1932

Abstract

The persistent triploid endosperms of cereal crops are the most important source of human food and animal feed. The development of cereal endosperms progresses through coenocytic nuclear division, cellularization, aleurone and starchy endosperm differentiation, and storage product accumulation. In the past few decades, the cell biological processes involved in endosperm formation in most cereals have been described. Molecular genetic studies performed in recent years led to the identification of the genes underlying endosperm differentiation, regulatory network governing storage product accumulation, and epigenetic mechanism underlying imprinted gene expression. In this article, we outline recent progress in this area and propose hypothetical models to illustrate machineries that control aleurone and starchy endosperm differentiation, sugar loading, and storage product accumulations. A future challenge in this area is to decipher the molecular mechanisms underlying coenocytic nuclear division, endosperm cellularization, and programmed cell death.

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2022-05-20
2024-06-19
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Literature Cited

  1. 1.
    Abu-Zaitoon YM, Bennett K, Normanly J, Nonhebel HM 2012. A large increase in IAA during development of rice grains correlates with the expression of tryptophan aminotransferase OsTAR1 and a grain-specific YUCCA. Physiol. Plant. 146:487–99
    [Google Scholar]
  2. 2.
    Ajadi AA, Tong X, Wang H, Zhao J, Tang L et al. 2019. Cyclin-dependent kinase inhibitors KRP1 and KRP2 are involved in grain filling and seed germination in rice (Oryza sativa L.). Int. J. Mol. Sci. 21:245
    [Google Scholar]
  3. 3.
    Albani D, Hammond-Kosack MC, Smith C, Conlan S, Colot V et al. 1997. The wheat transcriptional activator SPA: a seed-specific bZIP protein that recognizes the GCN4-like motif in the bifactorial endosperm box of prolamin genes. Plant Cell 9:171–84
    [Google Scholar]
  4. 4.
    Bai A-N, Lu X-D, Li D-Q, Liu J-X, Liu C-M. 2016. NF-YB1-regulated expression of sucrose transporters in aleurone facilitates sugar loading to rice endosperm. Cell Res 26:384–88Identified the NF-YB1 transcription factor that activates sugar transporters in rice aleurone for grain filling.
    [Google Scholar]
  5. 5.
    Barkan A, Small I. 2014. Pentatricopeptide repeat proteins in plants. Annu. Rev. Plant Biol. 65:415–42
    [Google Scholar]
  6. 6.
    Barrôco RM, Peres A, Droual A-M, De Veylder L, Nguyen LSL et al. 2006. The cyclin-dependent kinase inhibitor Orysa;KRP1 plays an important role in seed development of rice. Plant Physiol 142:1053–64
    [Google Scholar]
  7. 7.
    Basunia MA, Nonhebel HM. 2019. Hormonal regulation of cereal endosperm development with a focus on rice (Oryza sativa). Funct. Plant Biol. 46:493–506
    [Google Scholar]
  8. 8.
    Bate NJ, Niu X, Wang Y, Reimann KS, Helentjaris TG. 2004. An invertase inhibitor from maize localizes to the embryo surrounding region during early kernel development. Plant Physiol 134:246–54
    [Google Scholar]
  9. 9.
    Batista RA, Köhler C. 2020. Genomic imprinting in plants—revisiting existing models. Gene Dev 34:24–36
    [Google Scholar]
  10. 10.
    Bauer MJ, Birchler JA. 2006. Organization of endoreduplicated chromosomes in the endosperm of Zea mays L. Chromosoma 115:383–94
    [Google Scholar]
  11. 11.
    Becraft PW, Gutierrez-Marcos J. 2012. Endosperm development: dynamic processes and cellular innovations underlying sibling altruism. WIREs Dev. Biol. 1:579–93
    [Google Scholar]
  12. 12.
    Becraft PW, Stinard PS, McCarty DR. 1996. CRINKLY4: a TNFR-like receptor kinase involved in maize epidermal differentiation. Science 273:1406–9
    [Google Scholar]
  13. 13.
    Becraft PW, Yi G. 2011. Regulation of aleurone development in cereal grains. J. Exp. Bot. 62:1669–75
    [Google Scholar]
  14. 14.
    Bello BK, Hou Y, Zhao J, Jiao G, Wu Y et al. 2019. NF-YB1-YC12-bHLH144 complex directly activates Wx to regulate grain quality in rice (Oryza sativa L.). Plant Biotechnol. J. 17:1222–35
    [Google Scholar]
  15. 15.
    Bernardi J, Lanubile A, Li QB, Kumar D, Kladnik A et al. 2012. Impaired auxin biosynthesis in the defective endosperm18 mutant is due to mutational loss of expression in the ZmYuc1 gene encoding endosperm-specific YUCCA1 protein in maize. Plant Physiol 160:1318–28
    [Google Scholar]
  16. 16.
    Boudet J, Merlino M, Plessis A, Gaudin JC, Dardevet M et al. 2019. The bZIP transcription factor SPA Heterodimerizing Protein represses glutenin synthesis in Triticum aestivum. Plant J 97:858–71
    [Google Scholar]
  17. 17.
    Brown RC, Lemmon BE, Olsen OA. 1994. Endosperm development in barley microtubule involvement in the morphogenetic pathway. Plant Cell 6:1241–52
    [Google Scholar]
  18. 18.
    Brown RC, Lemmon BE, Olsen OA. 1996. Polarization predicts the pattern of cellularization in cereal endosperm. Protoplasma 192:168–77
    [Google Scholar]
  19. 19.
    Brugière N, Humbert S, Rizzo N, Bohn J, Habben JE. 2008. A member of the maize isopentenyl transferase gene family, Zea mays isopentenyl transferase 2 (ZmIPT2), encodes a cytokinin biosynthetic enzyme expressed during kernel development: cytokinin biosynthesis in maize. Plant Mol. Biol. 67:215–29
    [Google Scholar]
  20. 20.
    Brugière N, Jiao S, Hantke S, Zinselmeier C, Roessler JA et al. 2003. Cytokinin oxidase gene expression in maize is localized to the vasculature, and is induced by cytokinins, abscisic acid, and abiotic stress. Plant Physiol 132:1228–40
    [Google Scholar]
  21. 21.
    Campbell WP, Lee JW, O'Brien TP, Smart MG. 1981. Endosperm morphology and protein body formation in developing wheat grain. Funct. Plant Biol. 8:5–19
    [Google Scholar]
  22. 22.
    Chen L-Q, Lin IW, Qu X-Q, Sosso D, McFarlane HE et al. 2015. A cascade of sequentially expressed sucrose transporters in the seed coat and endosperm provides nutrition for the Arabidopsis embryo. Plant Cell 27:607–19
    [Google Scholar]
  23. 23.
    Chen X, Feng F, Qi W, Xu L, Yao D et al. 2017. Dek35 encodes a PPR protein that affects cis-splicing of mitochondrial nad4 intron 1 and seed development in maize. Mol. Plant 10:427–41
    [Google Scholar]
  24. 24.
    Cheng WH, Taliercio EW, Chourey PS. 1996. The Miniature1 seed locus of maize encodes a cell wall invertase required for normal development of endosperm and maternal cells in the pedicel. Plant Cell 8:971–83Identified a cell wall invertase that cleaves sucrose to glucose and fructose, which are then loaded to the endosperm.
