水稻胚乳发育遗传调控的研究进展

doi:10.16819/j.1001-7216.2021.210307

摘要/Abstract

摘要：

胚乳是被子植物双受精产物之一,为种子发育提供营养;同时,水稻胚乳也是人类口粮的重要来源。胚乳组织约占水稻种子干质量的70%以上,其发育直接影响稻米产量和品质。目前我们对水稻胚乳发育调控的分子机制有了较为深入的认识,克隆了一些重要基因,同时发现表观遗传调控在胚乳发育中也发挥重要作用。本文主要以水稻为例,同时穿插拟南芥和玉米等植物的相关研究进展,系统总结了胚乳细胞化、糊粉层细胞分化、储藏物质积累等胚乳发育重要生物学事件的遗传调控机制。最后我们也指出了关于胚乳发育过程中有待进一步深入研究的科学问题,以期能为今后相关研究提供一些思路。

关键词: 水稻, 胚乳发育, 遗传调控, 细胞分化, 基因组印记

Abstract:

Endosperm, a product of double fertilization, is a tissue nourishing the developing embryo. In addition, cereal endosperm is the primary source of calories of humans. Rice endosperm consists more than seventy percent of the seed dry mass, whose development directly determines the yield and quality of rice grains. To date, we have had a profound understanding of the genetic and epigenetic regulations for rice endosperm development. Many of the important genes required for rice endosperm development have been identified. In this review, we summarize the molecular controls of the key events/processes of endosperm development, including the syncytium-cellualrization transition, endosperm cell differentiation and storage compounds accumulation. We mainly focus on the new findings in rice; however, some important findings in Arabidopsis and other cereal crops are also introduced. We also discuss some questions that need to be elucidated in the future for rice endosperm development.

Key words: rice, endosperm development, genetic regulation, cell differentiation, genomic imprinting

张娟, 牛百晓, 鄂志国, 陈忱. 水稻胚乳发育遗传调控的研究进展[J]. 中国水稻科学, 2021, 35(4): 326-341.

Juan ZHANG, Baixiao NIU, Zhiguo E, Chen CHEN. Towards Understanding the Genetic Regulations of Endosperm Development in Rice[J]. Chinese Journal OF Rice Science, 2021, 35(4): 326-341.

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参考文献 137

[1]	Wu X, Liu J, Li D, Liu C M.Rice caryopsis development: I. Dynamic changes in different cell layers[J]. Journal of Integrative Plant Biology, 2016, 58(9): 772-785.
[2]	Zhang M, Xie S, Dong X, Zhao X, Zeng B, Chen J, Li H, Yang W, Zhao H, Wang G, Chen Z, Sun S, Hauck A, Jin W, Lai J.Genome-wide high resolution parental-specific DNA and histone methylation maps uncover patterns of imprinting regulation in maize[J]. Genome Research, 2014, 24(1): 167-176.
[3]	Zemach A, Kim M Y, Silva P, Rodrigues J A, Dotson B, Brooks M D, Zilberman D.Local DNA hypomethylation activates genes in rice endosperm[J]. Proceedings of the National Academy of Sciences, 2010, 107(43): 18729-18734.
[4]	Hsieh T F, Ibarra C I, Silva P, Zemach A, Eshed-Williams L, Fischer R L, Zilberman D.Genome- wide demethylation of Arabidopsis endosperm[J]. Science, 2009, 324: 1451-1454.
[5]	Wu X, Liu J, Li D, Liu C M.Rice caryopsis development II: Dynamic changes in the endosperm[J]. Journal of Integrative Plant Biology, 2016, 58(9): 786-798.
[6]	Chen C, Begcy K, Liu K, Folsom J J, Wang Z, Zhang C, Walia H.Heat stress yields a unique MADS box transcription factor in determining seed size and thermal sensitivity[J]. Plant Physiology, 2016, 171(1): 606-622.
[7]	Paul P, Dhatt B K, Sandhu J, Hussain W, Irvin L, Morota G, Staswick P, Walia H.Divergent phenotypic response of rice accessions to transient heat stress during early seed development[J]. Plant Direct, 2020, 4(1): 1-13.
[8]	Zhang H, Luo M, Johnson S D, Zhu X, Liu L, Huang F, Liu Y, Xu P, Wu X.Parental genome imbalance causes post-zygotic seed lethality and deregulates imprinting in rice[J]. Rice, 2016, 9(1): 43.
[9]	Sekine D, Ohnishi T, Furuumi H, Ono A, Yamada T, Kurata N, Kinoshita T.Dissection of two major components of the post-zygotic hybridization barrier in rice endosperm[J]. The Plant Journal, 2013, 76(5): 792-799.
[10]	Tonosaki K, Sekine D, Ohnishi T, Ono A, Furuumi H, Kurata N, Kinoshita T.Overcoming the species hybridization barrier by ploidy manipulation in the genus Oryza[J]. Plant Journal, 2018, 93(3): 534-544.
[11]	Ishikawa R, Ohnishi T, Kinoshita Y, Eiguchi M, Kurata N, Kinoshita T.Rice interspecies hybrids show precocious or delayed developmental transitions in the endosperm without change to the rate of syncytial nuclear division[J]. The Plant Journal, 2011, 65(5): 798-806.
[12]	Mozgova I, Hennig L.The polycomb group protein regulatory network[J]. Annual Review of Plant Biology, 2015, 66(1): 269-296.
[13]	Holec S, Berger F.Polycomb group complexes mediate developmental transitions in plants[J]. Plant Physiology, 2012, 158(1): 35-43.
[14]	Ohad N, Margossian L, Hsu Y C, Williams C, Repetti P, Fischer R L.A mutation that allows endosperm development without fertilization[J]. Proceedings of the National Academy of Sciences of the United States of America, 1996, 93(11): 5319-5324.
[15]	Eacock W J A P, Chaudhury A M, Ming L, Miller C, Craig S, Dennis E S, Peacock W J. Fertilization- independent seed development in Arabidopsis thaliana[J]. Proceedings of the National Academy of Sciences of the United States of America, 1997, 94: 4223-4228.
[16]	Kiyosue T, Ohad N, Yadegari R, Hannon M, Dinneny J, Wells D, Katz A, Margossian L, Harada J J, Goldberg R B, Fischer R L.Control of fertilization-independent endosperm development by the MEDEA polycomb gene in Arabidopsis[J]. Proceedings of the National Academy of Sciences, 1999, 96(7): 4186-4191.
