, Thier A, Galston W (1982) In vitro culture of four Amaranthus species through protein electropho-
grain and vegetable Amaranths (Amaranthus spp.). retic patterns and their amino acid composition.
Am J Bot 69(7):1049–1054 J Agric Food Chem 39(5):851–854
Foy CD, Campbell TA (1984) Differential tolerance of Gorinstein S, Zemser M, Paredes-Lopez O (1996)
Amaranthus strains to high levels of aluminium and Structural stability of globulins. J Agric Food Chem
manganese in acid soils. J Plant Nutr 71:1365–1388 44:100–105
Franssen AS, Skinner DZ, Al-Khatib K et al (2001a) Gorinstein S, Zemser M, Fliess A et al (1998)
Pollen morphological differences in Amaranthus spe- Computational analysis of the amino acid residue
cies and interspecific hybrids. Weed Sci 49:732–737 sequences of Amaranth and some other proteins.
Franssen AS, Skinner DZ, Al-Khatib K et al (2001b) Biosci Biotechnol Biochem 62(10):1845–1851
Interspecific hybridization and gene flow of ALS Gorinstein S, Jaramillo NO, Medina OJ et al (1999)
resistance in Amaranthus species. Weed Sci Evaluation of some cereals, plants and tubers through
49:598–606 protein composition. J Protein Chem 18:687–693
Frisch M, Bohn M, Melchinger AE (1999) Minimum Gorinstein S, Pawelzik E, Delgado-Licon E et al (2002)
sample size and optimal positioning of flanking mark- Characterization of pseudocereals and cereals proteins
ers in marker assisted backcrossing for transfer of a by protein and amino acid analyses. J Sci Food Agric
target gene. Crop Sci 39:967–975 82(8):886–891
Funke M (2011) Evaluation of nutrient contents of Gornal RJ, Bohm BA (1980) Flavonoid and taxonomy of
Amaranth leaves prepared using different cooking Boykinia and related genera. Can J Bot
methods. Food Nutr Sci 2:249–252 58:1768–1779
Gaines TA, Henry WB, Byrne PF et al (2008) Jointed Goss WL (1924) The vitality of buried seeds. J Agric Res
goatgrass (Aegilops cylindrica) by imidazolinone- 29(7):349–362
References 195
Grant WF (1959a) Cytogenetic studies in Amaranthus J (ed) Perspectives on new crops and new uses. ASHS
I. Cytological aspects of sex determination in dioe- Press, Alexandria
cious species. Can J Bot 37:413–417 Gupta VK (1986) Grain amaranths in Kenya. In:
Grant WF (1959b) Cytogenetic studies in Amaranthus Procedings of the third Amaranth conference. Rodale
II. Natural interspecific hybridization between A. Press, Inc, Emmaus
dubius and A. spinosus. Can J Bot 37:1063–1070 Gupta VK, Gudu S (1990) Inheritance of some morpho-
Grant WF (1959c) Cytogenetic studies in Amaranthus logical trait in grain amaranths. Euphytica 46:79–84
III. Chromosome numbers and phylogenetic aspects. Gupta VK, Gudu S (1991) Interspecific hybrids and pos-
Can J Genet Cytol 1:313–328 sible phylogenetic relations in grain amaranths.
Gravis A, Constantinesco C (1907) Contribution a l’ anat- Euphytica 52:33–38
omie des Amaranthaceae. Archieves de I’Institut Guttieri MJ, Eberlein CV, Mallory-Smith CA et al (1996)
Botanique de I’Universite de Liege, pp 1–65 Molecular genetics of target site resistance to acetolac-
Greizerstein EJ, Poggio L (1992) Estudios citogenetico de tate synthase inhibiting herbicide. In: Brown TM (ed)
seis hibridos inter-especificos de Amaranthus. Molecular genetics and evolution of pesticide resis-
Darwiniana 31:159–165 tance. American Chemical Society, Washington, DC,
Greizerstein EJ, Poggio L (1994) Karyological studies in pp 10–16
grain amaranths. Cytologia (Tokyo) 59:25–30 Hake S, Ross-Ibarra J (2015) Genetic, evolutionary, and
Greizerstein EJ, Poggio L (1995) Meiotic studies of spon- plant breeding insights from the domestication of
taneous hybrids of Amaranthus: genome analysis. maize. eLife 4:1–8
Plant Breed 114:448–450 Hamdi A, El-Ghareib AA, Shafey SA et al (2003) Genetic
Greizerstein EJ, Naranjo CA, Poggio L (1997) variability, heritability and expected genetic advance
Karyological studies in five wild species of for earliness and seed yield from selection in lentil.
Amaranthus. Cytologia 62:115–120 Egypt J Agric Res 81(1):125–137
Grenier JCM, Godron DA (1856) Flore de France 3. Savy, Hamrick J, Godt H (1996) Conservation genetics of
F, Paris, p 779 endemic plant species. In: Avise JC, Hamrick JL (eds)
Grobelnik MS, Bavec M, Turinek M et al (2009a) Conservation genetics, case histories from nature.
Rheological properties of dongh made from grain Chapman and Hall, New York, pp 281–304
amaranth-cereal composite flours based on wheat and Han S, Xu B (2014) Bioactive Components of Leafy
spett. Czech J. Food Sci 27:309–319 Vegetable Edible Amaranth (Amaranthus mangosta-
Grobelnik MS, Turinek M, Jakop M et al (2009b) nus L.) as Affected by Home Cooking Manner. J Food
Nutrition value and use of grain amaranth: potential Sci Technol 2(4):122–127
future application in bread making. Agriculture Hanelt P (1968) Beitrage zur Kulturpflanzen-Flora I:
6:5–15 Bemerkungen zur Syatematik and Anbaugeschichte
Grubben GJH (1976) The cultivation of amaranth as a einiger Amaranthus-Arten. Kulturpflanze 16:127–149
tropical leaf vegetable with special reference to South Hardys H, Balick M, Schierwater B (1992) Applications
Dahomey, Report 67. Tropical Research Institute, of random amplified polymorphic DNA (RAPD) in
Amsterdam molecular ecology. Mol Ecol 1(1):55–63
Grubben GJH, Denton OA (ed) (2004) Plant resources of Harlan JR (1975) Crops and man. American Society of
tropical Africa 2. vegetables. PROTA Foundation, Agronomy, Madison
Wageningen, Netherlands/Backhuys Publishers, Hartmann HT, Kester DE, FTD JR et al (2011) Plant prop-
Leiden, Netherlands/CTA, Wageningen, Netherlands. agation principles and practices, 7th edn. Prentice Hall
Kingdom Plantae.net, pp 63–89 Publishers, Englewood Cliffs, pp 281–340
Grubben GJH, van Sloten DH (1981) Genetic resources of Hauptli H, Jain SK (1978) Biosystematics and agronomic
Amaranths. The International Board for Plant Genetic potential of some weedy and cultivate amaranths.
Resources Food and Agriculture Organization, Rome, Theor Appl Genet 52:177–185
p 45 Hauptli H, Jain S (1984) Allozyme variation and evolu-
Gudu S, Gupta VK (1988a) Male-sterility in the grain tionary relationships of grain amaranths (Amaranthus
amaranth (Amaranthus hypochondriacus ex-Nepal) spp.). Theor Appl Genet 69:153–165
variety Jumla. Euphytica 37:23–26 Hauptli H, Jain S (1985) Genetic variation in outcrossing
Gudu S, Gupta VK (1988b) Electrophoretic identification rate and correlated floral traits in a population of grain
of grain amaranth species and cultivars. Acta Hortic amaranth (Amaranthus cruentus L.). Genetica
218:231–238 66:21–27
Guidea SD, Babeanu N, Popa O et al (2012) Preliminary Hauptli H, Robin L, Jain SK (1979) Germplasm explora-
studies on in vitro behaviour of varios somatic explants tion in Central and South America. In: Proceedings of
from some cultivated Amaranthus genotypes. the second Amaranth conference, Kutztown, Emmaus,
Scientific Bulletin, Series, F. Biotechnologies PA, USA
16:9–14 Hauptli H, Lutz RL, Jain SK (1980) Germplasm explora-
Guillen FR, Baltensperger DD, Nelson LA et al (1999) tion in Central and South America. In: Proceedings of
Variability in ‘Plainsman’ grain amaranth. In: Janick the second Amaranth Conference, Kutztown, PA, 13–14
Sept 1979. Rodale Press, Emmaus, PA, pp 117–122
196 References
Heap IM (2002) The world’s wrost herbiside resistant Iamonico D (2010b) Biology, life strategy and invasive-
weed. Proc Weed Sci Soc Am 42:227 (Abstract) ness of Amaranthus retroflexus L. (Amaranthaceae) in
Heap IM (2008) The International survey of herbiside Central Italy: Preliminary research. Bot Serbica
resistant weeds. http://www.weedscience.com. 34:137–145
Accessed 18 Apr 2008 Iamonico D (2012) Amaranthus powellii subsp. cacciatoi
Heap IM (2010) The International survey of herbiside comb. et stat. nov. (Amaranthaceae). Nord J Bot
resistant weeds. http://www.weedscience.com. 30:12–16
Accessed 29 June 2010 Iamonico D (2013) Amaranthus blitum L. s.l. In: Invasive
Heap IM (2011) The international survey of herbiside species compendium. Wallingford, CAB International,
resistant weed. http://www.weedscience.com. UK. www.cabi.org/isc. Accessed 31 May 2014
Accessed 24 Feb 2011 Iamonico D (2014a) Lectotypification of Linnaean names
Henrickson J (1993) Amaranthaceae. In: Hickman JC (ed) in the genus Amaranthus L. (Amaranthaceae). Taxon
Jepson manual: higher plants of California, Berkeley, 63(1):146–150
pp 130–134 Iamonico D (2014b) Amaranthus gangeticus
Hermandez-Bermejo JE, Leon J (eds) (1994). Neglected (Amaranthaceae), a name incertae sedis. Phytotaxa
crops: 1492 from a different perspective. FAO plant 162(5):299–300
production and protection series. No. 26. Food and Iamonico D (2014c) Nomenclature survey of the genus
Agricultural Organization of the United Nations, Amaranthus (Amaranthaceae). 3. Names to the Italian
Rome Flora. Plant Biosystems (in press)
Hershkovitz MA (1989) Phylogenetic studies in Iamonico D (2014d) Amaranthus graecizans s.l.
Centrospermae: a brief appraisal. Taxon 38:602–608 (Amaranthacee) in Italia: note tassonomiche e distrib-
Hirschberg J, McIntosh I (1983) Molecular basis of herbi- utive. Informatore Botanico Italiano (in press)
side resistance in Amaranthus hybridus. Science Iamonico D, Iberite M (2012) Amaranthaceae and
222:1346–1349 Chenopodiaceae in Italy: current understanding and
Hitchcock AS, Green ML (1929) Standard species of future prospective. In: Timonin AK, Sukhorukov AP,
Linnaean genera of Phanerogamae. In: International Harper GH, Nilova MV (eds) Proceedings of the sym-
botanical congress, Cambridge 1930, Nomenclature posium “Caryophyllales”: new insights into the phy-
prepared by British botanists, London, pp 110–199 logeny, systematics and morphological evolution of
Holm GL, Plucknett DL, Pancho JV et al (1991) The the order. M V Lomonosov State University, Moscow,
world’s worst weeds: distribution and biology. Krieger 24–27 September 2012, pp 65–69
Publishing Company, Florida International Board for Plant Genetic Resources (IBPGR)
Holm GL, Doll J, Holm E et al (1997) World weeds. (1984) The potential for using in vitro techniques for
Natural histories and distribution. Wiley, New York germplasm collection. IBPGR Publications,