    [Google Scholar]
  25. 25.
    Cheng X, Pan M, E Z, Zhou Y, Niu B, Chen C 2020. Functional divergence of two duplicated Fertilization Independent Endosperm genes in rice with respect to seed development. Plant J 104:124–37
    [Google Scholar]
  26. 26.
    Cheng X, Pan M, E Z, Zhou Y, Niu B, Chen C 2020. The maternally expressed polycomb group gene OsEMF2a is essential for endosperm cellularization and imprinting in rice. Plant Commun 2:100092
    [Google Scholar]
  27. 27.
    Choi SB, Wang C, Muench DG, Ozawa K, Franceschi VR et al. 2000. Messenger RNA targeting of rice seed storage proteins to specific ER subdomains. Nature 407:765–67
    [Google Scholar]
  28. 28.
    Choi Y, Gehring M, Johnson L, Hannon M, Harada JJ et al. 2002. DEMETER, a DNA glycosylase domain protein, is required for endosperm gene imprinting and seed viability in Arabidopsis. Cell 110:33–42
    [Google Scholar]
  29. 29.
    Chou HL, Tian L, Kumamaru T, Hamada S, Okita TW 2017. Multifunctional RNA binding protein OsTudor-SN in storage protein mRNA transport and localization. Plant Physiol 175:1608–23
    [Google Scholar]
  30. 30.
    Chourey PS, Hueros G 2017. The basal endosperm transfer layer (BETL): gateway to the maize kernel. Maize Kernel Development BA Larkin 56–67 Boston, MA: CABI
    [Google Scholar]
  31. 31.
    Costa LM, Yuan J, Rouster J, Paul W, Dickinson H, Gutierrez-Marcos JF 2012. Maternal control of nutrient allocation in plant seeds by genomic imprinting. Curr. Biol. 22:160–65
    [Google Scholar]
  32. 32.
    Crofts AJ, Washida H, Okita TW, Satoh M, Ogawa M et al. 2005. The role of mRNA and protein sorting in seed storage protein synthesis, transport, and deposition. Biochem. Cell Biol. 83:728–37
    [Google Scholar]
  33. 33.
    Cross FR, Umen JG. 2015. The Chlamydomonas cell cycle. Plant J 92:370–92
    [Google Scholar]
  34. 34.
    Danilevskaya ON, Hermon P, Hantke S, Muszynski MG, Kollipara K, Ananiev EV 2003. Duplicated fie genes in maize: expression pattern and imprinting suggest distinct functions. Plant Cell 15:425–38
    [Google Scholar]
  35. 35.
    Dante RA, Larkins BA, Sabelli PA 2014. Cell cycle control and seed development. Front. Plant Sci. 5:493
    [Google Scholar]
  36. 36.
    Dante RA, Sabelli PA, Nguyen HN, Leiva-Neto JT, Tao Y et al. 2014. Cyclin-dependent kinase complexes in developing maize endosperm: evidence for differential expression and functional specialization. Planta 239:493–509
    [Google Scholar]
  37. 37.
    Deng X, Song XF, Wei L, Liu C, Cao X. 2016. Epigenetic regulation and epigenomic landscape in rice. Natl. Sci. Rev. 3:309–27
    [Google Scholar]
  38. 38.
    Diaz I, Vicente-Carbajosa J, Abraham Z, Martínez M, Moneda IIL, Carbonero P. 2002. The GAMYB protein from barley interacts with the DOF transcription factor BPBF and activates endosperm-specific genes during seed development. Plant J 29:453–64
    [Google Scholar]
  39. 39.
    Emes MJ, Bowsher CG, Hedley C, Burrell MM, Scrase-Field ESF, Tetlow IJ. 2003. Starch synthesis and carbon partitioning in developing endosperm. J. Exp. Bot. 54:569–75
    [Google Scholar]
  40. 40.
    Feng F, Qi W, Lv Y, Yan S, Xu L et al. 2018. OPAQUE11 is a central hub of the regulatory network for maize endosperm development and nutrient metabolism. Plant Cell 30:375–96
    [Google Scholar]
  41. 41.
    Forestan C, Meda S, Varotto S 2010. ZmPIN1-mediated auxin transport is related to cellular differentiation during maize embryogenesis and endosperm development. Plant Physiol 152:1373–90
    [Google Scholar]
  42. 42.
    Fu F-F, Xue H-W. 2010. Coexpression analysis identifies Rice Starch Regulator1, a rice AP2/EREBP family transcription factor, as a novel rice starch biosynthesis regulator. Plant Physiol 154:927–38
    [Google Scholar]
  43. 43.
    Furbank RT, Scofield GN, Hirose T, Wang XD, Patrick JW, Offler CE 2001. Cellular localisation and function of a sucrose transporter OsSUT1 in developing rice grains. Aust. J. Plant Physiol. 28:1187–96
    [Google Scholar]
  44. 44.
    Gao Y, An K, Guo W, Chen Y, Zhang R et al. 2021. The endosperm-specific transcription factor TaNAC019 regulates glutenin and starch accumulation and its elite allele improves wheat grain quality. Plant Cell 33:603–22Discovered the TaNAC019 transcription factor that regulates the accumulation of both storage proteins and starch in wheat endosperm.
    [Google Scholar]
  45. 45.
    Gehring M, Bubb KL, Henikoff S. 2009. Extensive demethylation of repetitive elements during seed development underlies gene imprinting. Science 324:1447–51
    [Google Scholar]
  46. 46.
    Gehring M, Missirian V, Henikoff S. 2011. Genomic analysis of parent-of-origin allelic expression in Arabidopsis thaliana seeds. PLOS ONE 6:e23687
    [Google Scholar]
  47. 47.
    Geisler-Lee J, Gallie DR 2005. Aleurone cell identity is suppressed following connation in maize kernels. Plant Physiol 139:204–12
    [Google Scholar]
  48. 48.
    Gong Z, Morales-Ruiz T, Ariza RR, Roldán-Arjona T, David L, Zhu J-K 2002. ROS1, a repressor of transcriptional gene silencing in Arabidopsis, encodes a DNA glycosylase/lyase. Cell 111:803–14
    [Google Scholar]
  49. 49.
    Gontarek BC, Neelakandan AK, Wu H, Becraft PW. 2016. NKD transcription factors are central regulators of maize endosperm development. Plant Cell 28:2916–36
    [Google Scholar]
  50. 50.
    Grafi G, Larkins BA. 1995. Endoreduplication in maize endosperm: involvement of M phase–promoting factor inhibition and induction of S phase–related kinases. Science 269:1262–64
    [Google Scholar]
  51. 51.
    Grossniklaus U, Vielle-Calzada J-P, Hoeppner MA, Gagliano WB. 1998. Maternal control of embryogenesis by MEDEA, a Polycomb group gene in Arabidopsis. Science 280:446–50
    [Google Scholar]
  52. 52.
    Gutiérrez-Marcos JF, Costa LM, Dal Prà M, Scholten S, Kranz E et al. 2006. Epigenetic asymmetry of imprinted genes in plant gametes. Nat. Genet. 38:876–78
    [Google Scholar]
  53. 53.