[17]	Köhler C, Hennig L, Bouveret R, Gheyselinck J, Grossniklaus U, Gruissem W.Arabidopsis MSI1 is a component of the MEA/FIE Polycomb group complex and required for seed development[J]. The EMBO Journal, 2003, 22(18): 4804-4814.
[18]	Luo M, Platten D, Chaudhury A, Peacock W J, Dennis E S.Expression, imprinting, and evolution of rice homologs of the polycomb group genes[J]. Molecular Plant, 2009, 2(4): 711-723.
[19]	Li S, Zhou B, Peng X, Kuang Q, Huang X, Yao J, Du B, Sun M X.OsFIE2 plays an essential role in the regulation of rice vegetative and reproductive development[J]. New Phytologist, 2014, 201(1): 66-79.
[20]	Zhang L, Cheng Z, Qin R, Qiu Y, Wang J L, Cui X, Gu L, Zhang X, Guo X, Wang D, Jiang L, Wu C Y, Wang H, Cao X, Wan J.Identification and characterization of an Epi-allele of FIE1 reveals a regulatory linkage between two epigenetic marks in rice[J]. The Plant Cell, 2012, 24(11): 4407-4421.
[21]	Cheng X, Pan M, E Z, Zhou Y, Niu B, Chen C, Cheng X, Niu B, Chen C, Pan M, Zhiguo E, Zhou Y, Niu B, Chen C,. Functional divergence of two duplicated fertilization independent endosperm genes in rice with respect to seed development[J]. The Plant Journal, 2020, 104(1): 124-137.
[22]	Furihata H Y, Suenaga K, Kawanabe T, Yoshida T, Kawabe A.Gene duplication, silencing and expression alteration govern the molecular evolution of PRC2 genes in plants[J]. Genes & Genetic Systems, 2016, 91(2): 85-95.
[23]	Tonosaki K, Kinoshita T.Possible roles for polycomb repressive complex 2 in cereal endosperm[J]. Frontiers in Plant Science, 2015, 6: 1-5.
[24]	Cheng X, Pan M, E Z G, Zhou Y, Niu B, Chen C. The maternally expressed polycomb group gene OsEMF2a is essential for endosperm cellularization and imprinting in rice[J]. Plant Communications, 2020, 2: 100092.
[25]	Tonosaki K, Ono A, Kunisada M, Nishino M, Nagata H, Sakamoto S, Kijima S T, Furuumi H, Nonomura K I, Sato Y, Ohme-Takagi M, Endo M, Comai L, Hatakeyama K, Kawakatsu T, Kinoshita T.Mutation of the imprinted gene OsEMF2a induces autonomous endosperm development and delayed cellularization in rice[J]. The Plant Cell, 2021, 33(1): 85-103.
[26]	Deng L, Zhang S, Wang G, Fan S, Li M, Chen W, Tu B, Tan J, Wang Y, Ma B, Li S, Qin P.Down-regulation of OsEMF2b caused semi-sterility due to anther and pollen development defects in rice[J]. Frontiers in Plant Science, 2017, 8: 1998.
[27]	Xie S, Chen M, Pei R, Ouyang Y, Yao J.OsEMF2b acts as a regulator of flowering transition and floral organ identity by mediating H3K27me3 deposition at OsLFL1 and OsMADS4 in rice[J]. Plant Molecular Biology Reporter, 2015, 33(1): 121-132.
[28]	Conrad L J, Khanday I, Johnson C, Guiderdoni E, An G, Vijayraghavan U, Sundaresan V.The polycomb group gene EMF2B is essential for maintenance of floral meristem determinacy in rice[J]. Plant Journal, 2014, 80(5): 883-894.
[29]	Chen M, Xie S, Ouyang Y, Yao J.Rice PcG gene OsEMF2b controls seed dormancy and seedling growth by regulating the expression of OsVP1[J]. Plant Science, 2017, 260: 80-89.
[30]	Zhong J, Peng Z, Peng Q, Cai Q, Peng W, Chen M, Yao J.Regulation of plant height in rice by the Polycomb group genes OsEMF2b, OsFIE2 and OsCLF[J]. Plant Science, 2018, 267: 157-167.
[31]	Smaczniak C, Immink R G H, Angenent G C, Kaufmann K. Developmental and evolutionary diversity of plant MADS-domain factors: Insights from recent studies[J]. Development, 2012, 139(17): 3081-3098.
[32]	Masiero S, Colombo L, Grini P E, Schnittger A, Kater M M.The emerging importance of type I MADS box transcription factors for plant reproduction[J]. The Plant Cell, 2011, 23(3): 865-872.
[33]	Kang I H, Steffen J G, Portereiko M F, Lloyd A, Drews G N.The AGL62 MADS domain protein regulates cellularization during endosperm development in Arabidopsis[J]. The Plant Cell, 2008, 20(3): 635-647.
[34]	Portereiko M F, Lloyd A, Steffen J G, Punwani J A, Otsuga D, Drews G N.AGL80 is required for central cell and endosperm development in Arabidopsis[J]. The Plant Cell, 2006, 18(8): 1862-1872.
[35]	Kohler C.The Polycomb-group protein MEDEA regulates seed development by controlling expression of the MADS-box gene PHERES1[J]. Genes & Development, 2003, 17(12): 1540-1553.
[36]	Walia H, Josefsson C, Dilkes B, Kirkbride R, Harada J, Comai L.Dosage-dependent deregulation of an AGAMOUS-LIKE gene cluster contributes to interspecific incompatibility[J]. Current Biology, 2009, 19(13): 1128-1132.
[37]	Nam J, Kim J, Lee S, An G, Ma H, Nei M.Type I MADS-box genes have experienced faster birth-and -death evolution than type II MADS-box genes in angiosperms[J]. Proceedings of the National Academy of Sciences, 2004, 101(7): 1910-1915.
[38]	Gramzow L, Theißen G.Phylogenomics of MADS-box genes in plants: Two opposing life styles in one gene family[J]. Biology, 2013, 2(3): 1150-1164.
[39]	Paul P, Dhatt B K, Miller M, Folsom J J, Wang Z, Krassovskaya I, Liu K, Sandhu J, Yu H, Zhang C, Obata T, Staswick P, Walia H.MADS78 and MADS79 are essential regulators of early seed development in rice[J]. Plant Physiology, 2020, 182(2): 933-948.
[40]	Olsen O A.ENDOSPERM DEVELOPMENT: Cellularization and cell fate specification[J]. Annual Review of Plant Physiology and Plant Molecular Biology, 2001, 52(1): 233-267.
[41]	Dante R, Larkins B, Sabelli P.Cell cycle control and seed development[J]. Frontiers in Plant Science, 2014, 5: 493.