Hong QB, Hou L, Luo XY et al (2001) Using RAPD for AGP:IBPGR/83/108, IBPGR, Rome
evaluating genetic background among naked barley International Plant Genetic Resources Institute (IPGRI)
varieties in Sichuan Northwestern region. Sci Agric (1999) Directory of Germplasm collection. Online
Sin 34(20):133–138 data base, Rome. http://www.cgiar.org/ipgri/doc/dbin-
Hooker JD (1885) Flora of British India, vol 4. London tro.htm
Horak MJ, Peterson DE, Chessman DJ et al (1994) ISTA (International Seed Testing Association) (1985)
Pigweed identification: a pictorial guide to the com- International rules for seed testing. Seed Sci Technol
mon pigweeds of the Great Plains. Kansas State 13:299–513
University, Manhattan ISTA (International Seed Testing Association) (2010)
Huang H, Dane F, Kubisiak TL (1998) Allozyme and International rules for seed testing. International Seed
RAPD analysis of the genetic diversity and geographic Testing Association, Bassersdorf
variation in wild polulation of American Chestnut Iudina RS, Zheleznova NB, Zakharova OV et al (2005)
(Fagaceae). Am J Bot 85(7):1013–1021 Isozyme variation in a genetic collection of amaranths
Hugin G (1987) Einige Bemerkungen zu wenig bekannten (Amaranthus L.). Genetika 41:1681–1687
Amaranthus- Sippen (Amaranthaceae) Mitteleuropas. Jacobsen SE, Mujica A (2001) Advances en el cons-
Willdenowia 16:453–478 cimiento de resistencia a factories abióticos adversos
Hunziker AT (1965) Estudios sobre Amaranthus. en la quinoa (Chenopodium quinoa Willd.). In:
V. Revisión de las especies americanas con 1–4 estam- Jacobsen SE, Portillo Z (eds) Memorias, Primer Taller
bres, 1 a 5 tépalos, e inflorescencias exclusivamente Internacional Sobre Quinoa-Recurses Geneticos y
axilares. Kurtziana 2:27–52 sistems de producción. Universidad Nacional Agraria
Hurro JB, Cees MK (1991) The dual role of temperature La Molina, Lima, pp 10–14
in the regulation of theseasonal changes in dormancy Jain SM (2000) Mechanisms of spontaneous and
and germination of seeds of Polygonum. Oecologia induced mutations in plants. In: Moriarty M,
90:88–94 Mothersill C, Seymour C, Edington M, Ward JF, Fry
Iamonico D (2010a) On the presence of Amaranthus RJM (eds) Radiation research, vol 2. International
polygonoides L. (Amaranthaceae) in Europe. Phyton Association for Radiation Research, Lawrence,
(Horn, Austria) 50(2):205–219 pp 255–258
References 197
Jain SM (2009) Mutation induced genetic improvement of Joshi BD (1988) Under-utilized food plants of hilly
neglected crops. In: International conference on ‘New regions. Invited status paper presented in the national
approaches to orphan crops improvement in Africa.’ seminar on the under-utilized crop plants held at
September 2007, Bern, Switzerland, pp 115–126 NBPGR, New Delhi, India, p 21
Jain SM (2010) Mutagenesis in crop improvement under Joshi BD, Rana RS (1991) Grain amaranths: the future
the climate change. Rom Biotechnol Lett food crops, Shimla science monograph 3. National
15(2):88–106 Bureau of Plant Genetic Resources, New Delhi
Jain SK, Hauptli H, Vaidya KR (1982) Out crossing rate Joshi BD, Mehra KL, Sharma SD (1983) Cultivation of
in grain amaranths. J Hered 73(1):71–72 grain amaranth in the North-Western hills. Indian
Jain SK, Kulakow PA, Peters I (1984) Genetics and breed- Farm 32(12):35–37
ing of grain amaranth. In: Proceedings of the 3rd Juan R, Pastor J, Alaiz M et al (2007) Electrophoretic
Amaranth conference, Rodale Research Center, characterization of Amaranthus L. seed proteins and
Kutztown, PA, September 11–13, pp 174–191. Rodale its systematic implication. Bot J Linn Soc
Press, Emmaus, PA 155(1):57–63
Janovska D, Eepkova P, Bradova J (2008) Comparison of Jugran A, Bhatt ID, Rawal RS (2010) Characterisation of
SDS PAGE and chip electrophoresis in Amaranthus agro-diversity by seed storage protein electrophoresis:
species assesment, In J.Prohens J, Badenes ML (eds) focus on rice germplasm from Uttarakhand Himalaya,
Proceedings of the 18th EUCARPIA general congressí, India. Rice Sci 17(2):122–128
Valencia, Spain, 9–12 September 2008, pp 139–142 Karp A, Isaac PG, Ingram GS (1998) Molecular tools for
Jansen van Rensburg WS, Van Averbeke W, Slabbert R screening biodiversity: plant and animals. Chapman &
et al (2007) African leafy vegetables in South Africa. Hall, Thompson Science, London, pp 10–17
Water SA 3(3) (Special edition) Kauffman CS (1979) Grain amaranth research: an
Jensen HF (1970) A diallel selection mating system for approach to the development of a new crop. In:
cereal breeding. Crop Sci 10:629–635 Proceedings of the second amaranth Conference.
Jie H, Huang L, Chow WS et al (1993) Effect of supple- Rodale Press, Inc., Emmaus, pp 81–90
mentary Ultraviolet B radiation on rice and pea plants. Kauffman CS (1980) Grain amaranth research: an
Aust J Plant Physiol 20:129–142 approach to the development of a new crop. In:
Jisha KC, Vijayakumari K, Puthur JT (2013) Seed prim- Proceedings of the 2nd Amaranth Conference. Rodale
ing for abiotic stress tolerance: an overview. Acta Press, Emmaus, PA, p 81
Physiol Plant 35(5):1381–1396 Kauffman CS (1981a) Grain amaranth varietal improve-
Jofre-Garfias AE, Villegas-Sepulveda N, Cabrera- ment: breeding program. Rodale Press Inc, Emmaus
Ponce JL et al (1997) Agrobacterium mediated Kauffman CS (1981b) Grain amaranth varietal improve-
transformation of Amaranthus hypochondriacus: ment; breeding programme report 81–3, New crop
light and tissue specific expression of a pea chloro- Department. Organic Gardening and farming research
phyll a/b – binding protein promoter. Plant Cell Rep center. Rodale Press Inc, Emmaus
1(12):847–852 Kauffman CS (1982) Improved grain amaranth variety
Johnson SD, Linder HP, Steiner KE (1998) Phylogeny and their yields, Rodale Research Report, NC-81-1.
and radiation of pollination system in Disa Rodale Press Inc, Emmaus
(Orchidaceae). Am J Bot 85:402–411 Kauffman CS (1984) Thoughts on the development of
Jordan N (1996) Effects of the Triazine resistance muta- improved varieties of grain amaranths In: Proceedings
tion on fitness in Amaranthus hybridus (smooth pig- of 3rd Amaranth Conference, Rodale Research Center,
weed). J Appl Ecol 33:141–150 Kutztown, PA, 11–13 Sept 1984. Rodale Press,
Jordan D, Tao Ygodwin I, Henzell R et al (2003) Prediction Emmaus, PA
of hybrid performance in grain sorghum using RFLP Kauffman CS (1992) Realizing the potential of grain ama-
markers. Theor Appl Genet 106:559–567 ranths. Food Rev Int 8:5–21
Jorge Mario G, Bressani R (1987) A guide to amaranth Kauffman CS, Reider C (1986) Rodale amaranth germ-
grain cultivation. Amaranth News Lett 2:5–7 plasm collection. Rodale Press Inc, Emmaus
Joshi BD (1981a) Exploration for amaranth in North-West Kaul HP, Aufhammer W, Laibe B et al (1996) Suitability
India. Plant genetic resources news letter no. 48. of amaranth genotypes for grain and fodder use in
International Board for Plant Genetic Resources, FAO, Central Europe. Bodenkultur 47:173–181
Rome, Italy, pp 41–52 Keeley PE, Carter CH, Thullen RJ (1987) Influence of
Joshi BD (1981b) Catalogue on amaranth germplasm. planting date on growth of Palmer Amaranth
Regional Station, National Bureau of Plant Genetic (Amaranthus palmeri). Weed Sci 35:199–204
Resources (NBPGR), Phagli, Shimla,171004, India, Kehinde TO, Ajala MO, Daniel IO et al (2013)
p 42 Physiological and Genetic Integrity of Amaranth
Joshi BD (1985) Annapurna, a new variety of grain ama- (Amaranthus spp.) Seeds during storage. Int J Plant
ranth. Indian Farming 25(8):29–31 Breed Genet 7:35–46
Joshi BD (1986) Genetic variability in grain amaranths. Kendrick RE, Frankland B (1969) Photocontrol of germi-
Indian J Agric Sci 56:574–576 nation in Amaranthus caudatus. Planta 85:326–339
198 References
Ker YC, Chen RH, Wu CS (1993) Relationship of second- Konishi Y, Yoshimoto H (1989) Amaranth globulin as a
ary structure, microstructure, mechanical properties of heat stable emulsifying agent. Agric Biol Chem
heat-induced gel of Soy 11S globulin. Biosci Biotech 53:3327–3328
Biochem 57:536–541 Konishi Y, Nojima H, Okuno K et al (1985)
Kesarwani M, Md A, Natarajan K et al (2000) Molecular Characterization of starch granules from waxy, non-
cloning and the over expression to confer resistance to waxy, and hybrid seeds of Amaranthus hypocyondria-
fungal infection in transgenic tobacco and tomato. cus L. Agric Biol Chem 49(7):1965–1971
J Biol Chem 275:230–238 Koracev I (1969) Material on the phylogeny of the genus
Kgang IE, Emmenes L, Laloo N et al (2008) Amaranthus (Pollen morphology). Nauc Trud
Characterization of drought tolerant A. tricolor mutant Visselskostop. Int V Kolavov Raster Plovdiv
plants. In: Libiakova G, Gajdosova A (eds) 5th 28(1):133–136
International symposium of the European Amaranth Kowal T (1954) Cechy morfologiczne I anatomicznena-
Association on Amaranth – Plant for the Future, sion rodzaju Amaranthus L. oraz klucze do ich oznac-
Instutute of Plant Genetics and Biotechnology SAS, zania. Monogr Bot (Warszawa):170–193
Nitra, Slovak Republic, 9–14 November 2008, Kremer RJ (1993) Management of weed seed banks with
pp 66–69 microorganisms. Ecol Appl 3:42–52
Khoshoo TN (1995) Census of Indian biodiversity, tasks Krishna A, Kumar A (2005) Evaluation of radio protective
ahead. Curr Sci 69(1):14–17 effects of Rajgira (Amaranthus paniculatus) extract in
Khoshoo TN, Pal M (1972) Cytogenetic pattern in Swiss albino mice. J Radiat Res 46:233–239
Amaranthus: Chromosomes. Today 3:259–267 Kuhn M, Wagner S, Aufhammer W et al (1996) Einflub
Khurana DS, Singh J, Kaur B (2013) Genetic variability, von pflanzenbaulischer Mabnahmen auf die
correlation and path coefficient analysis in Mineralstoffgehalte von Amaranths, Buchweizen,
Amaranthus. Veg Sci 40(2):238–240 Reismelde und Hafer. DT Lebensm Rundshau
Kietlinski KD, Jimenez F, Jellen EN et al (2013) 92(1):47–152
Relationships between the Weedy Amaranthus hybri- Kulakow PA (1986) Genetic analysis of morphology and
dus (Amaranthaceae) and the Grain Amaranths. Crop flowering in cultivated and weedy Amaranthus.
Sci 54(1):220–228 Dissertation Abstracts, International B.S. Science and
Kigel J (1994) Development and ecophysiology of ama- Engineering 47(6):2294B. University of California,