    Hamada S, Ishiyama K, Sakulsingharoj C, Choi SB, Wu Y et al. 2003. Dual regulated RNA transport pathways to the cortical region in developing rice endosperm. Plant Cell 15:2265–72
    [Google Scholar]
  54. 54.
    Hara T, Katoh H, Ogawa D, Kagaya Y, Sato Y et al. 2015. Rice SNF2 family helicase ENL1 is essential for syncytial endosperm development. Plant J 81:1–12
    [Google Scholar]
  55. 55.
    He Y, Ning T, Xie T, Qiu Q, Zhang L et al. 2011. Large-scale production of functional human serum albumin from transgenic rice seeds. PNAS 108:19078–83
    [Google Scholar]
  56. 56.
    He Y, Wang J, Qi W, Song R 2019. Maize Dek15 encodes the cohesin-loading complex subunit SCC4 and is essential for chromosome segregation and kernel development. Plant Cell 31:465–85
    [Google Scholar]
  57. 57.
    He Y, Yang Q, Yang J, Wang YF, Sun X et al. 2021. shrunken4 is a mutant allele of ZmYSL2 that affects aleurone development and starch synthesis in maize. Genetics 218:iyab070
    [Google Scholar]
  58. 58.
    Hibara K, Obara M, Hayashida E, Abe M, Ishimaru T et al. 2009. The ADAXIALIZED LEAF1 gene functions in leaf and embryonic pattern formation in rice. Dev. Biol. 334:345–54
    [Google Scholar]
  59. 59.
    Holding DR, Otegui MS, Li B, Meeley RB, Dam T et al. 2007. The maize Floury1 gene encodes a novel endoplasmic reticulum protein involved in zein protein body formation. Plant Cell 19:2569–82
    [Google Scholar]
  60. 60.
    Hsieh TF, Ibarra CA, Silva P, Zemach A, Eshed-Williams L et al. 2009. Genome-wide demethylation of Arabidopsis endosperm. Science 324:1451–54
    [Google Scholar]
  61. 61.
    Hu T, Tian Y, Zhu J, Wang Y, Jing R et al. 2018. OsNDUFA9 encoding a mitochondrial complex I subunit is essential for embryo development and starch synthesis in rice. Plant Cell Rep 37:1667–79
    [Google Scholar]
  62. 62.
    Huang X, Peng X, Sun MX. 2017. OsGCD1 is essential for rice fertility and required for embryo dorsal-ventral pattern formation and endosperm development. New Phytol 215:1039–58
    [Google Scholar]
  63. 63.
    Huang Y, Wang H, Huang X, Wang Q, Wang J et al. 2019. Maize VKS1 regulates mitosis and cytokinesis during early endosperm development. Plant Cell 31:1238–56Identified maize VKS1, which encodes a kinesin that is important for mitosis and cytokinesis during early endosperm development.
    [Google Scholar]
  64. 64.
    Ingram GC. 2017. Dying to live: cell elimination as a developmental strategy in angiosperm seeds. J. Exp. Bot. 68:785–96
    [Google Scholar]
  65. 65.
    Ishimaru K, Hirotsu N, Madoka Y, Murakami N, Hara N et al. 2013. Loss of function of the IAA-glucose hydrolase gene TGW6 enhances rice grain weight and increases yield. Nat. Genet. 45:707–11
    [Google Scholar]
  66. 66.
    Jääskeläinen AS, Holopainen-Mantila U, Tamminen T, Vuorinen T 2013. Endosperm and aleurone cell structure in barley and wheat as studied by optical and Raman microscopy. J. Cereal Sci. 57:543–50
    [Google Scholar]
  67. 67.
    Jameson PE, Song J 2016. Cytokinin: a key driver of seed yield. J. Exp. Bot. 67:593–606
    [Google Scholar]
  68. 68.
    Jeon JS, Ryoo N, Hahn TR, Walia H, Nakamura Y. 2010. Starch biosynthesis in cereal endosperm. Plant Physiol. Biochem. 48:383–92
    [Google Scholar]
  69. 69.
    Kang BH, Xiong Y, Williams DS, Pozueta-Romero D, Chourey PS. 2009. Miniature1-encoded cell wall invertase is essential for assembly and function of wall-in-growth in the maize endosperm transfer cell. Plant Physiol 151:1366–76
    [Google Scholar]
  70. 70.
    Kapazoglou A, Tondelli A, Papaefthimiou D, Ampatzidou H, Francia E et al. 2010. Epigenetic chromatin modifiers in barley: IV. The study of barley Polycomb group (PcG) genes during seed development and in response to external ABA. BMC Plant Biol 10:73
    [Google Scholar]
  71. 71.
    Kawakatsu T, Takaiwa F. 2010. Cereal seed storage protein synthesis: fundamental processes for recombinant protein production in cereal grains. Plant Biotechnol. J. 8:939–53
    [Google Scholar]
  72. 72.
    Kawakatsu T, Takaiwa F 2019. Rice proteins and essential amino acids. Rice: Chemistry and Technology J Bao 109–30 Duxford, UK: Woodhead Publ.
    [Google Scholar]
  73. 73.
    Kawakatsu T, Yamamoto MP, Touno SM, Yasuda H, Takaiwa F 2009. Compensation and interaction between RISBZ1 and RPBF during grain filling in rice. Plant J 59:908–20
    [Google Scholar]
  74. 74.
    Khoury CK, Bjorkman AD, Dempewolf H, Ramirez-Villegas J, Guarino L et al. 2014. Increasing homogeneity in global food supplies and the implications for food security. PNAS 111:4001–6
    [Google Scholar]
  75. 75.
    Kim SR, Yang JI, Moon S, Ryu CH, An K et al. 2009. Rice OGR1 encodes a pentatricopeptide repeat-DYW protein and is essential for RNA editing in mitochondria. Plant J 59:738–49
    [Google Scholar]
  76. 76.
    Kobayashi H. 2019. Variations of endoreduplication and its potential contribution to endosperm development in rice (Oryza sativa L.). Plant Prod. Sci. 22:227–41
    [Google Scholar]
  77. 77.
    Köhler C, Hennig L, Bouveret R, Gheyselinck J, Grossniklaus U, Gruissem W 2003. Arabidopsis MSI1 is a component of the MEA/FIE Polycomb group complex and required for seed development. EMBO J 22:4804–14
    [Google Scholar]
  78. 78.
    Kühn C, Grof CP. 2010. Sucrose transporters of higher plants. Curr. Opin. Plant Biol. 13:288–98
    [Google Scholar]
  79. 79.
    Kumamaru T, Uemura Y, Inoue Y, Takemoto Y, Siddiqui SU et al. 2010. Vacuolar processing enzyme plays an essential role in the crystalline structure of glutelin in rice seed. Plant Cell Physiol 51:38–46
    [Google Scholar]
  80. 80.
    Lau MMH, Stewart CEH, Liu Z, Bhatt H, Rotwein P, Stewart CL 1994. Loss of the imprinted IGF2/cation-independent mannose 6-phosphate receptor results in fetal overgrowth and perinatal lethality. Genes Dev 8:2953–63
    [Google Scholar]
  81. 81.
    LeClere S, Schmelz EA, Chourey PS. 2010. Sugar levels regulate tryptophan-dependent auxin biosynthesis in developing maize kernels. Plant Physiol 153:306–18
    [Google Scholar]
  82. 82.