[42]	Guo J, Wang F, Song J, Sun W, Zhang X S.The expression of Orysa;CycB1;1 is essential for endosperm formation and causes embryo enlargement in rice[J]. Planta, 2010, 231(2): 293-303.
[43]	Mizutani M, Naganuma T, Tsutsumi K, Saitoh Y.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]. Journal of Experimental Botany, 2010, 61(3): 791-798.
[44]	Barrôco R M, Peres A, Droual A M, De Veylder L, Nguyen L S L, De Wolf J, Mironov V, Peerbolte R, Beemster G T S, Inzé D, Broekaert W F, Frankard V. The cyclin-dependent kinase inhibitor Orysa;KRP1 plays an important role in seed development of rice[J]. Plant Physiology, 2006, 142(3): 1053-1064.
[45]	Hara T, Katoh H, Ogawa D, Kagaya Y, Sato Y, Kitano H, Nagato Y, Ishikawa R, Ono A, Kinoshita T, Takeda S, Hattori T.Rice SNF2 family helicase ENL1 is essential for syncytial endosperm development[J]. The Plant Journal, 2015, 81(1): 1-12.
[46]	Huang X, Peng X, Sun M X. OsGCD1 is essential for rice fertility and required for embryo dorsal-ventral pattern formation and endosperm development[J]. New Phytologist, 2017, 215(3): 1039-1058.
[47]	Becraft P W, Yi G.Regulation of aleurone development in cereal grains[J]. Journal of Experimental Botany, 2011, 62(5): 1669-1675.
[48]	Yan D, Duermeyer L, Leoveanu C, Nambara E.The functions of the endosperm during seed germination[J]. Plant & Cell Physiology, 2014, 55(9): 1521-1533.
[49]	Wu H, Gontarek B C, Yi G, Beall B D, Neelakandan A K, Adhikari B, Chen R, McCarty D R, Severin A J, Becraft P W. The thick aleurone1 gene encodes a NOT1 subunit of the CCR4-NOT complex and regulates cell patterning in endosperm[J]. Plant Physiology, 2020, 184: 00703.2020. DOI: 10.1104/pp.20.00703
[50]	Lid S E, Gruis D, Jung R, Lorentzen J A, Ananiev E, Chamberlin M, Niu X, Meeley R, Nichols S, Olsen O A.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[J]. Proceedings of the National Academy of Sciences of the United States of America, 2002, 99(8): 5460-5465.
[51]	Yi G, Neelakandan A K, Gontarek B C, Vollbrecht E, Becraft P W.The naked endosperm genes encode duplicate INDETERMINATE domain transcription factors required for maize endosperm cell patterning and differentiation[J]. Plant Physiology, 2015, 167(2): 443-456.
[52]	Gontarek B C, Neelakandan A K, Wu H, Becraft P W.NKD transcription factors are central regulators of maize endosperm development[J]. Plant Cell, 2016, 28(12): 2916-2936.
[53]	Pu C X, Ma Y, Wang J, Zhang Y C, Jiao X W, Hu Y H, Wang L L, Zhu Z G, Sun D, Sun Y.Crinkly4 receptor-like kinase is required to maintain the interlocking of the palea and lemma, and fertility in rice, by promoting epidermal cell differentiation[J]. The Plant Journal, 2012, 70(6): 940-953.
[54]	Hibara K ichiro, Obara M, Hayashida E, Abe M, Ishimaru T, Satoh H, Itoh J, Nagato Y. The ADAXIALIZED LEAF1 gene functions in leaf and embryonic pattern formation in rice[J]. Developmental Biology, 2009, 334(2): 345-354.
[55]	Kawakatsu T, Yamamoto M P, Touno S M, Yasuda H, Takaiwa F.Compensation and interaction between RISBZ1 and RPBF during grain filling in rice[J]. The Plant Journal, 2009, 59(6): 908-920.
[56]	Qi X, Li S, Zhu Y, Zhao Q, Zhu D, Yu J.ZmDof3, a maize endosperm-specific Dof protein gene, regulates starch accumulation and aleurone development in maize endosperm[J]. Plant Molecular Biology, 2017, 93(1-2): 7-20.
[57]	Liu J, Wu X, Yao X, Yu R, Larkin P J, Liu C M.Mutations in the DNA demethylase OsROS1 result in a thickened aleurone and improved nutritional value in rice grains[J]. Proceedings of the National Academy of Sciences, 2018, 115(44): 11327-11332.
[58]	Liu E, Zeng S, Zhu S, Liu Y, Wu G, Zhao K, Liu X, Liu Q, Dong Z, Dang X, Xie H, Li D, Hu X, Hong D.Favorable alleles of GRAIN-FILLING RATE1 increase the grain-filling rate and yield of rice[J]. Plant Physiology, 2019, 181(3): 1207-1222.
[59]	Scofield G N, Hirose T, Gaudron J A, Upadhyaya N M, Ohsugi R, Furbank R T.Antisense suppression of the rice sucrose transporter gene, OsSUT1, leads to impaired grain filling and germination but does not affect photosynthesis[J]. Functional Plant Biology, 2002, 29(7): 815-826.
[60]	Sosso D, Luo D, Li Q B, Sasse J, Yang J, Gendrot G, Suzuki M, Koch K E, McCarty D R, Chourey P S, Rogowsky P M, Ross-Ibarra J, Yang B, Frommer W B. Seed filling in domesticated maize and rice depends on SWEET-mediated hexose transport[J]. Nature Genetics, 2015, 47(12): 1489-1493.
[61]	Ma L, Zhang D, Miao Q, Yang J, Xuan Y, Hu Y.Essential Role of Sugar Transporter OsSWEET11 during the early stage of rice grain filling[J]. Plant and Cell Physiology, 2017, 58(5): 863-873.
[62]	Yang J, Luo D, Yang B, Frommer W B, Eom J S.SWEET11 and 15 as key players in seed filling in rice[J]. The New Phytologist, 2018, 218(2): 604-615.
[63]	Wang E, Wang J, Zhu X, Hao W, Wang L, Li Q, Zhang L, He W, Lu B, Lin H, Ma H, Zhang G, He Z.Control of rice grain-filling and yield by a gene with a potential signature of domestication[J]. Nature Genetics, 2008, 40(11): 1370-1374.
[64]	Asano T, Kunieda N, Omura Y, Ibe H, Kawasaki T, Takano M, Sato M, Furuhashi H, Mujin T, Takaiwa F, Wu Cy C, Tada Y, Satozawa T, Sakamoto M, Shimada H.Rice SPK, a calmodulin-like domain protein kinase, is required for storage product accumulation during seed development: phosphorylation of sucrose synthase is a possible factor[J]. The Plant Cell, 2002, 14(3): 619-628.
