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Comparison of endosperm development in weedy rice (Oryza sativa f. spontanea) and cultivated rice (Oryza sativa). Microstructure of endosperm cell of the weedy rice and associated cultivated rice (ACR). The transverse sections of endosperm at 3, 5, 7, and 9 d post-anthesis (DPA) were used for the observation of microstructure. N, nucleus; ES, endosperm; A, amyloplast; Cv, central vacuole; Nu, nucellus; NE, nucellar epidermis; FN, free nucleus; NP, nucellar projection; DV, dorsal vascular bundle; AL, aleurone layer; Pe, pericarp; Enc, endocarp. Ecw, endosperm cell wall; NP, nucellar projection; PB, pigment bundle; II, inner integument.

Comparison of endosperm development in weedy rice (Oryza sativa f. spontanea) and cultivated rice (Oryza sativa). Microstructure of endosperm cell of the weedy rice and associated cultivated rice (ACR). The transverse sections of endosperm at 3, 5, 7, and 9 d post-anthesis (DPA) were used for the observation of microstructure. N, nucleus; ES, endosperm; A, amyloplast; Cv, central vacuole; Nu, nucellus; NE, nucellar epidermis; FN, free nucleus; NP, nucellar projection; DV, dorsal vascular bundle; AL, aleurone layer; Pe, pericarp; Enc, endocarp. Ecw, endosperm cell wall; NP, nucellar projection; PB, pigment bundle; II, inner integument.

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Early maturity allows weedy rice ( Oryza sativa L. f. spontanea ) to persist by escaping harvest in paddy fields. A shorter grain-filling period contributes to the early maturity of weedy rice. However, the differences in morphology and endosperm development in the caryopsis between weedy and cultivated rice are largely unexplored. Here, we selecte...

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... change in color of weedy rice was 2 to 5 d earlier than that in cultivated rice ( The endosperm cells of weedy rice completed the cellularization stage at 3 DPA and had more amyloplasts. Compared with that of ACR, the starch accumulation of weedy rice was faster, and the starch grains in the endosperm were arranged more closely (Figure 3). The outer layers of endosperm cells transformed into aleurone layer cells 7 d after flowering in the small-grain cultivated varieties but 10 d after flowering in the large-grain cultivated varieties ( Chen et al. 2009). ...
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... has been reported that at 1 to 2 d after anthesis, the endosperm of rice was in the coenocyte stage ( Wei et al. 2002;Yu et al. 2014), which is consistent with our results. We showed that the endosperm cells of weedy and cultivated rice were in the coenocyte stage at 1 to 2 d after anthesis (Figure 3). At 3 DPA, the endosperm cells of weedy rice cellularized earlier and more rapidly than those of ACR. ...
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... formed 6 layers of endosperm cells at 3 DPA, while TZCR remained at the coenocytic stage at 3 DPA. From 5 DPA to 9 DPA, both TZWR and TZCR rapidly accumulated starch, but the starch grains in weedy rice were arranged more closely than those in cultivated rice (Figure 3). YZWR had finished the cellularization stage at 3 DPA, and a small amount of starch grains had accumulated in the endosperm cells, while YZCR had formed only 6 to 7 layers of endosperm cells at 3 DPA without stored starch grains. ...
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... YZWR and YZCR differentiated the aleurone layer from 5 DPA to 9 DPA, and there was a large amount of starch accumulation in the endosperm. Compared with those of YZCR, the amyloplasts of YZWR were larger, and the starch grains were arranged more closely (Figure 3). MMWR had finished the cellularization stage at 3 DPA, and its outer layer cells of endosperm had already stored some small amyloplasts, while MMCR had formed only 3 to 4 layers of endosperm cells at 3 DPA without stored starch grains (Figure 3). ...
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... with those of YZCR, the amyloplasts of YZWR were larger, and the starch grains were arranged more closely (Figure 3). MMWR had finished the cellularization stage at 3 DPA, and its outer layer cells of endosperm had already stored some small amyloplasts, while MMCR had formed only 3 to 4 layers of endosperm cells at 3 DPA without stored starch grains (Figure 3). DDWR had already finished cellularization at 3 DPA, and the endosperm cells were fully formed in the endosperm with some small amyloplasts stored, while DDCR had formed only 2 layers of endosperm cells at 3 DPA (Figure 3). ...
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... had finished the cellularization stage at 3 DPA, and its outer layer cells of endosperm had already stored some small amyloplasts, while MMCR had formed only 3 to 4 layers of endosperm cells at 3 DPA without stored starch grains (Figure 3). DDWR had already finished cellularization at 3 DPA, and the endosperm cells were fully formed in the endosperm with some small amyloplasts stored, while DDCR had formed only 2 layers of endosperm cells at 3 DPA (Figure 3). Li et al. (2014) found that compared with the late-maturing cultivar, the early-maturing cultivar had faster grain development, which was consistent with our observations. ...
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... cell proliferation mainly occurs at the cellularization stage ( Wei et al. 2002;Yu et al. 2014). We determined that weedy rice had finished cellularization at 3 DPA and already stored some small starch grains in endosperm cells (Figures 3 and 4). However, the cultivated rice was still at the cellularization stage, had formed fewer cell layers, and had no starch grain stored in the endosperm at 3 DPA. ...