    Leiva-Neto JT, Grafi G, Sabelli PA, Dante RA, Woo YM et al. 2004. A dominant negative mutant of cyclin-dependent kinase A reduces endoreduplication but not cell size or gene expression in maize endosperm. Plant Cell 16:1854–69
    [Google Scholar]
  83. 83.
    Li C, Yue Y, Chen H, Qi W, Song R. 2018. The ZmbZIP22 transcription factor regulates 27-kD γ-zein gene transcription during maize endosperm development. Plant Cell 30:2402–24
    [Google Scholar]
  84. 84.
    Li DQ, Wu XB, Wang HF, Feng X, Yan SJ et al. 2021. Defective mitochondrial function by mutation in THICK ALEURONE 1 encoding a mitochondrion-targeted single-stranded DNA-binding protein leads to increased aleurone cell layers and improved nutrition in rice. Mol. Plant 14:1343–61Showed that mutations of mitochondria-targeted SSB1 led to thick aleurone and improved nutrition in rice.
    [Google Scholar]
  85. 85.
    Li J, Berger F 2012. Endosperm: food for humankind and fodder for scientific discoveries. New Phytol 195:290–305
    [Google Scholar]
  86. 86.
    Li S, Zhou B, Peng X, Kuang Q, Huang X et al. 2014. OsFIE2 plays an essential role in the regulation of rice vegetative and reproductive development. New Phytol 201:66–79
    [Google Scholar]
  87. 87.
    Li X, Franceschi VR, Okitat TW. 1993. Segregation of storage protein mRNAs on the rough endoplasmic reticulum membranes of rice endosperm cells. Cell 72:869–79
    [Google Scholar]
  88. 88.
    Li XJ, Zhang YF, Hou M, Sun F, Shen Y et al. 2014. Small kernel 1 encodes a pentatricopeptide repeat protein required for mitochondrial nad7 transcript editing and seed development in maize (Zea mays) and rice (Oryza sativa). Plant J 79:797–809Showed that PPR-protein-mediated RNA editing in mitochondria is critical for grain filling in maize.
    [Google Scholar]
  89. 89.
    Lid SE, Gruis D, Jung R, Lorentzen JA, Ananiev E et al. 2002. The defective kernel 1 (dek1) gene required for aleurone cell development in the endosperm of maize grains encodes a membrane protein of the calpain gene superfamily. PNAS 99:5460–65
    [Google Scholar]
  90. 90.
    Liu C, Lu F, Cui X, Cao X 2010. Histone methylation in higher plants. Annu. Rev. Plant Biol. 61:395–420
    [Google Scholar]
  91. 91.
    Liu F, Ren Y, Wang Y, Peng C, Zhou K et al. 2013. OsVPS9A functions cooperatively with OsRAB5A to regulate post-Golgi dense vesicle-mediated storage protein trafficking to the protein storage vacuole in rice endosperm cells. Mol. Plant 6:1918–32
    [Google Scholar]
  92. 92.
    Liu G, Wu Y, Xu M, Gao T, Wang P et al. 2016. Virus-induced gene silencing identifies an important role of the TaRSR1 transcription factor in starch synthesis in bread wheat. Int. J. Mol. Sci. 17:1557
    [Google Scholar]
  93. 93.
    Liu J, Wu X, Yao X, Yu R, Larkin PJ, Liu CM 2018. Mutations in the DNA demethylase OsROS1 result in a thickened aleurone and improved nutritional value in rice grains. PNAS 115:11327–32Showed that weak alleles of ROS1 mutations exhibited thick aleurone and enhanced nutrition in rice.
    [Google Scholar]
  94. 94.
    Liu L, Tong H, Xiao Y, Che R, Xu F et al. 2015. Activation of Big Grain1 significantly improves grain size by regulating auxin transport in rice. PNAS 112:11102–7
    [Google Scholar]
  95. 95.
    Lu J, Magnani E 2018. Seed tissue and nutrient partitioning, a case for the nucellus. Plant Reprod. 31:309–17
    [Google Scholar]
  96. 96.
    Luo M, Bilodeau P, Koltunow A, Dennis ES, Peacock WJ, Chaudhury AM. 1999. Genes controlling fertilization-independent seed development in Arabidopsis thaliana. PNAS 96:296–301
    [Google Scholar]
  97. 97.
    Luo M, Platten D, Chaudhury A, Peacock WJ, Dennis ES. 2009. Expression, imprinting, and evolution of rice homologs of the polycomb group genes. Mol. Plant 2:711–23
    [Google Scholar]
  98. 98.
    Luo M, Taylor JM, Spriggs A, Zhang H, Wu X et al. 2011. A genome-wide survey of imprinted genes in rice seeds reveals imprinting primarily occurs in the endosperm. PLOS Genet 7:e1002125
    [Google Scholar]
  99. 99.
    Ma L, Zhang D, Miao Q, Yang J, Xuan Y, Hu Y 2017. Essential role of sugar transporter OsSWEET11 during the early stage of rice grain filling. Plant Cell Physiol 58:863–73
    [Google Scholar]
  100. 100.
    Marzábal P, Busk PK, Ludevid MD, Torrent M. 1998. The bifactorial endosperm box of γ-zein gene: characterisation and function of the Pb3 and GZM cis-acting elements. Plant J 16:41–52
    [Google Scholar]
  101. 101.
    Mena M, Vicente-Carbajosa J, Schmidt RJ, Carbonero P. 1998. An endosperm-specific DOF protein from barley, highly conserved in wheat, binds to and activates transcription from the prolamin-box of a native B-hordein promoter in barley endosperm. Plant J 16:53–62
    [Google Scholar]
  102. 102.
    Milne RJ, Grof CP, Patrick JW. 2018. Mechanisms of phloem unloading: shaped by cellular pathways, their conductances and sink function. Curr. Opin. Plant Biol. 43:8–15
    [Google Scholar]
  103. 103.
    Mizutani M, Naganuma T, Tsutsumi K, Saitoh Y 2010. The syncytium-specific expression of the Orysa;KRP3 CDK inhibitor: implication of its involvement in the cell cycle control in the rice (Oryza sativa L.) syncytial endosperm. J. Exp. Bot. 61:791–98
    [Google Scholar]
  104. 104.
    Moreno-Romero J, Jiang H, Santos-González J, Köhler C. 2016. Parental epigenetic asymmetry of PRC2-mediated histone modifications in the Arabidopsis endosperm. EMBO J 35:1298–311
    [Google Scholar]
  105. 105.
    Müntz K. 1998. Deposition of storage proteins. Plant Mol. Biol. 38:77–99
    [Google Scholar]
  106. 106.
    Nayar S, Kapoor M, Kapoor S. 2014. Post-translational regulation of rice MADS29 function: homo-dimerization or binary interactions with other seed-expressed MADS proteins modulate its translocation into the nucleus. J. Exp. Bot. 65:5339–50
    [Google Scholar]
  107. 107.
    Offler CE, McCurdy DW, Patrick JW, Talbot MJ 2003. Transfer cells: cells specialized for a special purpose. Annu. Rev. Plant Biol. 54:431–54
    [Google Scholar]
  108. 108.