[65]	Jeon J S, Ryoo N, Hahn T R, Walia H, Nakamura Y.Starch biosynthesis in cereal endosperm[J]. Plant Physiology and Biochemistry, Elsevier Masson SAS, 2010, 48(6): 383-392.
[66]	Wei X, Jiao G, Lin H, Sheng Z, Shao G, Xie L, Tang S, Xu Q, Hu P.GRAIN INCOMPLETE FILLING 2 regulates grain filling and starch synthesis during rice caryopsis development[J]. Journal of Integrative Plant Biology, 2017, 59(2): 134-153.
[67]	Wang J C C, Xu H, Zhu Y, Liu Q Q Q, Cai X L L. OsbZIP58, a basic leucine zipper transcription factor, regulates starch biosynthesis in rice endosperm[J]. Journal of Experimental Botany, 2013, 64(11): 3453-3466.
[68]	Yin L L, Xue H W.The MADS29 transcription factor regulates the degradation of the nucellus and the nucellar projection during rice seed development[J]. The Plant Cell, 2012, 24(3): 1049-1065.
[69]	Fu F F, Xue H W.Coexpression analysis identifies Rice Starch Regulator1, a rice AP2/EREBP family transcription factor, as a novel rice starch biosynthesis regulator[J]. Plant Physiology, 2010, 154(2): 927-938.
[70]	Laloum T, De Mita S, Gamas P, Baudin M, Niebel A.CCAAT-box binding transcription factors in plants: Y so many?[J]. Trends in Plant Science, 2013, 18(3): 157-166.
[71]	Zhiguo E, Li T, Zhang H, Liu Z, Deng H, Sharma S, Wei X, Wang L, Niu B, Chen C.A group of nuclear factor y transcription factors are sub-functionalized during endosperm development in monocots[J]. Journal of Experimental Botany, 2018, 69(10): 2495-2510.
[72]	Xu J J, Zhang X F, Xue H W.Rice aleurone layer specific OsNF-YB1 regulates grain filling and endosperm development by interacting with an ERF transcription factor[J]. Journal of Experimental Botany, 2016, 67(22): 6399-6411.
[73]	Bello B K, Hou Y, Zhao J, Jiao G, Wu Y, Li Z, Wang Y, Tong X, Wang W, Yuan W, Wei X, Zhang J.NF-YB1-YC12-bHLH144 complex directly activates Wx to regulate grain quality in rice (Oryza sativa L.)[J]. Plant Biotechnology Journal, 2019, 17(7): 1222-1235.
[74]	Niu B, Deng H, Li T, Sharma S, Yun Q, Li Q, E Z G, Chen C. OsbZIP76 interacts with OsNF-YBs and regulates endosperm cellularization in rice(Oryza sativa)[J]. Journal of Integrative Plant Biology, 2020, 62(12): 1983-1996.
[75]	Bai A N, Lu X D, Li D Q, Liu J X, Liu C M.NF-YB1-regulated expression of sucrose transporters in aleurone facilitates sugar loading to rice endosperm[J]. Cell Research, 2016, 26(3): 384-388.
[76]	Pelletier J M, Kwong R W, Park S, Le B H, Baden R, Cagliari A, Hashimoto M, Munoz M D, Fischer R L, Goldberg R B, Harada J J.LEC1 sequentially regulates the transcription of genes involved in diverse developmental processes during seed development[J]. Proceedings of the National Academy of Sciences, 2017, 114(32): E6710-E6719.
[77]	Jo L, Pelletier J M, Hsu S, Baden R, Goldberg R B, Harada J J.Combinatorial interactions of the LEC1 transcription factor specify diverse developmental programs during soybean seed development[J]. Proceedings of the National Academy of Sciences, 2020, 117(2): 1223-1232.
[78]	Zhang Z, Dong J, Ji C, Wu Y, Messing J.NAC-type transcription factors regulate accumulation of starch and protein in maize seeds[J]. Proceedings of the National Academy of Sciences, 2019, 116(23): 11223-11228.
[79]	Wang J, Chen Z, Zhang Q, Meng S, Wei C.The NAC transcription factors OsNAC20 and OsNAC26 regulate starch and storage protein synthesis[J]. Plant Physiology, 2020, 184(4): 1775-1791.
[80]	Ren Y, Huang Z, Jiang H, Wang Z, Wu F, Xiong Y, Yao J.A heat stress responsive NAC transcription factor heterodimer plays key roles in rice grain filling[J]. Journal of Experimental Botany, 2021, 72(8): 2947-2964.
[81]	Mathew I E, Priyadarshini R, Mahto A, Jaiswal P, Parida S K, Agarwal P.SUPER STARCHY1/ONAC025 participates in rice grain filling[J]. Plant Direct, 2020, 4(9): 1-25.
[82]	Zhou H, Xia D, He Y Q.Rice grain quality-traditional traits for high quality rice and health-plus substances[J]. Molecular Breeding, 2020, 40(1): 1. DOI:10.1007/ s11032-019-1080-6.
[83]	Liu L, Waters D L E, Rose T J, Bao J, King G J. Phospholipids in rice: Significance in grain quality and health benefits: A review[J]. Food Chemistry, 2013, 139(1-4): 1133-1145
[84]	Hu Z L, Li P, Zhou M Q, Zhang Z H, Wang L X, Zhu L H, Zhu Y G.Mapping of quantitative trait loci (QTLs) for rice protein and fat content using doubled haploid lines[J]. Euphytica, 2004, 135(1-2): 47-54.
[85]	Liu W, Zeng J, Jiang G, He Y.QTLs identification of crude fat content in brown rice and its genetic basis analysis using DH and two backcross populations[J]. Euphytica, 2009, 169(2): 197-205.
[86]	Qin Y, Kim S M, Zhao X, Lee H S, Jia B, Kim K M, Eun M Y, Sohn J K.QTL detection and MAS selection efficiency for lipid content in brown rice (Oryza sativa L.)[J]. Genes and Genomics, 2010, 32(6): 506-512.
[87]	Liu H L, Yin Z J, Xiao L, Xu Y N, Qu L Q.Identification and evaluation of ω-3 fatty acid desaturase genes for hyperfortifying α-linolenic acid in transgenic rice seed[J]. Journal of Experimental Botany, 2012, 63(8): 3279-3287.