ranth. In: Paredes-Lopeez O (ed) Amaranth: biology, Davis, USA
chemistry and technology. CRC Press, Boca Raton, Kulakow PA (1987) Genetics of grain amaranths II. The
pp 185–205 inheritance of determinance, panicle orientation,
Kihara H, Matsumura S (1935) Genetische Studien bei dwarfism, and embryo colour in Amaranthus cauda-
Amaranthus tricolor L. I. Verurbung der Blattfarbung tus. J Hered 78:293–297
und gestalt (Japanisch mit deutscher zusammenfas- Kulakow PA, Jain SK (1985) The inheritance of flowering
sung). Bot Zool 3 time in Amaranthus species. J Genet 64:85–100
King JC (2002) Biotecghnilogy: a solution for improving Kulakow PA, Jain SK (1987) Genetics of grain amaranths.
nutrient bioavailability. Int J Vitam Nutr Res IV. Variation and early generation response to selec-
72(1):7–12 tion in Amaranthus cruentus L. Theor Appl Genet
Kintzios SE (2006) Terrestrial plant derives anticancer 74:113–120
agents and plants used in anticancer research. Crit Rev Kulakow PA, Jain SK (1990a) Grain amaranths crop spe-
Plant Sci 25:79–113 cies, evolution and genetics. In: Minnesota Agriculture
Kirkpatrick BA (1995) Interspecific relationships within Proc, Fourth amaranth Conference, St. Paul University
the genus Amaranthus (Amaranthaceae). Ph.D. thesis. of Minnesota, USA, pp 105–114
Texas A & M University, USA Kulakow PA, Jain SK (1990b) Genetics of grain ama-
Kirtikar KR, Basu BD (1987) Indian medicinal plants, vol ranths. IV. Varieties and early generation response to
3, 2nd edn. International Book Distributors, India, selection in Amaranthus cruentus L. Theor Appl Genet
Dehra Dun, pp 2061–2062 74:113–120
Klopper K, Robel J (1989) Beitrge zur Systematik mor- Kulakow PA, Hauptli H, Jain S (1985) Genetics of grain
phologie und anatomie der gattung Amaranthus L. 2. Amaranthus. Mendalian analysis of six colour charac-
Samenmorphologie und-anatomie ausgewahtter ver- teristics. J Hered 76:27–30
treter. Gleditschia 17:171–182 Kumpulainen JT, Salonen JT (1999) Natural antioxidants
Koebner RMD, Powell W, Donini P (2001) Contribution and anticarcinogens in nutrition, health and disease.
of DNA molecular marker technologies to the genetics Woodhead Publishing, Cambrdige
and breeding of wheat and barley. Plant Breed Rev Kunth KS (1838) Flora Berolinensis 2. Duncker et
21:181–220 Humbolt, Berlin, p 438
Kolano B, Pando LG, Maluszynska J (2001) Molecular Lakhsminarasimhan P, Godbole A (2001) Amaranthaceae.
cytogenetic studies in Chenopodium quinoa and In: Singh NP et al (eds) Flora of Maharashtra State
Amaranthus caudatus. Acta Societatis Botanicorum dicotyledons, vol 2. Botanical Survey of India,
Poloniae 70(2):85–90 Calcutta, pp 777–793
References 199
Lanoue KZ, Wolf PG, Browning S et al (1996) Madhav CK, Sivaji K, Tulasi RK (2008) Flowering plants
Phylogenetic analysis of restriction site variation in of Chitoor district, 1st edn. Students Offset Printers,
wild and cultivated Amaranthus species Tirupati, p 291
(Amaranthaceae). Theor Appl Genet 93:722–732 Madhusoodanan KJ (1978) Cytogenetical studies on veg-
Laude HN (1957) Comparative pre-emergence heat toler- etable amaranths. Ph.D thesis, Bhopal University,
ance of some seeded grasses and of weeds. Bot Gaz Bhopal
119(1):44–46 Madhusoodanan KJ, Nazeer MA (1983) Comparative
Lee SW, Ledig FT, Johnson DR (2002) Genetic variation morphology of the somatic karyotypes of vegetable
at allozyme and RAPD markers in Pinus longaeva amaranths and its phylogenetic significance. Cytologia
(Pinaceae) of the White Mountains, California. Am 48:237–244
J Bot 89:566–577 Madhusoodanan KJ, Pal M (1981) Cytology of vegetable
Lee JR, Hong GY, Dixit A et al (2008) Characterization of amaranths. Bot J Linn Soc 82(1):61–68
microsatellite loci developed for Amaranthus hypo- Madhusoodanan KJ, Pal M (1984) Autotetraploids in
chondriacus and their cross amplification in wild spe- Amaranthus tricolor Linn. Indian J Genet
cies. Conserv Genet 9:243–246 44:181–185
Lee RM, Thimmapuram J, Thinglum KA et al (2009) Maity S, Banerjee G, Royl M et al (2000) Chemical
Sampling the waterhemp (Amaranthus tuberculatus) induced prolongation of seed viability and stress toler-
genome using pyrosequencing technology. Weed Sci ance capacity of mung bean seedlings. Seed Sci
57:463–469 Technol 28:155–162
Legere A (2005) Risks and consequences of gene flow Makus DJ (1990) Composition and nutritive value of veg-
from herbicide-resistant crops: canola (Brassica napus etable amaranth as affected by stage of growth, envi-
L) as a case study. Pest Manag Sci 61:292–300 ronment and method of preparation. In: Proceeding of
Legleiter TR, Bradley KW (2008) Glyphosate and multi- fourth Amaranth symposium Minnesota, Minnesota
ple herbicide resistance in waterhemp (Amaranthus Agricultural University, Saint Paul, pp 35–46
rudis) populations from Missouri. Weed Sci Malik SS, Singh SP (2006) Role of plant genetic resources
56:582–587 in sustainable agriculture. Indian J Crop Sci
Lehman JW (1990) The potential of grain amaranth in the 1(1–2):21–28
1990’s and beyond. Amaranth: perspectives on pro- Mallory MA, Hall RV, McNabb AR et al (2008)
duction, processing and marketing. Minneapolis, Development and characterization of microsatellite
Minnesota, USA, pp 1–15 markers for the grain amaranths. Crop Sci
Lehman JW, Clark RL, Frey KJ (1991) Biomass 48:1098–1106
heterosis and conbining ability in interspecific and Mandal N, Das PK (2002) Intra- and interspecific genetic
intraspecific matings of grain amaranths. Crop Sci diversity in grain amaranthus using ramdom amplified
31:1111–1116 polymorphic DNA markers. Plant Tissue Cult
Lemen C (1980) Allocation of reproductive effect to the Biotechnol 12(1):49–56
male and female strategies in wind-pollinated plants. Manhart JR, Rettig JH (1994) Gene sequence data. In:
Oecologia 45:156–159 Behnke H-D, Mabry TJ (eds) Caryophyllales: evolu-
Leon-Camacho M, Garcia-Gonzalez DL, Aparicio R tion and systematics. Springer, Berlin, pp 235–246
(2001) A detailed and comprehensive study of ama- Manikandaselvi S, Nithya V (2011) Development and
ranth (Amaranthus cruentus L.) oil fatty profile. Eur biochemical analysis of iron supplementary nutraceu-
Food Res Technol 213:349–355 ticals from Moringa oleifera and Amaranthus
Linnaeus C (1753) Species plantarum 2. Salvius, polygonoides. Adv Pharmacol Toxicol 12(3):47–51
Stockholm, p 1200 Marcone MF (1999) Evidence confirming the existence of
Linnaeus C (1763) Systema Naturae, 13th edn. Salvius, a 7S globulin-like storage protein in Amaranthus
Stockholm, p 559 hypochondriacus. Seed Food Chem 65:533–542
Lloyd DG (1982) Selection of combined versus separate Marcone MF, Yada RY (1998) Structural analysis of glob-
sexes in seed plants. Am Nat 120:571–585 ulins isolated from genetically different Amaranthus
Loiseleur JLA (1810) Notice sur les plantes a ajouter a hybrid lines. Food Chem 61:319–326
la Flore de France (Flora Gallica; avec quelques Margulies M, Egholm M, Altman WE et al (2005)
Correections et Observations. J B Sajou, Paris, Genome sequencing in microfabricated high-density
p 172 picolitre reactors. Nature 437:376–380
Londo JP, Chiang YC, Hung KH et al (2006) Martin FW, Telek L (1979) Vegetables for the hot humid
Phytogeography of Asian wild rice, Oryza rufipogon, tropics. Part 6: Amaranth and Celosia. U.S. Department
reveals multiple independent domestication of culti- of Agriculture, New Orleans, LA, pp 1–21
vated rice, Oryza sativa. Proc Natl Acad Sci Mathai PJ (1978) Amaranthus, a neglected vegetable.
103:9578–9583 Indian Farm 28(1):29–32
Lorenz K, Coulter L (1991) Quinoa flour in baked prod- Matsumura S (1938) Genetische Studien bei Amaranthus
ucts. Plant Foods Hum Nutr 41:213–223 tricolor L. Part 2. Weitere Untersuchungen uber
Lyon CK, Becker R (1987) Extraction and refining of oil Blattfarbung (in Japanese with German summary). Jpn
from amaranth seed. J Am Chem Soc 64:233–236 J Genet 13:289–305
200 References
Maughan PJ, Sisneros N, Luo M et al (2008) Construction 33(3):369–376 (special edition). http://www.wrc.org.
of an Amaranthus hypochondriacus bacterial artificial za
chromosome library and genomic sequencing of her- Modi JD, Kulkarni PR (1976) New starches – the proper-
bicide target genes. Crop Sci 48:585–594 ties of the starch from Amaranthus paniculatus. Acta
Maughan PJ, Smith SM, Fairbanks DJ et al (2010) Aliment 5:399–402
Development characterization and linkage mapping of Mohindeen HK, Irulappan I (1993) Improvement in ama-
single nucleotide polymorphism in the Grain amaranths. In: Chadha KL, Kalloo G (eds) Advances in
ranths (Amaranthus sp.). Plant Genome 4(1):92–101 horticulture: vegetable crops, vol 5, Malhotra
Mayer AM, Mayber AP (1995) The germination of seeds, Publishing House. New Delhi, India, pp 305–323
2nd edn. Pergamon Press, Oxford, pp 237–247 Mohler CL (1993) A model of the effects of tillage on
Mayo CM, Horak MJ, Peterson DE et al (1995) emergence of weed seedlings. Ecol Appl 3(1):53–73
Differential control of 4 amaranthus species by 6 Mohler CL, Galford AE (1997) Weed seedling emergence
postemergence herbicides in soybean (Glycine max). and seed survival: separating the effects of seed posi-
Weed Technol 9:141–147 tion and soil modification by tillage. Weed Res
McClure JW, Alston RE (1966) Chemotaxonomy of 37(3):147–155
Lemnaceae. Am J Bot 53:849–859 Moquin-Tandon CHBA (1849) Amaranthaceae. In: De
McCullum CM, Comai L, Greene EA et al (2000) Candolle A (ed) Prodromus systematis naturalis regni
Targetting Induced Local lesions INGenomics vegetabilis 13, vol 2. Victoris Masson, Paris,
(TILLING) for plant functional genomics. Plant pp 231–424
Physiol 123(2):439–442 Moscova C (2012) Influence of depth placement and dura-
McLean KS, Roy KM (1988) Incidence of Colletotrichum tion of stay in the soil of Amaranthus species seeds on
domatium an prickly side, spotted spurge and smooth rest and germination. Sci Pap Ser A Agron LV:306–309
pigweed and pathogenicity to soybean. Plant Dis Mosyakin SL (2005) On the origin of dioecious ama-
72(5):390–393 ranths (Amaranthus L., Amaranthaceae Juss.).
McNeill J, Barrie FR, Buck WR et al (eds) (2012) Ukrainskyl botanichnyi zhurnal 62(1):3–9
International code of nomenclature for algae, fungi Mosyakin SL, Robertson KR (1996) New infrageneric