    Ohad N, Yadegari R, Margossian L, Hannon M, Michaeli D et al. 1999. Mutations in FIE, a WD Polycomb group gene, allow endosperm development without fertilization. Plant Cell 11:407–15
    [Google Scholar]
  109. 109.
    Olsen OA. 2004. Nuclear endosperm development in cereals and Arabidopsis thaliana. Plant Cell 16:S214–27
    [Google Scholar]
  110. 110.
    Oñate L, Vicente-Carbajosa J, Lara P, Díaz I, Carbonero P 1999. Barley BLZ2, a seed-specific bZIP protein that interacts with BLZ1 in vivo and activates transcription from the GCN4-like motif of B-hordein promoters in barley endosperm. J. Biol. Chem. 274:9175–82
    [Google Scholar]
  111. 111.
    Ono A, Yamaguchi K, Fukada-Tanaka S, Terada R, Mitsui T, Iida S. 2012. A null mutation of ROS1a for DNA demethylation in rice is not transmittable to progeny. Plant J 71:564–74
    [Google Scholar]
  112. 112.
    Onodera Y, Suzuki A, Wu CY, Washida H, Takaiwa F. 2001. A rice functional transcriptional activator, RISBZ1, responsible for endosperm-specific expression of storage protein genes through GCN4 motif. J. Biol. Chem. 276:14139–52
    [Google Scholar]
  113. 113.
    Paine JA, Shipton CA, Chaggar S, Howells RM, Kennedy MJ et al. 2005. Improving the nutritional value of Golden Rice through increased pro-vitamin A content. Nat. Biotechnol. 23:482–87
    [Google Scholar]
  114. 114.
    Pan Z, Ren X, Zhao H, Liu L, Tan Z, Qiu F 2019. A mitochondrial transcription termination factor, ZmSmk3, is required for nad1 intron4 and nad4 intron1 splicing and kernel development in maize. G3 9:2677–86
    [Google Scholar]
  115. 115.
    Panda BB, Sekhar S, Dash SK, Behera L, Shaw BP. 2018. Biochemical and molecular characterisation of exogenous cytokinin application on grain filling in rice. BMC Plant Biol 18:89
    [Google Scholar]
  116. 116.
    Park K, Kim MY, Vickers M, Park JS, Hyun Y et al. 2016. DNA demethylation is initiated in the central cells of Arabidopsis and rice. PNAS 113:15138–43
    [Google Scholar]
  117. 117.
    Pedroza-Garcia JA, Eekhout T, Achon I, Nisa M-U, Coussens G et al. 2021. Maize ATR safeguards genome stability during kernel development to prevent early endosperm endocycle onset and cell death. Plant Cell 33:2662–84
    [Google Scholar]
  118. 118.
    Pu C-X, Ma Y, Wang J, Zhang Y-C, Jiao X-W et al. 2012. Crinkly4 receptor-like kinase is required to maintain the interlocking of the palea and lemma, and fertility in rice, by promoting epidermal cell differentiation. Plant J 70:940–53
    [Google Scholar]
  119. 119.
    Qi W, Tian Z, Lu L, Chen X, Chen X et al. 2017. Editing of mitochondrial transcripts nad3 and cox2 by Dek10 is essential for mitochondrial function and maize plant development. Genetics 205:1489–501
    [Google Scholar]
  120. 120.
    Qi W, Yang Y, Feng X, Zhang M, Song R. 2017. Mitochondrial function and maize kernel development requires Dek2, a pentatricopeptide repeat protein involved in nad1 mRNA splicing. Genetics 205:239–49
    [Google Scholar]
  121. 121.
    Qi X, Li S, Zhu Y, Zhao Q, Zhu D, Yu J 2017. ZmDof3, a maize endosperm-specific Dof protein gene, regulates starch accumulation and aleurone development in maize endosperm. Plant Mol. Biol. 93:7–20
    [Google Scholar]
  122. 122.
    Qian D, Qiu B, Zhou N, Takaiwa F, Yong W, Qu LQ 2020. Hypotensive activity of transgenic rice seed accumulating multiple antihypertensive peptides. J. Agric. Food Chem. 68:7162–68
    [Google Scholar]
  123. 123.
    Qiao Z, Qi W, Wang Q, Feng Y, Yang Q et al. 2016. ZmMADS47 regulates zein gene transcription through interaction with Opaque2. PLOS Genet 12:e1005991
    [Google Scholar]
  124. 124.
    Qin P, Zhang G, Hu B, Wu J, Chen W et al. 2021. Leaf-derived ABA regulates rice seed development via a transporter-mediated and temperature-sensitive mechanism. Sci. Adv. 7:eabc8873
    [Google Scholar]
  125. 125.
    Qu LQ, Takaiwa F. 2004. Evaluation of tissue specificity and expression strength of rice seed component gene promoters in transgenic rice. Plant Biotechnol. J. 2:113–25
    [Google Scholar]
  126. 126.
    Radchuk V, Riewe D, Peukert M, Matros A, Strickert M et al. 2017. Down-regulation of the sucrose transporters HvSUT1 and HvSUT2 affects sucrose homeostasis along its delivery path in barley grains. J. Exp. Bot. 68:4595–612
    [Google Scholar]
  127. 127.
    Ren Y, Wang Y, Liu F, Zhou K, Ding Y et al. 2014. GLUTELIN PRECURSOR ACCUMULATION3 encodes a regulator of post-Golgi vesicular traffic essential for vacuolar protein sorting in rice endosperm. Plant Cell 26:410–25
    [Google Scholar]
  128. 128.
    Ren Y, Wang Y, Pan T, Wang Y, Wang Y et al. 2020. GPA5 encodes a Rab5a effector required for post-Golgi trafficking of rice storage proteins. Plant Cell 32:758–77
    [Google Scholar]
  129. 129.
    Rodrigues JA, Ruan R, Nishimura T, Sharma MK, Sharma R et al. 2013. Imprinted expression of genes and small RNA is associated with localized hypomethylation of the maternal genome in rice endosperm. PNAS 110:7934–39
    [Google Scholar]
  130. 130.
    Rodrigues JC, Luo M, Berger F, Koltunow AM. 2010. Polycomb group gene function in sexual and asexual seed development in angiosperms. Sex Plant Reprod 23:123–33
    [Google Scholar]
  131. 131.
    Sabelli PA, Larkins BA. 2009. The development of endosperm in grasses. Plant Physiol 149:14–26
    [Google Scholar]
  132. 132.
    Sabelli PA, Liu Y, Dante RA, Lizarraga LE, Nguyen HN et al. 2013. Control of cell proliferation, endoreduplication, cell size, and cell death by the retinoblastoma-related pathway in maize endosperm. PNAS 110:E1827–36Revealed the critical role of the RBR pathway in cell cycle regulation of maize endosperms.
    [Google Scholar]
  133. 133.
    Sato H, Suzuki Y, Sakai M, Imbe T. 2002. Molecular characterization of Wx-mq, a novel mutant gene for low-amylose content in endosperm of rice (Oryza sativa L.). Breeding Sci. 52:131–35
    [Google Scholar]
  134. 134.
    Sattler SE, Singh J, Haas EJ, Guo L, Sarath G, Pedersen JF. 2009. Two distinct waxy alleles impact the granule-bound starch synthase in sorghum. Mol. Breeding 24:349–59
    [Google Scholar]
  135. 135.