[88]	Zaplin E S, Liu Q, Li Z, Butardo V M, Blanchard C L, Rahman S.Production of high oleic rice grains by suppressing the expression of the OsFAD2-1 gene[J]. Functional Plant Biology, 2013, 40(10): 996-1004.
[89]	Zhou H, Xia D, Li P, Ao Y, Xu X, Wan S, Li Y, Wu B, Shi H, Wang K, Gao G, Zhang Q, Wang G, Xiao J, Li X, Yu S, Lian X, He Y.Genetic architecture and key genes controlling the diversity of oil composition in rice grains[J]. Molecular Plant, 2021, 14(3): 456-469.
[90]	Zhang X F, Tong J H, Bai A N, Liu C M, Xiao L T, Xue H W.Phytohormone dynamics in developing endosperm influence rice grain shape and quality[J]. Journal of Integrative Plant Biology, 2020, 62(10): 1625-1637.
[91]	Zhao Y.Auxin biosynthesis[J]. The Arabidopsis Book, 2014, 12: e0173.
[92]	Figueiredo D D, Batista R A, Roszak P J, Köhler C.Auxin production couples endosperm development to fertilization[J]. Nature Plants, 2015, 1: 15184.
[93]	Batista R A, Figueiredo D D, Santos-González J, Köhler C.Auxin regulates endosperm cellularization in Arabidopsis[J]. Genes & Development, 2019, 33(7-8): 466-476.
[94]	Abu-Zaitoon Y M, Bennett K, Normanly J, Nonhebel H M. A large increase in IAA during development of rice grains correlates with the expression of tryptophan aminotransferase OsTAR1 and a grain-specific YUCCA[J]. Physiologia Plantarum, 2012, 146(4): 487-499.
[95]	Xu X, Zhang D, Niu B, Chen C, Yun Q, Zhou Y.OsYUC11-mediated auxin biosynthesis is essential for endosperm development of rice[J]. Plant Physiology, 2021, 185(3): 934-950.
[96]	Ishimaru K, Hirotsu N, Madoka Y, Murakami N, Hara N, Onodera H, Kashiwagi T, Ujiie K, Shimizu B I, Onishi A, Miyagawa H, Katoh E.Loss of function of the IAA-glucose hydrolase gene TGW6 enhances rice grain weight and increases yield[J]. Nature Genetics, 2013, 45(6): 707-711.
[97]	Lur H S, Setter T L.Role of auxin in maize endosperm development (timing of nuclear DNA endoreduplication, zein expression, and cytokinin)[J]. Plant Physiology, 1993, 103(1): 273-280.
[98]	Bernardi J, Lanubile A, Li Q B, Kumar D, Kladnik A, Cook S D, Ross J J, Marocco A, Chourey P S.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[J]. Plant Physiology, 2012, 160(3): 1318-1328.
[99]	Forestan C, Meda S, Varotto S.ZmPIN1-mediated auxin transport is related to cellular differentiation during maize embryogenesis and endosperm development[J]. Plant Physiology, 2010, 152(3): 1373-1390.
[100]	Wang Z, Xu Y, Chen T, Zhang H, Yang J, Zhang J.Abscisic acid and the key enzymes and genes in sucrose-to-starch conversion in rice spikelets in response to soil drying during grain filling[J]. Planta, 2015, 241(5): 1091-1107.
[101]	Zhang D, Zhang M, Zhou Y, Wang Y, Shen J, Chen H, Zhang L, Lü B, Liang G, Liang J.The rice G protein γ subunit DEP1/qPE9-1 positively regulates grain-filling process by increasing auxin and cytokinin content in rice grains[J]. Rice, 2019, 12(1): 91.
[102]	Qin P, Zhang G, Hu B, Wu J, Chen W, Ren Z, Liu Y, Xie J, Yuan H, Tu B, Ma B, Wang Y, Ye L, Li L, Xiang C, Li S. Leaf-derived ABA regulates rice seed development via a transporter-mediated and temperature-sensitive mechanism[J]. Science Advances, 2021, 7(3): eabc8873.
[103]	Sreenivasulu N, Radchuk V, Alawady A, Borisjuk L, Weier D, Staroske N, Fuchs J, Miersch O, Strickert M, Usadel B, Wobus U, Grimm B, Weber H, Weschke W.De-regulation of abscisic acid contents causes abnormal endosperm development in the barley mutant seg8[J]. Plant Journal, 2010, 64(4): 589-603.
[104]	Xing M Q, Zhang Y J, Zhou S R, Hu W Y, Wu X X, Ye Y J, Wu X T, Xiao Y P, Li X, Xue H W.Global analysis reveals the crucial roles of DNA methylation during rice seed development[J]. Plant Physiology, 2015, 168(4): 1417-1432.
[105]	Yang C, Ma B, He S, Xiong Q, Duan K, Yin C, Chen H, Lu X, Chen S, Zhang J.MAOHUZI6/ETHYLENE INSENSITIVE3-LIKE1 and ETHYLENE INSENSITIVE3- LIKE2 regulate ethylene response of roots and coleoptiles and negatively affect salt tolerance in rice[J]. Plant Physiology, 2015, 169(1): 148-165.
[106]	Yin C, Zhao H, Ma B, Chen S, Zhang J.Diverse roles of ethylene in regulating agronomic traits in rice[J]. Frontiers in Plant Science, 2017, 8: 1676. DOI:10.3389/fpls.2017.01676.
[107]	Huang X, Lu Z, Wang X, Ouyang Y, Chen W, Xie K, Wang D, Luo M, Luo J, Yao J.Imprinted gene OsFIE1 modulates rice seed development by influencing nutrient metabolism and modifying genome H3K27me3[J]. The Plant Journal, 2016, 87(3): 305-317.
[108]	Liu X, Wang P.SDG711 is involved in rice seed development through regulation of starch metabolism gene expression in coordination with other histone modi cations[J]. Rice, 2021, 14(1): 25.
[109]	Qin F J, Sun Q W, Huang L M, Chen X S, Zhou D X.Rice SUVH histone methyltransferase genes display specific functions in chromatin modification and retrotransposon repression[J]. Molecular Plant, 2010, 3(4): 773-782.
[110]	Qi D, Wen Q, Meng Z, Yuan S, Guo H, Zhao H, Cui S.OsLFR is essential for early endosperm and embryo development by interacting with SWI/SNF complex members in Oryza sativa[J]. Plant Journal, 2020, 104(4): 901-916.
[111]	Zhang H, Lu Y, Zhao Y, Zhou D X.OsSRT1 is involved in rice seed development through regulation of starch metabolism gene expression[J]. Plant Science, 2016, 248: 28-36.