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... The early maturation and shattering ability of weedy rice are well-recognized traits that can help its seeds to escape harvest, enter the soil seed bank, and hence persist in paddy fields over subsequent growing seasons (Gu et al., 2003;Thurber et al., 2010;Zhao et al., 2019). A less-well studied but potentially equally important trait is stem lodging, whereby the stems of the maturing plant fail to maintain erect growth and bend over at the stem base. ...
... Weedy rice has many morphological and physiological characteristics related to weediness, such as high phenotypic plasticity, rapid germination and growth, early seed maturity, easy seed shattering, long seed dormancy, and long lifespan (Delouche et al., 2007;Thurber et al., 2010;Andres et al., 2015;Dai et al., 2017;Zhao et al., 2019). The lodging ability of weedy rice is also a weediness trait that could synergize with early maturity, seed shattering, and dormancy to increase its continuity over successive growing seasons, which is an important survival strategy for it to persist in paddy fields. ...
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... Actin was used as the control. and black) or the elimination of color (white; Zhao et al., 2020). Chlorophyll is a highly photosensitive photosynthetic pigment, and temperature is also a critical abiotic factor influencing Chl accumulation (Ding et al., 2020). ...
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... Both WR and CR share similar morphological and physiological traits, which make controlling WR very difficult compared to other weeds (Chauhan, 2013). WR has many morphological and physiological characteristics related to weediness, such as a rapid growth rate, high phenotypic plasticity, early maturity, seed shattering, long seed dormancy, and longevity, which facilitate seed dispersal and persistence in paddy fields (Azmi and Karim, 2008;Burgos et al., 2014;Dai et al., 2014Dai et al., , 2017Zhao et al., 2018Zhao et al., , 2020. These characteristics contribute to its competitive advantage over CR in rice production areas globally (Delouche et al., 2007). ...
... Our previous study found that the shorter grain-filling period promotes early maturity in WR compared with the associated cultivated rice (ACR) (Zhao et al., 2018). Furthermore, the rapid development of endosperm cells and starch grains leads to the shorter grainfilling period of WR (Zhao et al., 2020). The endosperm accounts for 91-92% of the total weight of rice grains. ...
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... In addition, weedy rice has heavy shattering, which contribute to weedy rice escape from harvesting (Zhao et al. 2018). Furthermore, the rapid development of endosperm cells and starch grains leads to the shorter grain-lling period of weedy rice (Zhao et al. 2020). However, the relationship between the process of programmed cell death (PCD) in endosperm cells and the early maturity of weedy rice was unclear. ...
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