    Schmidt R, Schippers JHM, Mieulet D, Watanabe M, Hoefgen R et al. 2014. SALT-RESPONSIVE ERF1 is a negative regulator of grain filling and gibberellin-mediated seedling establishment in rice. Mol. Plant 7:404–21
    [Google Scholar]
  136. 136.
    Schweizer L, Yerk-Davis GL, Phillips RL, Srienc F, Jones RJ. 1995. Dynamics of maize endosperm development and DNA endoreduplication. PNAS 92:7070–74
    [Google Scholar]
  137. 137.
    Scofield GN, Hirose T, Gaudron JA, Upadhyaya NM, Ohsugi R, Furbank RT 2002. Antisense suppression of the rice transporter gene, OsSUT1, leads to impaired grain filling and germination but does not affect photosynthesis. Funct. Plant Biol. 29:815–26
    [Google Scholar]
  138. 138.
    Shen B, Li C, Min Z, Meeley RB, Tarczynski MC, Olsen OA. 2003. sal1 determines the number of aleurone cell layers in maize endosperm and encodes a class E vacuolar sorting protein. PNAS 100:6552–57
    [Google Scholar]
  139. 139.
    Shen S, Ma S, Chen X-M, Yi F, Li B-B et al. 2022. A transcriptional landscape underlying sugar import for grain set in maize. Plant J. 110:22842
    [Google Scholar]
  140. 140.
    Shewry PR, Napier JA, Tatham AS. 1995. Seed storage proteins: structures and biosynthesis. Plant Cell 7:945–56
    [Google Scholar]
  141. 141.
    Shimada T, Takagi J, Ichino T, Shirakawa M, Hara-Nishimura I. 2018. Plant vacuoles.. Annu. Rev. Plant Biol. 69:123–45
    [Google Scholar]
  142. 142.
    Slade AJ, Fuerstenberg SI, Loeffler D, Steine MN, Facciotti D. 2005. A reverse genetic, nontransgenic approach to wheat crop improvement by TILLING. Nat. Biotechnol. 23:75–81
    [Google Scholar]
  143. 143.
    Song Y, Luo G, Shen L, Yu K, Yang W et al. 2020. TubZIP28, a novel bZIP family transcription factor from Triticum urartu, and TabZIP28, its homologue from Triticum aestivum, enhance starch synthesis in wheat. New Phytol 226:1384–98
    [Google Scholar]
  144. 144.
    Sosso D, Luo D, Li QB, Sasse J, Yang J et al. 2015. Seed filling in domesticated maize and rice depends on SWEET-mediated hexose transport. Nat. Genet. 47:1489–93
    [Google Scholar]
  145. 145.
    Sparla F, Falini G, Botticella E, Pirone C, Talamè V et al. 2014. New starch phenotypes produced by TILLING in barley. PLOS ONE 9:e107779
    [Google Scholar]
  146. 146.
    Sreenivasulu N, Radchuk V, Alawady A, Borisjuk L, Weier D et al. 2010. De-regulation of abscisic acid contents causes abnormal endosperm development in the barley mutant seg8. Plant J 64:589–603
    [Google Scholar]
  147. 147.
    Strobbe S, Verstraete J, Stove C, Van Der Straeten D. 2021. Metabolic engineering of rice endosperm towards higher vitamin B1 accumulation. Plant Biotechnol. J. 19:1253–67
    [Google Scholar]
  148. 148.
    Su J, Hu C, Yan X, Jin Y, Chen Z et al. 2015. Expression of barley SUSIBA2 transcription factor yields high-starch low-methane rice. Nature 523:602–6
    [Google Scholar]
  149. 149.
    Sun C, Palmqvist S, Olsson H, Boren M, Ahlandsberg S, Jansson C 2003. A novel WRKY transcription factor, SUSIBA2, participates in sugar signaling in barley by binding to the sugar-responsive elements of the iso1 promoter. Plant Cell 15:2076–92
    [Google Scholar]
  150. 150.
    Su'udi M, Cha JY, Ahn IP, Kwak YS, Woo YM, Son D. 2012. Functional characterization of a B-type cell cycle switch 52 in rice (OsCCS52B). Plant Cell Tiss. . Organ. Cult. 111:101–11
    [Google Scholar]
  151. 151.
    Su'udi M, Cha JY, Jung MH, Ermawati N, Han CD et al. 2012. Potential role of the rice OsCCS52A gene in endoreduplication. Planta 235:387–97
    [Google Scholar]
  152. 152.
    Suzuki A, Wu CY, Washida H, Takaiwa F. 1998. Rice MYB protein OsMYB5 specifically binds to the AACA motif conserved among promoters of genes for storage protein glutelin. Plant Cell Physiol 39:555–59
    [Google Scholar]
  153. 153.
    Takemoto Y, Coughlan SJ, Okita TW, Satoh H, Ogawa M, Kumamaru T. 2002. The rice mutant esp2 greatly accumulates the glutelin precursor and deletes the protein disulfide isomerase. Plant Physiol 128:1212–22
    [Google Scholar]
  154. 154.
    Tetlow IJ. 2010. Starch biosynthesis in developing seeds. Seed Sci. Res. 21:5–32
    [Google Scholar]
  155. 155.
    Tian L, Chou HL, Zhang L, Hwang SK, Starkenburg SR et al. 2018. RNA-binding protein RBP-P is required for glutelin and prolamine mRNA localization in rice endosperm cells. Plant Cell 30:2529–52
    [Google Scholar]
  156. 156.
    Tian L, Chou HL, Zhang L, Okita TW. 2019. Targeted endoplasmic reticulum localization of storage protein mRNAs requires the RNA-binding protein RBP-L. Plant Physiol 179:1111–31
    [Google Scholar]
  157. 157.
    Tian Q, Olsen L, Sun B, Lid SE, Brown RC et al. 2007. Subcellular localization and functional domain studies of DEFECTIVE KERNEL1 in maize and Arabidopsis suggest a model for aleurone cell fate specification involving CRINKLY4 and SUPERNUMERARY ALEURONE LAYER1. Plant Cell 19:3127–45
    [Google Scholar]
  158. 158.
    Tonosaki K, Kinoshita T. 2015. Possible roles for polycomb repressive complex 2 in cereal endosperm. Front. Plant Sci. 6:144
    [Google Scholar]
  159. 159.
    Tonosaki K, Ono A, Kunisada M, Nishino M, Nagata H et al. 2021. Mutation of the imprinted gene OsEMF2a induces autonomous endosperm development and delayed cellularization in rice. Plant Cell 33:85–103
    [Google Scholar]
  160. 160.
    Vamvaka E, Arcalis E, Ramessar K, Evans A, O'Keefe BR et al. 2016. Rice endosperm is cost-effective for the production of recombinant griffithsin with potent activity against HIV. Plant Biotechnol. J. 14:1427–37
    [Google Scholar]
  161. 161.
    Vicente-Carbajosa J, Moose SP, Parsons RL, Schmidt RJ. 1997. A maize zinc-finger protein binds the prolamin box in zein gene promoters and interacts with the basic leucine zipper transcriptional activator Opaque2. PNAS 94:7685–90
    [Google Scholar]
  162. 162.