[112]	La H, Ding B, Mishra G P, Zhou B, Yang H, Bellizzi M D R, Chen S, Meyers B C, Peng Z, Zhu J K, Wang G L. A 5-methylcytosine DNA glycosylase/lyase demethylates the retrotransposon Tos17 and promotes its transposition in rice[J]. Proceedings of the National Academy of Sciences USA, 2011, 108(37): 15498-15503.
[113]	Köhler C, Wolff P, Spillane C.Epigenetic mechanisms underlying genomic imprinting in plants[J]. Annual Review of Plant Biology, 2012, 63(1): 331-352.
[114]	Kermicle J L.Dependence of the R-mottled aleurone phenotype in maize on mode of sexual transmission[J]. Genetics, 1970, 66(1): 69-85.
[115]	Gehring M, Missirian V, Henikoff S.Genomic analysis of parent-of-origin allelic expression in Arabidopsis thaliana seeds[J]. PLoS ONE, 2011, 6(8): e23687. DOI: 10.1371/journal.pone.0023687
[116]	Yang G, Liu Z, Gao L, Yu K, Feng M, Yao Y, Peng H, Hu Z, Sun Q, Ni Z, Xin M.Genomic imprinting was evolutionarily conserved during wheat polyploidization[J]. Plant Cell, 2018, 30(1): 37-47.
[117]	Luo M, Taylor J M, Spriggs A, Zhang H, Wu X, Russell S, Singh M, Koltunow A.A genome-wide survey of imprinted genes in rice seeds reveals imprinting primarily occurs in the endosperm[J]. PLoS Genet, 2011, 7(6): e1002125.
[118]	Chen C, Li T, Zhu S, Liu Z, Shi Z, Zheng X, Chen R, Huang J, Shen Y, Luo S, Wang L, Liu Q Q, E Z G. Characterization of imprinted genes in rice reveals conservation of regulation and imprinting with other plant species[J]. Plant Physiology, 2018, 177(4): 1754-1771.
[119]	Florez-Rueda A M, Paris M, Schmidt A, Widmer A, Grossniklaus U, Städler T. Genomic imprinting in the endosperm is systematically perturbed in abortive hybrid tomato seeds[J]. Molecular Biology and Evolution, 2016, 33(11): 2935-2946.
[120]	Xu W, Dai M, Li F, Liu A.Genomic imprinting, methylation and parent-of-origin effects in reciprocal hybrid endosperm of castor bean[J]. Nucleic Acids Research, 2014, 42(11): 6987-6998.
[121]	Hsieh T-F F, Shin J, Uzawa R, Silva P, Cohen S, Bauer M J, Hashimoto M, Kirkbride R C, Harada J J, Zilberman D, Fischer R L. Regulation of imprinted gene expression in Arabidopsis endosperm[J]. Proceedings of the National Academy of Sciences of the United States of America, 2011, 108(5): 1755-1762.
[122]	Waters A J, Bilinski P, Eichten S R, Vaughn M W, Ross-Ibarra J, Gehring M, Springer N M.Comprehensive analysis of imprinted genes in maize reveals allelic variation for imprinting and limited conservation with other species[J]. Proceedings of the National Academy of Sciences, 2013, 110(48): 19639-19644.
[123]	Zhang M, Li N, He W, Zhang H, Yang W, Liu B.Genome-wide screen of genes imprinted in sorghum endosperm and the roles of allelic differential cytosine methylation[J]. The Plant Journal, 2015: 424-436.
[124]	Waters A J, Makarevitch I, Eichten S R, Swanson-Wagner R A, Yeh C T, Xu W, Schnable P S, Vaughn M W, Gehring M, Springer N M. Parent-of -origin effects on gene expression and DNA methylation in the maize endosperm[J]. The Plant Cell, 2011, 23(12): 4221-4233.
[125]	Hatorangan M R, Laenen B, Steige K, Slotte T, Köhler C.Rapid evolution of genomic imprinting in two species of the brassicaceae[J]. The Plant Cell, 2016, 28(8): 1815-1827.
[126]	Pignatta D, Erdmann R M, Scheer E, Picard C L, Bell G W, Gehring M.Natural epigenetic polymorphisms lead to intraspecific variation in Arabidopsis gene imprinting[J]. eLife, 2014: e03198.
[127]	Batista R A, Köhler C.Genomic imprinting in plants-revisiting existing models[J]. Genes & Development, 2020, 34(1-2): 24-36.
[128]	Huh J H, Bauer M J, Hsieh T F, Fischer R L.Cellular programming of plant gene imprinting[J]. Cell, 2008, 132(5): 735-744.
[129]	Burkart-Waco D, Ngo K, Lieberman M, Comai L.Perturbation of parentally biased gene expression during interspecific hybridization[J]. PloS ONE, 2015, 10(2): e0117293.
[130]	Kradolfer D, Wolff P, Jiang H, Siretskiy A, Kohler C.An imprinted gene underlies postzygotic reproductive isolation in Arabidopsis thaliana[J]. Development Cell, 2013, 26(5): 525-535.
[131]	Jullien P E, Berger F.Parental genome dosage imbalance deregulates imprinting in Arabidopsis[J]. PLoS Genetics, 2010, 6(3): e1000885.
[132]	Kinoshita T, Yadegari R, Harada J J, Goldberg R B, Fischer R L.Imprinting of the MEDEA polycomb gene in the Arabidopsis endosperm[J]. The Plant Cell, 1999, 11(10): 1945-1952.
[133]	Liu P, Qi M, Wang Y, Chang M, Liu C, Sun M, Yang W, Ren H.Arabidopsis RAN1 mediates seed development through its parental ratio by affecting the onset of endosperm cellularization[J]. Molecular Plant, 2014, 7(8): 1316-1328.
[134]	Wolff P, Jiang H, Wang G, Santos-González J, Köhler C.Paternally expressed imprinted genes establish postzygotic hybridization barriers in Arabidopsis thaliana[J]. eLife, 2015, 4: 1-14.
[135]	Wang G, Jiang H, Del Toro de León G, Martinez G, Köhler C. Sequestration of a transposon-derived sirna by a target mimic imprinted gene induces postzygotic reproductive isolation in Arabidopsis[J]. Developmental Cell, 2018, 46(6): 696-705.
[136]	Yuan J, Chen S, Jiao W, Wang L, Wang L, Ye W, Lu J, Hong D, You S, Cheng Z, Yang D L, Chen Z J.Both maternally and paternally imprinted genes regulate seed development in rice[J]. New Phytologist, 2017, 216(2): 373-387.