and plants (Melbourne Code): adopted by the eigh- taxa and combination in Amaranthus (Amaranthaceae).
teenth international botanical congress, Melbourne, Ann Bot Fenn 33:275–281
Australia, July 2011. Regnum Vegetabile 154, Mosyakin SL, Robertson KR (2003) Amaranthus
Ruggell Linnaeus. In: Flora of North America editorial com-
Mead A, Gray D (1999) Production of seed longevity: a mittee (ed) Flora of North America. North of Mexico,
modification of the shape of Ellis and Roberts. Seed vol 4: Magnoliophyta, Caryophillidae, Part I. Oxford
Sci Res 9:63–73 University Press, New York pp 410–435
Mehta A, Datta A (1991) Oxalate decarboxylase from Mujica A, Jacobsen SE (2003) The genetic resources of
Collybia velutipes, purification, characterization and Andean grain amaranths (Amaranthus caudatus L., A.
cDNA cloning. J Biol Chem 266:23548–23553 cruentus L. and A. hypochondriacus L.) in America.
Melchinger AE, Coors JG, Pandey S (1999) Genetic Plant Genet Resour Newsl 133:41–44
diversity and heterosis. In: The genetics and exploita- Mulugeta S, Sentayehu A, Kassahun B (2012) Genetic
tion of heterosis in crops. In: Proceedings of an inter- varaiability, heritability, correlation coefficient and
national symposium, CIMMYT, Mexico City, Mexico, path analysis for yield and yield related traits in upland
17–22 August 1997. CIMMYT, Mexico, pp 99–118 rice. J Plant Sci 7(1):13–22
Melvin HW (2010) Nutrition for health, fitness and sport. Munusamy U, Abdullah SNA, Aziz MA, Khazaai H
Mc Graw-Hill, p 284 (2013) Female reproductive system of Amaranthus as
Miller JS, Venable DL (2000) Polyploid and the evolution the target for Agrobacterium mediated transformation.
of gender dimorphism. Science 289:2335–2338 Adv Biosci Biotechnol 4(2):188–192
Merrill ED (1950) Observation on cultivated plants with Murashige T, Skoog F (1962) A revised medium for rapid
reference to certain American problems. Ceiba 1:3–36 growth and bioassay with tobacco tissue culture.
Miller A, Wing B, Huete C (1984) The agricultural poten- Physiol Plant 15:473–497
tial of selected C4 plants in arid environments. J Arid Murray MJ (1938) Interspecific and intergenetic crosses
Environ 7:275 in the family Amaranthaceae. PhD thesis, Cornell
Misra PS, Pal M, Mitra CR et al (1971) Chemurgic studies University, Ithaca, NY
on some diploid and tetraploid grain amaranths. Proc Murray MJ (1940) The genetics of sex determination in
Indian Acad Sci Ser B 74:155–160 the family Amaranthaceae. Genetics 25:409–431
Mitchell CH, Diggle PK (2005) Evolution of unisexual Murray MJ (1960) Environmental vegetative heterosis.
flowers: morphological and functional convergence Argon J 52:509
results from diverse developmental transitions. Am Musa M, Singh A, Lawal AA (2014) Influence of priming
J Bot 92:1068–1076 duration on the performance of amaranths (Amaranthus
Modi AT (2007) Growth temperature and plant age influ- cruentus L.) in Sokoto semiarid zone of Nigeria. Int
ence on nutrional quality of Amaranthus leaves and J Agron:1–4
seed germination capacity. Water South Africa
References 201
Muthomi J, Musyimi DM (2009) Growth responses of Nowicke JW (1993) Pollen morphology and exine ultra-
african nightshades (Solanum scabrum mill) seedlings structure in Caryophyllales. In: Behnke HD, Mahbri
to water deficit. J Agric Biol Sci 5:2006–2009 TJ (eds) Evolution and systematic in Caryophyllales.
Muyonga JH, Nabakabya D, Nakimbungwe DN et al Springer Berlin Heidelberg, Berlin, pp 165–221
(2008) Effects to promote Amaranth production and Okuno K (1985) Expression of mutant genes specifying
consumption in Uganda to fight malnutrition. In: starch synthesis in cereal grains. Gamma Field Symp
Robertson GL, Lubien JR (eds) Using food Science 24:39–62
and Technology to Improve Nutrition and Promote Okuno K, Sakoguchi S (1982) Inheritance of starch char-
National Development. International union of food acteristics in perisperm of Amaranthus hypochondria-
sciences & technology. London cus. J Hered 73:467
Myers N (1990) The biodiversity challanges: expanded Oleszek W, Junkuszew M, Stochmal A (1999)
hotspots analyses. Environmentalist 10:243–256 Determination and toxicity of saponins from
Myers RL (1996) Amaranths: New crop opportunity. In: Amaranthus cruentus seeds. J Agric Food Chem
Janick J (ed) Progress in new crops proceedings of the 47:3685–3687
Third National Conference in new crops, Indianapolis, Omami EN, Haigh AM, Medd RW et al (1999) Changes
22–25 October 1996, ASHS Press, Alexandria, VA, in germinability, dormancy and viability of
pp 207–220 Amaranthus retroflexus as affected by depth and dura-
Nandula NK, Wright AA, Bond JA et al (2014) EPSPS tion of burial. Weed Res 39(5):345–354
amplification in glyphosate-resistant spiny amaranth Ort DP, Ahreus WH, Martin B et al (1983) Comparison of
(Amaranthus spinosus): a case of gene transfer via photosynthetic performance in triazine-resistance and
interspecific hybridization from glyphosate-resistant susceptible biotypes of Amaranthus hybridus. Plant
palmer amaranth (Amaranthus palmeri). Pest Manag Physiol 72:925–930
Sci 70(12):1092–1909 Osborne TB (1907) The proteins of the Wheat Kernel.
Naseema AS, Balakrishanan S, Nair MC (1983) Pathology Carnegie Institution of Washington, Washington, DC,
and control of seed mycoflora of some vegetables in p 119
Kerala. Agric Res J Kerala 21(2):32–37 Pal M (1972) Evolution and improvement of cultivated
National Academy of Sciences (NAS) (1975) Under- amaranths. I. Breeding system and inflorescence struc-
utilized tropical plants with promising economic value ture. Proc Ind Natl Sci Acad 38:28–37
National Academy of Sciences (NAS) (1984) National Pal M, Khoshoo TN (1968) Cytogenetics of the new auto-
Research Council Committee on Amaranth: modern tetraploid Amaranthus edulis. Technical
Prospect for an ancient crop. National Academy of Communication of the National Botanic Gardens,
Science Press, Washington, DC, USA Lucknow, pp 25–36
National Academy of Sciences (NAS) (1989) National Pal M, Khoshoo TN (1972) Evolution and improvement
Research Council Committee on dietary allowances, of cultivated amaranths. V. Inviability, Weakness and
recommended dietary allowances, 10th edn. National sterility in hybrids. J Hered 63:78–82
Academy of Science Press, Washington, DC, USA Pal M, Khoshoo TN (1973a) Evolution and improvement
National Academy of Sciences (NAS) (1996) Lost crops of cultivated amaranths. VI. Cytogenetic relationship
of Africa, vol I: grains. National Academy Press, in grain types. Theor Appl Genet 43:343–350
Washington, DC, USA Pal M, Khoshoo TN (1973b) Evolution and improvement
National Research Council (1984) Amaranth: modern of cultivated amaranths. VII. Cytogenetic relationship
prospects for an ancient crop. National Academy invegetable Amaranths. Theor Appl Genet
Press, Washington, DC 43:343–350
Nayar MP (1996) Hotspots of endemic plants of India, Pal M, Khoshoo TN (1974) Grain amaranths. In:
Nepal and Bhutan. TBGRI, Thiruvananthpuram Hutchinson JB (ed) Evolutionary studies in world
Nayar MP, Sastry ARK (1987–1990) Red data book of crops: diversity and change in the Indian sub-
Indian plants, vol 3. Botanical Survey of India, continent. UK, pp 129–137
Kolkata Pal M, Khoshoo TN (1977) Evolution and improvement
Nilsson J, Stegmark R, Akesson B (2004) Total antioxi- of cultivated amaranths. VIII. Induced autotetraploidy
dant capacity in different pea (Pisum sativum) variet- in grain types. Z Pflanzenzuchtg 78:135–148
ies blanching and freezing. Food Chem 86:501–507 Pal M, Khoshoo TN (1982) Evolution and improvement
Nolan C, Noyes A, Bennett A et al (2010) Inter Simple of cultivated Amaranths IV. Cytogenetic relationship
Sequence Repeats (ISSRs) reveal genetic variation between the two basic chromosome numbers. J Hered
amomg mid-atlantic polulations of threatened African 73:353–356
Amaranthus pumilis and phylogenetic relationship. Pal M, Pandey RM (1982) Decrease in quadrivalent fre-
Castanea 75:506–516 quency over a 10 years period in autotetraploids in two
Norsworthy JK, Griffith GM, Scott RC et al (2008) species of grain amaranths. Cytologia 47:795–801
Confirmation and control of glyphosate-resistant Pal M, Pandey RM, Khoshoo TN (1982) Evolution and
Palmer amaranth (Amaranthus palmeri) in Arkansas. improvement of cultivated amaranths. IX. Cytogenetic
Weed Technol 22:108–113 relationship between the two basic chromosome chro-
mosome numbers. J Herd 73:353–356
202 References
Pal M, Pandey RM, Khoshoo TN (1982b) Evolution and Patzoldt WL, Hager AG, McCormick JS et al (2006) A
improvement of cultivated amaranths. IX. Cytogenetic codon deletion confers resistance to herbisides inhibit-
relationship between the two basic chrosome number. ing protoporphyrinogen oxidase. Proc Natl Acad Sci U
J Hered 73(3):353–356 S A 03:12329–12334
Pal M, Ohri D, Subrahmanyam (2000) A new basic chro- Pedersen B, Kalinowski LS, Eggum BO (1987) The nutri-
mosome number for Amaranthus (Amaranthaceae). tive value of amaranth grain (Amaranthus caudatus) 1.
Cytologia 65:13–16 Protein and minerals of row and processed grain. Plant
Pal A, Swain SS, Das AB et al (2013) Stable germ line trans- Foods Hum Nutr 36:309–324
formation a leafy vegetable crop amaranth (Amaranthus Peters I, Jain S (1985) Genetic-variation of sex expression
tricolor L.) mediated by Agrobacterium tumefacience. cultivated Amaranthus spp.—solution to many puz-
In Vitro Cell Dev Biol – Plant 49(2):114–128 zles. Am J Bot 72:878
Palomino G, Rubi R (1991) Diferencias cromosomicas Peters I, Jain S (1987) Genetics of grain amaranths III. 3.
entre algunas especias y tipos del genero Amaranthus Gene-cytoplasmic male-sterility. J Hered 78:
distribuidos en Mexico, Actas Primer Congreso 251–256
International del Amaranto. Morelos, Mexico Plate AYA, Areas JAG (2002) Cholesterol-lowering effect
Pammi S, Schertz K, Xu G et al (1994) Random amplified of extruded amaranth (Amaranthus caudatus L.) in
polymorphic DNA markers in Sorghum. Theor Appl hypercholesterolemic rabbits. Food Chem 76:1–6
Genet 89:80–88 Poets AM, Fang Z, Clegg MT et al (2015) Barley landra-
Pan RS, Sirohi PS, Sivakami N (1992) Genetic divergence ces are characterized by geographically heterogeneous