    Vicente-Carbajosa J, Oñate L, Lara P, Diaz I, Carbonero P. 1998. Barley BLZ1: a bZIP transcriptional activator that interacts with endosperm-specific gene promoters. Plant J 13:629–40
    [Google Scholar]
  163. 163.
    Wang C, Washida H, Crofts AJ, Hamada S, Katsube-Tanaka T et al. 2008. The cytoplasmic-localized, cytoskeletal-associated RNA binding protein OsTudor-SN: evidence for an essential role in storage protein RNA transport and localization. Plant J 55:443–54
    [Google Scholar]
  164. 164.
    Wang E, Wang J, Zhu X, Hao W, Wang L et al. 2008. Control of rice grain-filling and yield by a gene with a potential signature of domestication. Nat. Genet. 40:1370–74Identified a grain-vasculature-expressed cell wall invertase that is critical for grain filling in rice.
    [Google Scholar]
  165. 165.
    Wang G, Wang F, Wang G, Wang F, Zhang X et al. 2012. Opaque1 encodes a myosin XI motor protein that is required for endoplasmic reticulum motility and protein body formation in maize endosperm. Plant Cell 24:3447–62
    [Google Scholar]
  166. 166.
    Wang J, Chen Z, Zhang Q, Meng S, Wei C. 2020. The NAC transcription factors OsNAC20 and OsNAC26 regulate starch and storage protein synthesis. Plant Physiol 184:1775–91
    [Google Scholar]
  167. 167.
    Wang JC, Xu H, Zhu Y, Liu QQ, Cai XL. 2013. OsbZIP58, a basic leucine zipper transcription factor, regulates starch biosynthesis in rice endosperm. J. Exp. Bot. 64:3453–66
    [Google Scholar]
  168. 168.
    Wang P, Xia H, Zhang Y, Zhao S, Zhao C et al. 2015. Genome-wide high-resolution mapping of DNA methylation identifies epigenetic variation across embryo and endosperm in Maize (Zea may). BMC Genom 16:21
    [Google Scholar]
  169. 169.
    Wang R, Wang H, Liu X, Ji X, Chen L et al. 2018. Waxy allelic diversity in common millet (Panicum miliaceum L.) in China. Crop J. 6:377–85
    [Google Scholar]
  170. 170.
    Wang Y, Liu F, Ren Y, Wang Y, Liu X et al. 2016. GOLGI TRANSPORT 1B regulates protein export from the endoplasmic reticulum in rice endosperm cells. Plant Cell 28:2850–65
    [Google Scholar]
  171. 171.
    Wang Y, Liu W, Wang H, Du Q, Fu Z et al. 2019. ZmEHD1 is required for kernel development and vegetative growth through regulating auxin homeostasis. Plant Physiol 182:1467–80
    [Google Scholar]
  172. 172.
    Wang Y, Ren Y, Liu X, Jiang L, Chen L et al. 2010. OsRab5a regulates endomembrane organization and storage protein trafficking in rice endosperm cells. Plant J 64:812–24
    [Google Scholar]
  173. 173.
    Washida H, Kaneko S, Crofts N, Sugino A, Wang C, Okita TW 2009. Identification of cis-localization elements that target glutelin RNAs to a specific subdomain of the cortical endoplasmic reticulum in rice endosperm cells. Plant Cell Physiol 50:1710–14
    [Google Scholar]
  174. 174.
    Washida H, Sugino A, Doroshenk KA, Satoh-Cruz M, Nagamine A et al. 2012. RNA targeting to a specific ER sub-domain is required for efficient transport and packaging of α-globulins to the protein storage vacuole in developing rice endosperm. Plant J 70:471–79
    [Google Scholar]
  175. 175.
    Washida H, Sugino A, Messing J, Esen A, Okita TW 2004. Asymmetric localization of seed storage protein RNAs to distinct subdomains of the endoplasmic reticulum in developing maize endosperm cells. Plant Cell Physiol 45:1830–37
    [Google Scholar]
  176. 176.
    Wen S, Wen N, Pang J, Langen G, Brew-Appiah RA et al. 2012. Structural genes of wheat and barley 5-methylcytosine DNA glycosylases and their potential applications for human health. PNAS 109:20543–48
    [Google Scholar]
  177. 177.
    Wu CY, Suzuki A, Washida H, Takaiwa F. 1998. The GCN4 motif in a rice glutelin gene is essential for endosperm-specific gene expression and is activated by Opaque-2 in transgenic rice plants. Plant J 14:673–83
    [Google Scholar]
  178. 178.
    Wu C-Y, Washida H, Onodera Y, Harada K, Takaiwa F. 2000. Quantitative nature of the Prolamin-box, ACGT and AACA motifs in a rice glutelin gene promoter: minimal cis-element requirements for endosperm-specific gene expression. Plant J 23:415–21
    [Google Scholar]
  179. 179.
    Wu H, Gontarek BC, Yi G, Beall BD, Neelakandan AK et al. 2020. The thickaleurone1 gene encodes a NOT1 subunit of the CCR4-NOT complex and regulates cell patterning in endosperm. Plant Physiol 184:960–72
    [Google Scholar]
  180. 180.
    Wu J, Chen L, Chen M, Zhou W, Dong Q et al. 2019. The DOF-domain transcription factor ZmDOF36 positively regulates starch synthesis in transgenic maize. Front. Plant Sci. 10:465
    [Google Scholar]
  181. 181.
    Wu M, Ren Y, Cai M, Wang Y, Zhu S et al. 2019. Rice FLOURY ENDOSPERM10 encodes a pentatricopeptide repeat protein that is essential for the trans-splicing of mitochondrial nad1 intron 1 and endosperm development. New Phytol 223:736–50
    [Google Scholar]
  182. 182.
    Wu MW, Zhao H, Zhang JD, Guo L, Liu CM 2020. RADICLELESS 1 (RL1)-mediated nad4 intron 1 splicing is crucial for embryo and endosperm development in rice (Oryza sativa L.). Biochem. Biophys. Res. Commun. 523:220–25
    [Google Scholar]
  183. 183.
    Wu X, Liu J, Li D, Liu CM 2016. Rice caryopsis development I: dynamic changes in different cell layers. J. Integr. Plant Biol. 58:772–85
    [Google Scholar]
  184. 184.
    Wu X, Liu J, Li D, Liu CM 2016. Rice caryopsis development II: dynamic changes in the endosperm. J. Integr. Plant Biol. 58:786–98Performed a day-by-day cell biological examination of rice endosperm development.
    [Google Scholar]
  185. 185.
    Wu XY, Chen D, Lu YQ, Liu WH, Yang XM et al. 2017. Molecular characteristics of two new waxy mutations in China waxy maize. Mol. Breeding 37:27
    [Google Scholar]
  186. 186.
    Wu Y, Lee S-K, Yoo Y, Wei J, Kwon S-Y et al. 2018. Rice transcription factor OsDOF11 modulates sugar transport by promoting expression of Sucrose Transporter and SWEET genes. Mol. Plant 11:833–45
    [Google Scholar]
  187. 187.
    Xi DM, Zheng CC. 2011. Transcriptional regulation of seed storage protein genes in Arabidopsis and cereals. Seed Sci. Res. 21:247–54
    [Google Scholar]
  188. 188.