[137]	Folsom J J, Begcy K, Hao X, Wang D, Walia H.Rice fertilization-independent endosperm1 regulates seed size under heat stress by controlling early endosperm development[J]. Plant Physiology, 2014, 165(1): 238-248.

基因 Gene	基因登录号 Gene ID	编码蛋白 Encoded protein		突变体种子表型 Penotype of mutant	文献 Reference
OsFIE2	LOC_Os08g04270	PRC2复合体Esc家族	No	胚乳细胞化受阻	[20-21]
OsFIE1	LOC_Os08g04290	PRC2复合体Esc家族	MEG	休眠性降低	[20-21]
OsEMF2a	LOC_Os04g08034	PRC2复合体Su(z)12家族	MEG	胚乳细胞化受阻	[24-25]
OsEMF2b	LOC_Os09g13630	PRC2复合体Su(z)12家族	No	休眠性降低	[26-28]
OsMADS78	LOC_Os09g02830	I型MADS成员	Unknown	胚乳细胞化提前	[6, 39]
OsMADS79	LOC_Os01g74440	I型MADS成员	Unknown	胚乳细胞化提前	[6, 39]
OsMADS87	LOC_Os03g38610	I型MADS成员	MEG	胚乳细胞化提前	[6, 39]
Orysa;CycB1;1	LOC_Os01g59120	细胞周期蛋白	No	胚乳细胞化障碍导致无胚乳表型,胚增大	[42]
Oryza;KRP1	LOC_Os02g52480	细胞周期蛋白依赖性激酶抑制因子	No	过量表达时种子灌浆异常	[43-44]
ENL1	LOC_Os04g59624	SNF2解旋酶蛋白	No	无胚乳	[45]
OsGCD1	LOC_Os01g58750	GCD1同源基因	No	细胞化推迟	[46]
RPBF/OsDof3	LOC_Os02g15350	醇溶谷蛋白框结合蛋白, Dof转录因子	No	糊粉细胞层数变多	[55]
RISBZ1/bZIP58	LOC_Os07g08420	bZIP转录因子	No	单突无明显表型	[55]
OsROS1a	LOC_Os01g11900	DNA去甲基化酶	No	糊粉细胞层数显著增多	[57]
GFR1	LOC_Os10g36400	膜定位蛋白	No	显著降低了灌浆率	[58]
OsSUT1	LOC_Os03g07480	蔗糖转运蛋白	No	籽粒不能正常灌浆	[59]
OsSWEET4	LOC_Os02g19820	蔗糖转运蛋白	No	胚乳变小,结实率下降	[60]
OsSWEET11	LOC_Os08g42350	蔗糖转运蛋白	MEG	籽粒不充实	[61-62]
OsSWEET15	LOC_Os02g30910	蔗糖转运蛋白	No	籽粒不充实	[61-62]
GIF1	LOC_Os04g33740	细胞壁转化酶	No	灌浆不完全,粒重降低,粉质胚乳表型	[63]
GIF2	LOC_Os01g44220	AGP酶大亚基	No	种子淀粉合成障碍	[66]
SPK	LOC_Os10g39420	蔗糖合酶激酶	MEG	抑制淀粉在籽粒中的合成	[64]
OsMADS29	LOC_Os02g07430	II型MADS-box转录因子	No	细胞不能正常积累储藏物质,种子干瘪、败育	[68]
RSR1	LOC_Os05g03040	AP2家族转录因子	No	种子变大,直链淀粉含量升高,产量增加	[69]
OsNF-YB1	LOC_Os02g49410	NF-YB家族转录因子	No	结实率和千粒重均显著降低,种子空瘪萎缩	[72-75]
OsNF-YB9	LOC_Os06g17480	NF-YB家族转录因子	No	粒型变长,垩白增多	[72-76]
OsNF-YC12	LOC_Os10g11580	NF-YC家族转录因子	No	籽粒变小,千粒重下降	[73]
OsNF-YC10	LOC_Os01g24460	NF-YC家族转录因子	No	籽粒表现为细窄、变轻	[74]
OsbZIP76	LOC_Os09g34880	bZIP转录因子	MEG	细胞化进程提前,籽粒变小	[74]
OsNAC20	LOC_Os01g01470	NAC转录因子	No	和NAC26双突变体表型为粉质胚乳,粒宽、粒厚、千粒重降低	[79]
OsNAC26	LOC_Os01g29840	NAC转录因子	No	和NAC20双突变体表型为粉质胚乳,粒宽、粒厚、千粒重降低	[79]
ONAC127	LOC_Os11g31340	NAC转录因子	PEG	抑制种子中储藏物质的合成	[80]
ONAC129	LOC_Os11g31380	NAC转录因子	PEG	抑制种子中储藏物质的合成	[80]
OsFAD3	LOC_Os12g01370	脂肪酸去饱和酶	No	过表达导致亚麻酸（C18:3）含量显著提高	[87]
OsFAD2	LOC_Os02g48560	脂肪酸去饱和酶	No	籽粒中油脂的含量显著提高	[88]
OsYUC9	LOC_Os01g16714	黄素单加氧酶	No	种子粒重降低、垩白增加	[95]
OsYUC11	LOC_Os12g08780	黄素单加氧酶	PEG	种子灌浆缓慢	[95]
OsTAR1	LOC_Os05g07720	色氨酸转氨酶	No	种子较小、垩白增加,籽粒灌浆延迟	[95]
TGW6	LOC_Os06g41850	IAA-葡萄糖水解酶	No	显著提高千粒重	[96]
DG1	LOC_Os03g12790	ABA外排转运蛋白	No	灌浆延迟,垩白增加,高温下有空瘪籽粒	[102]
OsbHLH144	LOC_Os04g35010	bHLH转录因子	No	稻谷品质发生改变	[102]
OsCLF/SDG711	LOC_Os06g16390	PRC2复合体E(z)家族成员	No	抑制淀粉和储藏蛋白积累	[107-108]
SDG728/OsSET22	LOC_Os05g41172	组蛋白H3K9甲基转移酶	No	籽粒变小和粒重降低	[109]
OsLFR	LOC_Os07g41900	染色质重塑因子	No	胚乳早期游离核数目减少、不能细胞化	[110]
OsSRT1	LOC_Os04g20270	组蛋白去乙酰化酶基因	No	籽粒淀粉合成障碍	[111]
OsMET1b	LOC_Os07g08500	DNA甲基转移酶基因	PEG	籽粒皱缩败育	[112]