in vegetable amaranths. Indian J Hortic genomic origin. Genome Biol 16:173
49(2):183–186 Poggio L (1988) Aspectos citogenéticos de los amarantos
Pandey RM (1982) Genetics of Agronomic traits in ama- silvestres y cultivados. Actas de las Primeras Jornadas
ranths. SABRAO J 14:121–129 Nacionales sobre Amarantos (Santa Rosa, La Pampa)
Pandey RM (1984a) A study of heterosis in Amaranthus. pp 34–49
SABRAO J 16:93–99 Pond WG, Lehmann JW, Elmore R et al (1991) Feeding
Pandey RM (1984b) Genetic studies of yield contributing value of raw or heated grain amaranth germplasm.
traits in Amaranthus. Theor Appl Genet Anim Feed Sci 33:231–236
Pandey DN (1998) Ethno-forestry: local knowledge for Popa G, Cornea CP, Ciuca M et al (2010) Studies of
sustainable forestry and livelihood security. genetic diversity in Amaranthus species using the
Himangshu Publication, New Delhi RAPD markers. Analele Universitati din Oradee-
Pandey ML, Gupta RC (1985) Studies on leaf surface Fascicula Biol Tom 17:280–285
mycoflora of Amaranthus paniculatus grown in Possehlo GL (ed) (1993) Harrapan civilization: a recent
Almora hills. Madras Agric J 72(5):272–275 perspective. Oxford and IBH Publishers, New Delhi
Pandey RM, Pal M (1985) Genetics of grain protein in Prakash D, Pal M (1991) Nutritional and anti-nutritional
Amaranthus. Crop Improv 12:55–58 composition of vegetable and grain amaranth leaves.
Pandey RM, Singh R (2010) Genetic studies for biochem- J Sci Food Agric 57:573–583
ical and quantitative characters in grain amaranth Prakash D, Pal M (1992) Seed protein, fat and fatty acid
(Amaranthus hypochondriacus L.). Plant Omics profile of Amaranthus species. J Sci Food Agric
J 3(4):129–134 58:145–147
Pandey RM, Singh R (2011) Genetic divergence in grain Prashantha GS, Nagaraja TS (2011) Variability and
amaranth (Amaranthus hypochondriacus L.). Genetika genetic diversity studies in grain amaranth (Amaranthus
43(1):41–49 spp.). J Maharashtra Agric Univ 36(1):63–66
Pandey RM, Dwivedi M, Singh R (2012) Biochemical Pratt DB, Clark LG (2001) Amaranthus rudis and A.
analysis of 25 genotypes of grain amaranths tuberculatus– one species or two? J Torrey Bot Soc
(Amaranthus hypochondriacus). Global J Sci Front 128:282–296
Res Bio-Tech Genet 12(5):1–12 Qiang WX, Jin PY (2013) Comparison of genetic diver-
Paredes-Lopez O, Cervantes-Ceja ML, Vigna-Perez M sity among Amaranth accessions from South and
et al (2010) Berries: improving human health and South-East Asia using SSR markers. Korean J Med
healthy aging and promoting quality life: a review. Crop Sci 21(3):220–228
Plant Foods Hum Nutr 65:299–308 Rafinesque CS (1836) Flora Telluriana 3. Printed by the
Park Y-J, Nishikawa T (2012) Rapid identification of author, Philadelphia, p 100
Amaranthus caudatus and Amaranthus hypochondria- Ragot M, Sisco PH, Hoisington DA et al (1995)
cus by sequencing and PCR-RFLP analysis of two Molecular-marker mediated characterization of favor-
starch synthase gene. Genome 55(8):623–628 able exotic alleles at quantitative trait loci in maize.
Patzoldt WL, Tranel PJ (2007) Multiple mutations confer Crop Sci 35:1306–1315
herbicide resistance in waterhemp (Amaranthus tuber- Rahman MH (2001) Production of yellow –seeded bras-
culatus). Weed Sci 55:421–428 sica napas through interspecific crosses. Plant Breed
Patzoldt WL, Tranel PJ, Hager AG (2005) A waterhemp 120:463–472
(Amaranthus tuberculatus) biotype with multiple Raina A, Datta A (1992) Molecular cloning of a gene
resistance across three herbicide sites of action. Weed encoding a seed specific protein with nutritionally bal-
Sci 53:30–36
References 203
anced amino acid composition from Amaranthus. Proc Roxburgh W (1832) Flora Indica; or, description of indian
Natl Acad Sci U S A 89:11774–11778 plants, vol 3. Serampore: Printed for W. Thacker and
Rajan S, Markose BL (2007) Propagation of horticultural Co, Calcutta and Parbury, Aellen and Co, London,
crops. In: Peter KV (ed) New India Publishing Agency, p 875
New Delhi, India, pp 110–113 Russell J, Fuller J, Macaulay M et al (1997) Direct com-
Rana JC, Yadav SK, Mandal S et al (2005) Genetic diver- parison of levels of genrtic variation amomg barley
gence and interrelationship analysis in grain amaranth accessions detected by RFLP, AFLP, SSRs and
(Amaranthus hypochondriacus) germplasm. Indian RAPDs. Theor Appl Genet 95:714–722
J Genet 65(2):99–102 Ryan GF (1970) Resistance to common groundsel to
Ranade SA, Sane PV (1995) PCR in plant genetic simazine and atrazine. Weed Sci 18:359–389
research. J Indian Bot Soc A74:431–441 Sammons B, Barnett OW (1987) Tobacco ringspot virus
Ranade SA, Kumar A, Goswami M et al (1997) Genome from squash grown in South Carolina and transmis-
analysis of Amaranths: determination of inter- and sion of the virus through seed of smooth pigweed.
intra-species variation. J Biosci 22(4):457–464 Plant Dis 71(6):530–532
Rayburn AL, McCloskey R, Tatum TC et al (2005) Sammour RH (1991) Using electrophoretic techniques in
Genome size analysis of weedy Amaranthus species. varietal identification, biosystematic analysis, phylo-
Crop Sci 45:2557–2562 genetic relations and genetic resources management.
Razna K, Ziarovska J, Labazova M (2012) Genome J Islam Acad Sci 4(3):221–226
changes in mutant lines of Amaranthus AS detected by Sammour RH, Hammoud MA, Abd Alla SA (1993)
microsatellite- directed PCR. ARPN J Agric Biol Sci Electrophoretic variations in Amaranthus. Bot Bull
7(11):877–884 Acadamia Sinica 34:37–42
Reddy BN, Chakravorty BP, Shekhawat PC (1980) Sammour RH, El-Zahwar Mustafa A, Badr S et al (2007)
Nutriational rquirements of Xanthomonas amaranthi- Genetic variation in accessions of Lathyrus sativus
cola, casual organism of bacterial leaf spot of ama- L. Acta Bot Croat 66:1–13
ranths. Indian Phytopathol 32(4):581–586 Sanches-Marroquin A, Del Valle FR, Escobedo M et al
Reif JC, Melchinger AE, Xia XC et al (2003a) Use of (1986) Evaluation of whole amaranth (Amaranthus
SSRs for establishing heterotic groups in subtropical cruentus), its air-classfied fractions, and blends of
maize. Theor Appl Genet 107:947–957 these with wheat and oats as possible components for
Reif JC, Melchinger AE, Xia XC et al (2003b) Genetic infant formulas. J Food Sci 51:1231–1234
distance based on simple sequence repeats and hetero- Sanchez-Del Pino I, Olvera HF, Valdes J (1999) La familia
sis in tropical maize populations. Crop Sci Amaranthaceae en la flora halofila gipsofila de
43:1275–1282 Mexico. Anales del Instuto de Biologia de la
Renner SS, Ricklefs RE (1995) Dioecy and its correlates. Universidad Nacional Autonoma de Mexico, Serie
Am J Bot 82:596–606 Botanica 70:29–135
Renner SS, Won H (2001) Repeated evolution of monoecy Sanchez-Marrowquin A (1983) Two forgotten crops of
in Siparunaceae (Laurales). Syst Biol 50:700–712 agro-Induatrial importance: amaranth and quinoa.
Resio ANC, Tolaba MP, Suarez C (2000) Some physical Archinos Latino americanos de Nutricion 33:11–32
and thermal characteristic of amaranth starch. Food Sánchez-Monge E (2002) Diccionario de plantas de
Sci Technol Int 65:371–378 interés agrícola. Ministerio de Agricultura, Pesca y
Richards AJ (1997) Plant breeding systems. Chapman and Alimentación, Madrid
Hall, New York Sangameswaran B, Jayakar B (2008) Anti-diabetic, anti-
Riggins CW, Peng Y, Stewart CN Jr et al (2010) hyperlipidemic and spermatogenic effects of
Characterization of de novo transcriptome for water- Amaranthus spinosus Linn. onstreptozotocin-induced
hemp (Amaranthus tuberculatus) using GS-FLX 454 diabetic rats. J Nat Med 62:79–82
pyrosequencing target-site genes. Port Manag Sci Sauer JD (1950) The grain amaranths. A survey of their
66:1042–1052 history & classification. Ann Mo Bot Gard
Roberts HA (1986) Seed persistence in soil and seasonal 37:561–632
emergence in plant species from different habitats. Sauer JD (1953) Herbarium species as records of genetic
J Appl Ecol 23:639–656 research. Am Nat 187:155–156
Roberts EH (1988) Seed aging: the genome and its expres- Sauer JD (1955) Revision of the dioecious amaranths.
sion. In: Nooden LD, Leopold AC (eds) Senescence and Madrono 13:5–46
aging in plants. Academic, San Diego, pp 465–498 Sauer JD (1957) Recent migration and evolution of the
Robertson KR (1981) The genera of Amaranthaceae in the dioecious amaranths. Evolution 11:11–31
Southeastern United states. J Arnold Arbor Sauer JD (1967) The grain amaranths and their relatives: a
62:267–313 revised taxonomic and geographic survey. Ann Mo
Román B, Hernández R, Pujadas-Salvá A et al (2007) Bot Gard 54:103–137
Genetic diversity in two variants of Orobanche graci- Sauer JD (1972) The dioecious amaranths: a new species
lis Sm. [var. gracilis and var. deludens (Beck) name and major range extension. Madrono
A. Pujadas](Orobanchaceae) from different regions of 21:425–434
Spain. J Biotechnol 10:0717–3458. doi:10.2225/ Sauer JD (1976) Grain amaranths. In: Simmonds NW (ed)
vol10-issue-2-fulltext-6 Evolution of crop plants. Longman, London, pp 4–7
204 References
Sauer JD (1993) Amaranthaceae: Amaranth family. In: Ecology, biogeography and management of Pinus
Historical geography of crop plants. Boca Raton, halepensis and P. brutia forest ecosystems in the
p 9–14 Mediterranean Basin. Backhuys Publishers, Leiden,
Saunders RM, Becker R (1984) Amaranthus: a potential pp 1–14
food and feed resource. In: Advances in cereal science Shoup DE, Al-Khatib K, Peterson DE (2003) Common
and technology, volume VI. American Association of waterhemp (Amaranthus rudis) resistance to protopor-
Cereal Chemists, INC, MN, pp 357–397 phyrinogen oxidase-inhibiting herbicides. Weed Sci
Schinz H (1934) Amaranthaceae. In Engler and Prantal 51:145–150
Di-cnatonrlichen Pflanzen familien, vol 160, pp 7–85 Shukla S, Bhargava A, Chatterjee A et al (2004) Estimates
Schnetzler KA, Breene WM (1994) Food uses and ama- of genetic parameters to determine variability for foli-
ranth product research: a comprehensive review. In: age yield and its different quantitative and qualitative
Peredes-Lopez O (ed) Amaranth biology, chemistry traits in the vegetable amaranth (Amaranthus tricolor).
and technology. CRC Press, Boca Raton, J Genet Breed 58:169–176
pp 155–184 Shukla S, Bhargava A, Chatterjee A et al (2006) Mineral