    Xi X-Y, Ye B-X. 1995. Studies on endosperm development and deposition of storage reserves in Coix lacryma-jobi. J. Integr. Plant Biol. 37:118–24
    [Google Scholar]
  189. 189.
    Xiao Q, Wang Y, Du J, Li H, Wei B et al. 2017. ZmMYB14 is an important transcription factor involved in the regulation of the activity of the ZmBT1 promoter in starch biosynthesis in maize. FEBS J 284:3079–99
    [Google Scholar]
  190. 190.
    Xu W, Fiume E, Coen O, Pechoux C, Lepiniec L, Magnani E 2016. Endosperm and nucellus develop antagonistically in Arabidopsis seeds. Plant Cell 28:1343–60
    [Google Scholar]
  191. 191.
    Xu X, E Z, Zhang D, Yun Q, Zhou Y et al. 2021. OsYUC11-mediated auxin biosynthesis is essential for endosperm development of rice. Plant Physiol 185:934–50
    [Google Scholar]
  192. 192.
    Yamamoto MP, Onodera Y, Touno SM, Takaiwa F. 2006. Synergism between RPBF Dof and RISBZ1 bZIP activators in the regulation of rice seed expression genes. Plant Physiol 141:1694–707
    [Google Scholar]
  193. 193.
    Yang J, Luo D, Yang B, Frommer WB, Eom JS 2018. SWEET11 and 15 as key players in seed filling in rice. New Phytol 218:604–15
    [Google Scholar]
  194. 194.
    Yang T, Guo L, Ji C, Wang H, Wang J et al. 2021. The B3 domain-containing transcription factor ZmABI19 coordinates expression of key factors required for maize seed development and grain filling. Plant Cell 33:104–28
    [Google Scholar]
  195. 195.
    Yang Y, Crofts AJ, Crofts N, Okita TW 2014. Multiple RNA binding protein complexes interact with the rice prolamine RNA cis-localization zipcode sequences. Plant Physiol 164:1271–82
    [Google Scholar]
  196. 196.
    Yao D, Qi W, Li X, Yang Q, Yan S et al. 2016. Maize opaque10 encodes a cereal-specific protein that is essential for the proper distribution of zeins in endosperm protein bodies. PLOS Genet 12:e1006270
    [Google Scholar]
  197. 197.
    Yi G, Lauter AM, Scott MP, Becraft PW. 2011. The thick aleurone1 mutant defines a negative regulation of maize aleurone cell fate that functions downstream of defective kernel1. Plant Physiol 156:1826–36
    [Google Scholar]
  198. 198.
    Yi G, Neelakandan AK, Gontarek BC, Vollbrecht E, Becraft PW 2015. The naked endosperm genes encode duplicate INDETERMINATE domain transcription factors required for maize endosperm cell patterning and differentiation. Plant Physiol 167:443–56Reported two homologous transcription factors that regulate endosperm patterning and cell differentiation.
    [Google Scholar]
  199. 199.
    Yin LL, Xue HW. 2012. The MADS29 transcription factor regulates the degradation of the nucellus and the nucellar projection during rice seed development. Plant Cell 24:1049–65
    [Google Scholar]
  200. 200.
    Young TE, Gallie DR. 1999. Analysis of programmed cell death in wheat endosperm reveals differences in endosperm development between cereals. Plant Mol. Biol. 39:915–26
    [Google Scholar]
  201. 201.
    Young TE, Gallie DR. 2000. Programmed cell death during endosperm development. Plant Mol. Biol. 44:283–301
    [Google Scholar]
  202. 202.
    Zemach A, Kim MY, Silva P, Rodrigues JA, Dotson B et al. 2010. Local DNA hypomethylation activates genes in rice endosperm. PNAS 107:18729–34
    [Google Scholar]
  203. 203.
    Zhang H, Lang Z, Zhu JK 2018. Dynamics and function of DNA methylation in plants. Nat. Rev. Mol. Cell Biol. 19:489–506
    [Google Scholar]
  204. 204.
    Zhang L, Cheng Z, Qin R, Qiu Y, Wang JL et al. 2012. Identification and characterization of an epi-allele of FIE1 reveals a regulatory linkage between two epigenetic marks in rice. Plant Cell 24:4407–21
    [Google Scholar]
  205. 205.
    Zhang M, Xie S, Dong X, Zhao X, Zeng B et al. 2014. Genome-wide high resolution parental-specific DNA and histone methylation maps uncover patterns of imprinting regulation in maize. Genome Res 24:167–76
    [Google Scholar]
  206. 206.
    Zhang M, Zhao H, Xie S, Chen J, Xu Y et al. 2011. Extensive, clustered parental imprinting of protein-coding and noncoding RNAs in developing maize endosperm. PNAS 108:20042–47
    [Google Scholar]
  207. 207.
    Zhang QF. 2021. Purple tomatoes, black rice and food security. Nat. Rev. Genet. 22:414
    [Google Scholar]
  208. 208.
    Zhang XF, Tong JH, Bai AN, Liu CM, Xiao LT, Xue HW 2020. Phytohormone dynamics in developing endosperm influence rice grain shape and quality. J. Integr. Plant Biol. 62:1625–37
    [Google Scholar]
  209. 209.
    Zhang Z, Dong J, Ji C, Wu Y, Messing J 2019. NAC-type transcription factors regulate accumulation of starch and protein in maize seeds. PNAS 116:11223–28
    [Google Scholar]
  210. 210.
    Zhang Z, Yang J, Wu Y 2015. Transcriptional regulation of zein gene expression in maize through the additive and synergistic action of opaque2, prolamine-box binding factor, and O2 heterodimerizing proteins. Plant Cell 27:1162–72
    [Google Scholar]
  211. 211.
    Zhang Z, Zheng X, Yang J, Messing J, Wu Y 2016. Maize endosperm-specific transcription factors O2 and PBF network the regulation of protein and starch synthesis. PNAS 113:10842–47
    [Google Scholar]
  212. 212.
    Zheng XM, Chen J, Pang HB, Liu S, Gao Q et al. 2019. Genome-wide analyses reveal the role of noncoding variation in complex traits during rice domestication. Sci. Adv. 5:eaax3619
    [Google Scholar]
  213. 213.
    Zhou YF, Zhang YC, Sun YM, Yu Y, Lei MQ et al. 2021. The parent-of-origin lncRNA MISSEN regulates rice endosperm development. Nat. Commun. 12:6525
    [Google Scholar]
  214. 214.
    Zhu C, Jin G, Fang P, Zhang Y, Feng X et al. 2019. Maize pentatricopeptide repeat protein DEK41 affects cis-splicing of mitochondrial nad4 intron 3 and is required for normal seed development. J. Exp. Bot. 70:3795–808
    [Google Scholar]
  215. 215.
    Zhu Q, Yu S, Zeng D, Liu H, Wang H et al. 2017. Development of “Purple Endosperm Rice” by engineering anthocyanin biosynthesis in the endosperm with a high-efficiency transgene stacking system. Mol. Plant 10:918–29
    [Google Scholar]
  216. 216.
    Zhu Y, Cai XL, Wang ZY, Hong MM 2003. An interaction between a MYC protein and an EREBP protein is involved in transcriptional regulation of the rice Wx gene. J. Biol. Chem. 278:47803–11
    [Google Scholar]
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