ROS1c/DNG701	LOC_Os05g37350	DNA去甲基化酶	No	15%的种子皱缩,不能正常灌浆	[112]
PEG1	LOC_Os01g08570	依赖酮戊二酸和铁的加氧酶	PEG	成熟籽粒空瘪	[136]

基因 Gene	基因登录号 Gene ID	编码蛋白 Encoded protein		突变体种子表型 Penotype of mutant	文献 Reference
OsFIE2	LOC_Os08g04270	PRC2复合体Esc家族	No	胚乳细胞化受阻	[20-21]
OsFIE1	LOC_Os08g04290	PRC2复合体Esc家族	MEG	休眠性降低	[20-21]
OsEMF2a	LOC_Os04g08034	PRC2复合体Su(z)12家族	MEG	胚乳细胞化受阻	[24-25]
OsEMF2b	LOC_Os09g13630	PRC2复合体Su(z)12家族	No	休眠性降低	[26-28]
OsMADS78	LOC_Os09g02830	I型MADS成员	Unknown	胚乳细胞化提前	[6, 39]
OsMADS79	LOC_Os01g74440	I型MADS成员	Unknown	胚乳细胞化提前	[6, 39]
OsMADS87	LOC_Os03g38610	I型MADS成员	MEG	胚乳细胞化提前	[6, 39]
Orysa;CycB1;1	LOC_Os01g59120	细胞周期蛋白	No	胚乳细胞化障碍导致无胚乳表型,胚增大	[42]
Oryza;KRP1	LOC_Os02g52480	细胞周期蛋白依赖性激酶抑制因子	No	过量表达时种子灌浆异常	[43-44]
ENL1	LOC_Os04g59624	SNF2解旋酶蛋白	No	无胚乳	[45]
OsGCD1	LOC_Os01g58750	GCD1同源基因	No	细胞化推迟	[46]
RPBF/OsDof3	LOC_Os02g15350	醇溶谷蛋白框结合蛋白, Dof转录因子	No	糊粉细胞层数变多	[55]
RISBZ1/bZIP58	LOC_Os07g08420	bZIP转录因子	No	单突无明显表型	[55]
OsROS1a	LOC_Os01g11900	DNA去甲基化酶	No	糊粉细胞层数显著增多	[57]
GFR1	LOC_Os10g36400	膜定位蛋白	No	显著降低了灌浆率	[58]
OsSUT1	LOC_Os03g07480	蔗糖转运蛋白	No	籽粒不能正常灌浆	[59]
OsSWEET4	LOC_Os02g19820	蔗糖转运蛋白	No	胚乳变小,结实率下降	[60]
OsSWEET11	LOC_Os08g42350	蔗糖转运蛋白	MEG	籽粒不充实	[61-62]
OsSWEET15	LOC_Os02g30910	蔗糖转运蛋白	No	籽粒不充实	[61-62]
GIF1	LOC_Os04g33740	细胞壁转化酶	No	灌浆不完全,粒重降低,粉质胚乳表型	[63]
GIF2	LOC_Os01g44220	AGP酶大亚基	No	种子淀粉合成障碍	[66]
SPK	LOC_Os10g39420	蔗糖合酶激酶	MEG	抑制淀粉在籽粒中的合成	[64]
OsMADS29	LOC_Os02g07430	II型MADS-box转录因子	No	细胞不能正常积累储藏物质,种子干瘪、败育	[68]
RSR1	LOC_Os05g03040	AP2家族转录因子	No	种子变大,直链淀粉含量升高,产量增加	[69]
OsNF-YB1	LOC_Os02g49410	NF-YB家族转录因子	No	结实率和千粒重均显著降低,种子空瘪萎缩	[72-75]
OsNF-YB9	LOC_Os06g17480	NF-YB家族转录因子	No	粒型变长,垩白增多	[72-76]
OsNF-YC12	LOC_Os10g11580	NF-YC家族转录因子	No	籽粒变小,千粒重下降	[73]
OsNF-YC10	LOC_Os01g24460	NF-YC家族转录因子	No	籽粒表现为细窄、变轻	[74]
OsbZIP76	LOC_Os09g34880	bZIP转录因子	MEG	细胞化进程提前,籽粒变小	[74]
OsNAC20	LOC_Os01g01470	NAC转录因子	No	和NAC26双突变体表型为粉质胚乳,粒宽、粒厚、千粒重降低	[79]
OsNAC26	LOC_Os01g29840	NAC转录因子	No	和NAC20双突变体表型为粉质胚乳,粒宽、粒厚、千粒重降低	[79]
ONAC127	LOC_Os11g31340	NAC转录因子	PEG	抑制种子中储藏物质的合成	[80]
ONAC129	LOC_Os11g31380	NAC转录因子	PEG	抑制种子中储藏物质的合成	[80]
OsFAD3	LOC_Os12g01370	脂肪酸去饱和酶	No	过表达导致亚麻酸（C18:3）含量显著提高	[87]
OsFAD2	LOC_Os02g48560	脂肪酸去饱和酶	No	籽粒中油脂的含量显著提高	[88]
OsYUC9	LOC_Os01g16714	黄素单加氧酶	No	种子粒重降低、垩白增加	[95]
OsYUC11	LOC_Os12g08780	黄素单加氧酶	PEG	种子灌浆缓慢	[95]
OsTAR1	LOC_Os05g07720	色氨酸转氨酶	No	种子较小、垩白增加,籽粒灌浆延迟	[95]
TGW6	LOC_Os06g41850	IAA-葡萄糖水解酶	No	显著提高千粒重	[96]
DG1	LOC_Os03g12790	ABA外排转运蛋白	No	灌浆延迟,垩白增加,高温下有空瘪籽粒	[102]
OsbHLH144	LOC_Os04g35010	bHLH转录因子	No	稻谷品质发生改变	[102]
OsCLF/SDG711	LOC_Os06g16390	PRC2复合体E(z)家族成员	No	抑制淀粉和储藏蛋白积累	[107-108]
SDG728/OsSET22	LOC_Os05g41172	组蛋白H3K9甲基转移酶	No	籽粒变小和粒重降低	[109]
OsLFR	LOC_Os07g41900	染色质重塑因子	No	胚乳早期游离核数目减少、不能细胞化	[110]
OsSRT1	LOC_Os04g20270	组蛋白去乙酰化酶基因	No	籽粒淀粉合成障碍	[111]
OsMET1b	LOC_Os07g08500	DNA甲基转移酶基因	PEG	籽粒皱缩败育	[112]
ROS1c/DNG701	LOC_Os05g37350	DNA去甲基化酶	No	15%的种子皱缩,不能正常灌浆	[112]
PEG1	LOC_Os01g08570	依赖酮戊二酸和铁的加氧酶	PEG	成熟籽粒空瘪	[136]