Schulz-Schaeffer J, Baldridge DE, Bowman HF et al profile and variability in vegetable amaranth
(1991) Registration of ‘Amont’ grain amaranth. Crop (Amaranthus tricolor). Plant Foods Hum Nutr
Sci 31:482–483 61:23–28
Segura–Nieto M, de la Rosa AP B, Parades-Lopez O et al Shukla S, Bhargava A, Chatterjee A et al (2010) Genetic
(1994) Biochemistry of amaranth protein. In: interrelationship among nutritional and qualitative
Amaranth: biology, chemistry and technology. CRC traits in the vegetable amaranths. Crop Breed Appl
Press, Boca Raton, pp 75–106 Biotechnol 10:16–22
Sellers BA, Smeda RJ, Johnson WG et al (2003) Simmonds NW (1990) The social context of plant breed-
Comparative growth of six Amaranthus species in ing. Plant Breed Abstr 60:337–341
Missouri. Weed Sci 51:329–333 Singh BP, Whitehead WH (1993) Population density and
Senft J (1979) Protein quality of amaranth grain. In soil pH effects on vegetable amaranth production. In:
Proceedings of 2nd Amaranth Conference Rodale Janick J, Simon JE (eds) New crops, vol 15. Wiley,
Research Center Kutztown, PA, 13–14 September, New York, Amaranth, pp 562–564
Rodale Press, Emmaus, PA Singhal RS, Kulkarni PR (1988) Composition of the seeds
Senthong C (1986) Yield improvement, agronomy and of some amaranthus species. J Sci Food Agric
local use of amaranth. In: Proceedings of third 42:325–331
Amaranth conference. Rodale Press Inc, Emmaus, PA Slabbert MM, de Rondi K, Caetano T et al (2003)
Shanna PN, Chowla SC (1987) Studies on a mosaic dis- Development and evaluation of mutant germplasm of
ease of amaranth (A. caudatus) in Himachal Pradesh. Amaranthus. In: genetic improvement of under-
Indian J Mycol Plant Pathol 16(3):349–350 utilized and neglected crops in low income food deficit
Shanna PN, Thakur V, Munjal RL (1981) Seed mycoflora countries, through irradiation and related techniques.
of Amaranthus caudatus, its pathology and control. Prcceedings of a final research coordination meeting,
Indian Phytopathol 33(2):242–244 organized by Joint FAO/IAEA, Pretoria, South Africa,
Sharma AK, Banik M (1965) Cytological investigation of pp 13–23
different genera of Amaranthaceae with a view to Smith JM, Haigh J (1974) The hitch-hiking effect of a
trace their interrelationship. Bull Bot Soc Bengal favourable gene. Genet Res 23:23–35
19:40–50 Snezana DM, Marija K, Danijela R et al (2012)
Sharma AD, Thakur V, Munjal RL (1981) Seed microflora Assessment of genetic relatedness of the two
of Amaranthus caudatus, it’s pathology and control. Amaranthus retroflexus polulation by protein and
Indian Phytopathol 33(2):242–244 Random Amplified Poymorphic DNA (RAPD) mark-
Sharma S, Dawson I, Waugh R (1995) Relationships ers. Afr J Biotechnol 11:7331–7337
among cultivated and wild lentils revealed by RAPD Sooby J, Myers RI, Baltensperger DD et al (1998)
analysis. Theor Appl Genet 91:647–654 Amaranth: production guide for the Central Unites
Sharma JR, Mudgal V, Hajra PK (1997) Floristic diver- States, a guide to growing and marketing. University
sity- review, scope and perspective. In: Mudgal V, of Nebraska Cooperative Extension, EC 98-151-S
Hajra PK (eds) Floristic diversity and conservation Soomarin SJ, Alipoor SH, Mahmoodabad RZ (2010)
strategies in India, vol I. Botanical Survey India, Evaluation of sulfuric acid application in breaking
Kolkata, pp 1–45 dormancy of goosefoot and red-root amaranth seeds.
Shewry PR (2002) The major seed storage proteins of Plant Ecophysiol 2:127–131
Spelt Wheat, Sorghum, Millets and Pseudocereals. In: Sosnoskie LM, Webster TM, MacRae AW et al (2009)
Belton P, Taylor J (eds) Pseudocereals and less com- Movement of glyphosate-resistant Palmer amaranth
mon cereals, grain properties and utilization potential. (Amaranthus palmeri) pollen in-field. Proc South
Springer-Verlag Berlin Heidelberg, pp 1–24 Weed Sci Soc 62:227
Shmida A, Lev-Yadun S, Goubitz S et al (2000) Sexual Souci SW, Fachman W, Kraut H (2000) Food composition
allocation and gender segregation in Pinus halepensis, and nutritional table. Wissenschaft Verlago GmbH,
P. brutia and P. pinea. In: Ne’emen G, Trabaud L (eds) Stuttgart
References 205
Sravanti V, Begum H, Sunil N et al (2012) Variance com- vation, Chinese Academy of Agricultural Science,
ponent analysis for grain yield and agro-economic Beijing, China, pp 110–112
traits in green amaranths (Amaranthus spp.). Adv Sung ZR, Dudits D (1981) Carrot somatic cell genetics.
Agric Sci Eng Res 2(7):233–244 In: Panopoulos NJ (ed) Genetic engineering in the
Sreelathakumary I, Peter KV (1993) Amaranth- plant sciences. Praeger, New York, pp 11–37
Amaranthus spp. In: Kallo G, Bergh BO (eds) Genetic Sunil M, Hariharan A, Nayak S et al (2014) The draft
improvement of vegetable crops. Pergamon Press, genome and transcriptome of Amaranthus hypochon-
Oxford, pp 315–323 driacus: a C4 Dicot producing High-Lysine edible
Srivastava R, Roy BK (2012) Analysis of genetic variation pseudo-cereals. DNA Res 21(6):585–602
among accessions of wild and cultivated species of Suprasanna P, Jain SM, Ochatt SJ et al (2010) Applications
Amaranthus based on RAPD and SDS-PAGE mark- of in vitro techniques in mutation breeding of vegeta-
ers. Int J Pharm Bio Sci 3(4):168–178 tively propagated crops. Plant Mutation. Ed. Q. Shu.
Srivastava R, Roy BK (2014) A new chromosome number IAEA, Vienna pp 369–383
for Amaranthus blitum. J New Biol Report Svobadova A, Psotova J, Walterova D (2003) Natural phe-
3(2):111–114 nolics in the prevention of UV-induced skin damage.
Stallknecht GF, Schulz-Schaeffer JR (1993) Amaranth A review. Biomed Pap 147(2):137–145
rediscovered. In: Janick J, Simon JE (eds) New crops. Swain SS, Sahu L, Banik DP et al (2010) Agrobacterium
Wiley, New York, pp 211–218 x plant factors influencing transformation of Joseph’s
Stebbins GL (1940) The significance of polyploidy in coat (Amaranthus tricolor L.). Sci Hortic
plant evolution. Am Nat 74:54–66 125(3):461–468
Stebbins GL (1947) Types of polyploid. Their classifica- Sweat JK, Horak MJ, Peterson DE et al (1998) Herbicide
tion and significance. Adv Genet 1:403–429 efficacy on four Amaranthus species in soybean
Stebbins GL (1957) Self fertilization and population vari- (Glycine max). Weed Technol 12:315–321
ability in higher plants. Am Nat 91:337–354 Syamkumar S, Sasikumar B (2007) Molecular Mrker
Stebbins G (1971) Chromosomal evolution in higher based genetic diversity analysis of Curcuma species
plans. Edward Arnold, London from India. Sci Hortic 112(2):235–241
Steckel LE (2007) The dioiecious Amaranth spp.: here to Tague BW, Mantis J (2006) In planta Agrobacterium
stay. Weed Technol 21:567–570 mediated transformation by vacuum infiltration.
Steckel LE, Cristy LS, Edward WS et al (2004) Methods Mol Biol 323:215–223
Temperature effects on germination of nine Tanksley SD, Orton TJ (eds) (1983) Isozyme in plant
Amaranthus species. Weed Sci 52:217–221 genetics and breeding, part A. Elsevier Science
Steckel LE, Sprague CL, Stoller EW et al (2007) Tillage, Publication B. V, New York
cropping system, and soil depth effects on Common Taylor JRN, Parker ML (2002) Quinoa. In: Belton PS,
Waterhemp (Amaranthus rudis) seed-bank persis- Taylor JRN (eds) Pseudocereals and less common
tence. Weed Sci 55:235–239 cereals: grain properties and utilization. Springer,
Steckel LE, Main CL, Ellis AT et al (2008) Palmer Berlin, pp 93–122
amaranth (Amaranthus palmeri) in Tennessee has low Taylorson RB, Borthwick HA (1969) Light filtration by
level glyphosate resistance. Weed Technol foliar canopies: significance for light-controlled weed
22:119–123 seed germination. Weed Sci 17:48–51
Stefu Nova V, Bezo M, Labajova M et al (2014) Genetic Tazeen M, Nadia K, Farzana NN (2009) Heritability, phe-
analysis of three Amaranthus species using ISSR notypic correlation and path coefficient studies for
markers. Emir J Food Agric 26(1):35–45 some agronomic characters in synthetic elite lines of
Steinback KE, McIntosh L, Bogorad L et al (1981) wheat. J Food Agric Environ 7(3&4):278–282
Identification of the triazine receptor protein as a chlo- Teutonico R, Knorr D (1985a) Nondestructive method for
roplast gene product. Proc Natl Acad Sci U S A determination of water-soluble Oxalate in cultured
78:7463–7467 Amaranthus tricolor cells. J Agric Food Chem
Stetter MG, Muller T, Schmid KJ (2015) Incomplete 33:60–62
domestication of south American grain amaranths Teutonico RA, Knorr D (1985b) Amaranth: composition,
(Amaranthus caudatus) from its wild relatives. properties and applications of a rediscovered food
doi:http://dx.doi.org/10.1101/025866 crop. J Food Technol 39:49–60
Sumar KL, Blanco OG, Alvarez AC et al (1983) Catalogo Thakur V (1964) Amaranthus palmeri S. Watts) in India.
de la coleccion de Amaranthus del CICA. Universidad J Indian Bot Soc 43:573–576
Nacional del Cusco, Cusco, Peru Thellung A (1914) Amaranthus. In: Ascherson P, Graebner
Sumar L, Pacheco J, Roca Conacha AI et al (1992) Grain P (eds) Synopsis der Mitteleuropaischen Flora 5(1).
amaranth research in Peru. Food Rev Inst 8(1):125–142 Engelmann, Leipzig, pp 225–356
Sun Y, Yue S (1993) Research on polyploid grain ama- Thellung A (1919) Beitrage zur Adventivflora der
ranth- a preliminary study on selection of grain ama- Schweiz 3. Vierteljahrsschr Natrforsch Ges Zurich
ranth with character of bigger seed (in Chinese). In: 64:684–815
Yue S (ed) The research and development of grain Thellung A (1926) Amaranthus bouchonii Thell. Monde
amaranth in China. Institue of crop breeding and culti- Plant 27:4–5
206 References
Thiers B (2011) Index herbariorum, a global directory of Tucker JM, Sauer JD (1958) Abarrant Amaranthus popu-
public herbaria and associated staff. New York lation of Sacramento- San Joaquin delta, California.
Botanical Garden’s Virtual Herbarium. Available Madrono 14:252–261
from http://sweetgum.nybg.org/ih/. Accessed 31 Turner BL (1954) Chromosome numbers and their phy-
May 2014. letic interpretation. In: Behenke HD, Mabry TJ (eds)
Tironi VA, Añón MC (2010) Amaranth proteins as a Caryophyllales evolution and systematic. New York,
source of antioxidant peptides: effect of proteolysis. pp 27–43
Food Res Int 43:315–322 Urvan I (1924) Sertum antillanum XX. Feddes Repertor
Toll J, Van Sloten DH (1982) Directory of germplasm col- 20:297–313
lections. International Board for Plant Genetic USDA-ARS (1999) National genetic resources program.
Resources, Via delle Terme di Caracalla, Rome, Germplasm Resources Information Network,
00100, Italy Beltsville
Toole EH (1946) Final results of the Duvel buried seed USDA-ARS (2001) National Genetic Resources
experiment. J Agric Res 72(6):201–210 Programme, Germplasm Resource Information
Tranel PJ, Wright TR (2002) Resistance of weeds to ALS- Network (GRIN). National Germplasm Resource
inhibiting herbiside: what have we learned ? Weed Sci Laboratory, Beltsville, MD. http://www.ars-grin.gov/
50:700–712 cgi-bin/npgs/acc/display.pl?1624948
Trannel PJ, Trucco F (2009) 21st century weed science: a Vaidya KR (1984) genetic varistion in landrace popula-
call for Amaranthus genomics. In: Stewart CN (ed) tion of Indian amaranths, Dessertatin abstracts. In:
Weedy and invasive plant genomics. Blackwell, Ames, International Bioscience and engineering, California,
pp 53–81 Davis, USA
Trannel PJ, Wassom JJ, Jeschke MR et al (2002) Vaidya KR, Jain SK (1987) Response to mass selection
Transmission of herbiside resistance from a monoe- for plant height and grain yield in amaranth
cious to a dioecious weedy Amaranthus species. Theor (Amaranthus spp.). Plant Breed 98(l):61–64
Appl Genet 105:674–679 Van-Rensburg J, Averbeke VW, Slabbert R et al (2007)
Transue DK, Fairabanks D, Robinson LR et al (1994) African leafy vegetables in South Africa. Water SA
Species identification by RAPD analysis of grain ama- 33(3):317–326
ranth genetic resources. Crop Sci 34:1385–1389 Varalakshmi B (2004) Characterization and preliminary
Trucco F, Tranel PL (2011) Amaranthus. In: Kole C (ed) evaluation of vegetable amaranth (Amaranthus spp.)
Wild crop relatives: genomics and breeding resources, germplasm. PGR News Lett 137:55–57
vegetables. Springer, Berlin/Heidelberg, pp 11–21 Vasil V, Castillo AM, Fromm ME et al (1992) Herbiside
Trucco F, Jeschke MR, Rayburn AL et al (2005a) resistant fertile transgenic wheat plant obtained by
Amaranthus hybridus can be pollinated frequently by microprojectile bombardment of regenerable embryo-
A. tuberculatus under field conditions. Heredity genic callus. Biotechnology 10:667–674
94:64–70 Vavilov N (1949) Origin, variation, immunity and breed-
Trucco F, Jeschke MR, Rayburn AL et al (2005b) ing of cultivated plants. Chron Bot 13:1–364
Promiscuity in weedy amaranths: high frequency of Vetter J (1994) Minerals and amino acids in the seeds of the
female tall waterhemp (Amaranthus tubercula- new, cultivated cereal-like species Amaranthus hypo-
tus) × smooth pigweed (A. hybridus) hybridization chondracus. Z. Lebensm. Unters Forsch 198:284–286
under field conditions. Weed Sci 53:46–54 Vietmeyer ND (1983) Amaranth: return of the Aztec mys-
Trucco F, Hager AG, Tranel PG (2006a) Acetolactate syn- tery crop. In: Year book of science in the future. P 188
thase mutation conferring imidazolinone-specific her- Vietmeyer ND (1984) Amaranth: modern prospects for an
bicide resistance in Amaranthus hybridus. J Plant ancient crop, The Board on Science and Technology
Physiol 163:475–479 for International Development (BOSTID). Rodale
Trucco F, Tatum T, Robertson KR et al (2006b) Research Centre of Rodale Press in Emmaus,
Characterization of waterhemp (Amaranthus tubercu- Pennsylvania, p 74
latus) x smooth pigweed (A.hybridus) F1 hybrids. Villars D (1807) Catalogue méthodique de plantes du
Weed Technol 20:14–21 jardin de Strasbourg. F G Levrault, Strasbourg,
Trucco F, Zheng D, Woodyard AJ et al (2007) Nonhybrid p 398
progeny from crosses of dioecious amaranths: impli- Virk PS, Ford-Lloyd BV, Jackson MT et al (1995) Use of
cations for gene-flow research. Weed Sci 55:119–122 RAPD for the study of diversity within plant diversity
Trucco F, Tatum T, Rayburn AL et al (2009) Out of the collection. Heredity 74:170–179
swamp: unidirectional hybridization with weedy spe- Vuylsteke MJR (1999) Genetic analysis of maize by using
cies may explain the prevalence of Amaranthus tuber- the AFLP® method. PhD thesis, University of
culatus as a weed. New Phytol 184:819–827 Wageningen, Netherlands
Tucker JB (1986) Amaranth: the once and future crop. Vuylsteke M, Kuiper M, Stam P (2000) Chromosomal
Bioscience 36(1):9–13 regions involved in hybrid performance and heterosis:
References 207
their AFLPbased identification and practical use in Wilkin P (1992) The status of Amaranthus bouchonii
prediction models. Heredity 85(3):208–218 Thell within Amaranthus L. section Amaranthus: new
Wallace TC (2011) Anthocyanins in cardiovascular dis- evidence from morphology and isoenzyme. J Linn Soc
ease. Adv Nutr 2:1–7 Bot 108:253–267
Walter CW (2001) Eat, drink and be healthy. Free Press, Williams JT, Brenner D (1995) Grain amaranths
New York, p 132 (Amaranthus species). In: Williams JT (ed) Cereals
Walter J, Dobes C (2004) Morphological characters, geo- and pseudocereals. Springer Berlin Heidelburg,
graphic distribution and ecology of neophytic London, pp 129–186
Amaranthus blitum L. subsp. emarginatus in Austria. Wilson CL (1924) Medullary bundle in relation to pri-
Annalen des Natuhristorischen Museums in Wien mary vascular system in Chenopodiaceae and
105(B):645–672 Amaranthaceae. Bot Gaz 78:175–199
Walters C (1998) Understanding the mechanism and Wilson MF (1979) Sexual selection in planta. Am Nat
kinetics of seed aging. Seed Sci Res 8:223–244 113:777–790
Walton PD (1968) Genetic homoeostasis in Amaranthus Wilson DO Jr, McDonald MB Jr (1992) Mechanical dam-
caudatus. J Hered 60:365–368 age in bean (Phaseolus vulgaris L.) seed in mecha-
Ward SM, Webster TM, Steckel LE (2013) Palmer nized and non-mechanized threshing systems. Seed
Amaranth (Amaranthus palmeri): a review. Weed Sci Technol 20:571–582
Technol 27:12–27 Wilson LT, Sterling WL, Rummel DR et al (1989)
Waselkov KE, Olsen KM (2014) population genetics and Quantitative sampling principles in cotton IPM. In:
origin of the native. North American Agricultural Friesbie RE et al (eds) Integrated pest management sys-
weed waterhemp (Amaranthus tuberculatus; tem and cotton production. Wiley, New York, pp 85–120
Amaranthaceae). Am J Bot 101(10):1726–1736 Wright R (1843) Icones plantarum Indiae Orientalis. J.B.
Wassom JJ, Tranel PJ (2005) Amplified fragment length Pharoah, Madras
polymorphism-based genetic relationship among Wu H (1998) Development of grain Amaranthus as a
weedy weedy Amaranthus species. J Hered starch crop in China. Ph.D. dissertation. University of
96:410–416 Hong Kong, Hong Kong
Wax LM (1995) Pigweeds of the Midwest—distribution, Wu H, Sun M, Yue S et al (2000) Field evaluation of an
importance, and management. Proc Iowa Integ Crop Amaranthus genetic resource collection in China.
Manage Conf 7:239–242 Genet Resour Crop Evol 47(1):43–53
Waycott W, Fort S (1994) Differentiation of nearly identi- Xia M, Wang D, Wang M et al (2004) Dracorhodin per-
cal germplasm by a combination of molecular and cholate induces apoptosis via activation of caspases
morphological analysis. Genome 85:239–244 and generation of reactive oxygen species. J Pharmacol
Wayne RB, Stanley TO, Mark SM et al (2000) Sci 95:273–283
Phytochemicals as bioactive agents. CRC Press, Xiao SG, Liu ZM, Song Y et al (2000) Classification of
Lancaster vegetable amaranth variety resources. J Hunan Agric
Weaver SE, McWilliams EL (1980) The biology of Univ 26(4):274–277
Canadian weeds. 44. Amaranthus retroflexus L., A. Xu F, Sun M (2001) Comparative analysis of phylogenetic
powellii S. Wats. and A. hybridus L. Can J Plant Sci relationships of grain amaranths and their wild rela-
60:1215–1234 tives (Amaranthus: Amaranthaceae) using internal
Webb DM, Smith CW, Schulz-Schaeffer J (1987) transcribed spacer, amplified fragment length poly-
Amaranth seedling emergence as affected by seeding morphism and double – primer fluorescent inter sim-
depth and temperature on a thermogradient plate. ple sequence repeat markers. Mol Phylogenet Evol
Agron J 79:23–26 21(3):372–387
Weber LE, Kauffman CS (1990) Amaranth grain produc- Yadav R, Rana JC, Ranjan JK (2014) Analysis of variabil-
tion guides. Rodale Press, Emmaus ity parameters for morphological and Agronomic traits
Wetzel DK, Horak MJ, Skinner DZ (1999a) Use of in grain amaranths (Amaranthus sp.) genotypes.
PCR-based molecular markers to identify weedy Bioscan 9(4):1661–1665
Amaranthus species. Weed Sci 47:518–523 Yang A, Su Q, An L (2009) Ovary-drip transformation: a
Wetzel DK, Horak MJ, Skinner DZ et al (1999b) Transfer simple method for directly generating vector and
of herbiside resistance from Amaranthus palmeri to marker free tranagenic maize (Zea mays L.) with lin-
Amaranthus rudis. Weed Sci 47:538–543 ear GFP cassette transformation. Planta 229:793–801
Wiese AM, Binning LK (1987) Calculating the threshold Yasmeen A, Mirza B, Inayatullah S et al (2009) In planta
temperature of development for weed. Weed Sci transformation of tomato. Plant Mol Biol Report
35:177–179 27:20–28
Wight R (1843) Icones plantarum Indiae Orientalis. J.B. Yoo K, Jang S (2003) Intra-specific relationship of
Pharoah, Madras Lactuca sativa var. capitata cultivars based on RAPD
analysis. Korean J Hortic Sci 21:273–278
208 References
Yoshimi Y, Chowdhury A, Hidenori K et al (2007) Sci Zharare GE (2012) Differential requirements for breaking
Hortic 112:366–375 seed dormancy in biotypes of Cleome gynandra and two
Yue S, Sun H (1993) The research and development of Amaranthus species. Afr J Agric Res 7(36):5049–5059
grain amaranths in China, pp 449–464. In: Yue S Zheleznov AV, Sonenko LR, Zheleznova NB (1997) Seed
(ed) The research and development of grain ama- proteins of the wild and cultivated Amaranthus spe-
ranths in China. Institute of Crop Breeding and cies. Euphytica 97:177–182
Cultivation, Chinese Academy of Agricultural Zoltan S (2009) Transgenic approach to improve the
Sciences, Beijing nutritional quality of wheat (Triticum aestivum L.).
Zebrowska J, Tyrka M (2003) The use of RAPD markers Ph.D. thesis, Tamásné Nyitrai Erzsébet Cecília Eötvös
for strawberry identification and genetic diversity Loránd University, Agricultural Research Institute of
studies. Food Agric Environ 1(1):115–117 the HAS, Martonvásár

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ISBN 978-981-10-1468-0 ISBN 978-981-10-1469-7 (eBook)

Library of Congress Control Number: 2016945253

© Springer Science+Business Media Singapore 2016

Printed on acid-free paper

This Springer imprint is published by Springer Nature

Taki, West Bengal, India Saubhik Das

4 Infrageneric Classification of Amaranths .................................. 49

8.4 Conventional Breeding by Selection .................................. 116

Appendices ............................................................................................ 173

References ............................................................................................. 189

© Springer Science+Business Media Singapore 2016 1

2.1 General Ghats, out of the 25 biodiversity hotspots (Fig.

© Springer Science+Business Media Singapore 2016 5

Fig. 2.2 (continued)

Fig. 2.3 Use of grain amaranths in ancient civilisations

3.1 General and A. blitum. The tender plant of grain species

© Springer Science+Business Media Singapore 2016 13

Fig. 3.1 Morphology of Amaranthus hypochondriacus L.

Fig. 3.3 Morphology of Amaranthus caudatus with characteristic drooping inflorescence

Fig. 3.4 Worldwide distribution of Amaranthus hypochondriacus L.

3.3.2 Amaranthus cruentus L. Southwestern USA, light brown grain type of A.

Fig. 3.6 Worldwide distribution of Amaranthus cruentus L.

Fig. 3.7 A. cruentus L.

Fig. 3.8 Worldwide distribution of Amaranthus caudatus L.

deltoid-ovate, pale-membranous, acuminate with 3.4 Potentiality of Grain

Fig. 3.9 A. caudatus L.

Table 3.2 Seed composition in different grain amaranths

Fig. 3.10 Different

Fig. 3.11 Worldwide distribution of Amaranthus tricolor L.

Fig. 3.12 Amaranthus

Fig. 3.13 Worldwide distribution of Amaranthus blitum L.

3.6.2 Amaranthus blitum L. base, glabrous, attaining a height of 10–30 cm.

in distal portion. Stems are green and branched.

3.6.3 Amaranthus dubius L. 3.6.4 Amaranthus cruentus L.

Fig. 3.15 Worldwide distribution of Amaranthus dubius Mart. ex Thell

in the cultivation field of the grain crop, uprooted

3.6.5 Amaranthus viridis L. terminal, often paniculate spikes, 2.5–12 cm long

Fig. 3.17 General plant morphology of (a) A. viridis L. (b) A. graecizans L.

8. Decreases risk of cardiovascular disease: ous/carcinogenic nitrosamines in the digestive

3.9 Culinary Properties The health beneficial antioxidant activities are

The word culinary means ‘related to cooking’; it

Fig. 3.18 (continued)

lanceolate-oblong or lanceolate, with subacute or

Fig. 3.20 Worldwide distribution of Amaranthus hybridus L.

Fig. 3.21 General plant morphology and inflorescence of Amaranthus hybridus L.

4.1 General 4.2 Synopsis of Infrageneric

© Springer Science+Business Media Singapore 2016 49

(Linnaeus 1753: 1027) Aellen ex K. R. Robertson 4.2.2 Amaranthus Subgen.

Key for the Infrageneric Delimitation of broadly triangular, occasionally

5.1 General grain species are closely related, but A. hypo-

© Springer Science+Business Media Singapore 2016 57

accessions of vegetable Amaranthus. Qualitative physiological and genetic integrity of nineteen

Fig. 5.2b Amaranthus

Fig. 5.3b Leaf phenolic chromatography differentiating grain amaranths

Fig. 5.4 Compiled

Amaranthus tricolor (Morphotype)

Fig. 5.5 Dendrogram computed from morphological data

Amaranthus tricolor (Morphotype)

Amaranthus tricolor var. tricolor

Amaranthus tricolor var. acutus

Amaranthus tricolor (Morphotype) Group A

Amaranthus tricolor (Morphotype)

Amaranthus hybridus (Morphotype)

10.1 General within 4–5 days after sowing. Amaranths do not

oisture content within medium-term gene-bank

10.3.4 Rainfall 10.4.1 Sowing of Seeds