水稻株型的分子机理研究进展

doi:10.16819/j.1001-7216.2023.221102

摘要/Abstract

摘要：

株型是水稻产量的重要决定因素，创制理想株型品种是提高水稻产量的重要途径。20世纪50年代的矮化育种和70年代的杂交水稻培育是水稻增产的两次革命，但近几年水稻产量增长放缓，通过理想株型与杂种优势的结合进而实现超高产将成为第三次育种革命的关键。本文简要回顾了水稻株型分子调控机理的最新研究概况，重点关注了叶形、穗型和粒型等方面所取得的进展，并展望了水稻理想株型未来的研究趋势，为通过分子设计育种创制水稻理想株型，进一步提高水稻产量提供参考。

关键词: 水稻, 理想株型, 叶形, 穗型, 粒型, 分子设计

Abstract:

The plant type of rice is one of the major factors determining rice yield. The ideotype of high-yielding rice cultivar is an important way to increase rice yield potential in rice breeding. Dwarf breeding in the 1950s and hybrid rice breeding in the 1970s are two revolutions in rice yield. However, the growth rate of rice yield has slowed down significantly in recent years. The third yield breakthrough depends on the super-high-yield breeding that combines ideotype with heterosis. We briefly reviewed the advances in the regulation mechanism of rice plant architecture, focusing on the progress achieved in leaf, panicle, grain and other aspects, and looked forward to the future research trends rice ideotype. It will lay a solid foundation for improving rice plant type and further increasing rice yield through molecular breeding.

Key words: rice, ideotype, leaf shape, panicle type, grain type, molecular breeding

兰金松, 庄慧. 水稻株型的分子机理研究进展[J]. 中国水稻科学, 2023, 37(5): 449-458.

LAN Jinsong, ZHUANG Hui. Advances in the Molecular Mechanism of Rice Plant Type[J]. Chinese Journal OF Rice Science, 2023, 37(5): 449-458.

图/表 1

参考文献 78

[1]	张启发. 绿色超级稻培育的设想[J]. 分子植物育种, 2005, 3(5): 601-602.
	Zhang Q F. Strategies for developing green super rice[J]. Molecular Plant Breeding, 2005, 3(5): 601-602. (in Chinese)
[2]	Boysen-Jensen P. Die Stoffproduktion der Pflanzen[J]. Protoplasma, 1933, 18(1): 311.
[3]	Heath O V S, Gregory F G. The constancy of the mean net assimilation rate and its ecological importance[J]. Annals of Botany, 1938, 2(4): 811-818.
[4]	Donald C M. The breeding of crop ideotypes[J]. Euphytica, 1968, 17(3): 385-403.
[5]	马梦影, 巩文靓, 康雪蒙, 段海燕. 水稻理想株型改良的研究进展[J]. 中国农学通报, 2020, 36(29): 1-6.
	Ma M Y, Gong W L, Kang X M, Duan H Y. The improvement of ideal plant type of rice: A review[J]. China Agricultural Science Bulletin, 2020, 36(29): 1-6. (in Chinese with English abstract)
[6]	黄耀祥. 水稻超高产育种研究[J]. 作物杂志, 1990(4): 1-2.
	Huang Y X. Study on super high yield rice breeding[J]. Crops, 1990(4): 1-2. (in Chinese)
[7]	杨守仁, 张龙步, 徐正进, 陈温福, 王进民, 董克. 水稻理想株形育种的基础研究及其与国内外同类研究的比较[J]. 沈阳农业大学学报, 1991(S1): 1-5.
	Yang S R, Zhang L B, Xu Z J, Chen W F, Wang J M, Dong K. The research results of ideal morphological breeding in rice and their comparisons with the same studies at home and abroad[J]. Journal of Shenyang Agricultural University, 1991(S1): 1-5. (in English with Chinese abstract)
[8]	杨守仁, 张龙步, 陈温福, 徐正进, 王进民. 水稻超高产育种的理论和方法[J]. 中国水稻科学, 1996, 10(2): 115-120.
	Yang S R, Zhang L B, Chen W F, Xu Z J, Wang J M. Theories and methods of rice breeding for maximum yield[J]. Chinese Journal of Rice Science, 1996, 10(2): 115-120. (in Chinese with English abstract)
[9]	周开达, 汪旭东, 李仕贵, 李平, 黎汉云, 黄国寿, 刘太清, 沈茂松. 亚种间重穗型杂交稻研究[J]. 中国农业科学, 1997(5): 92-94.
	Zhou K D, Wang X D, Li S G, Li P, Li H Y, Huang G S, Liu T Q, Shen M S. The study on heavy panicle type of inter-subspecific hybrid rice (Oryza sativa L.)[J]. Scientia Agricultura Sinica, 1997(5): 92-94. (in Chinese with English abstract)
[10]	汪开治. 国际水稻所将推出理想株型水稻新品种[J]. 中国农技推广, 1996(4): 26.
	Wang K Z. International Rice Research Institute will introduce a new ideal strain type of rice varieties[J]. China Agro-Technology Extension, 1996(4): 26. (in Chinese)
[11]	袁隆平. 杂交水稻超高产育种[J]. 杂交水稻, 1997, 12(6): 1-6.
	Yuan L P. Hybrid rice breeding for super high yield[J]. Hybrid Rice, 1997, 12(6): 1-6. (in Chinese)
[12]	Li S, Tian Y H, Wu K, Ye Y F, Yu J P, Zhang J Q, Liu Q, Hu M Y, Li H, Tong Y P, Nicholas Harberd P, Fu X D. Modulating plant growth-metabolism coordination for sustainable agriculture[J]. Nature, 2018, 560(7720): 595-600.
[13]	胡茂龙. 水稻光合功能相关性状QTL分析及转绿型白叶突变体基因的图位克隆[D]. 南京: 南京农业大学, 2006.
	Hu M L. QTL analysis for traits associated with photosynthetic functions and map-based cloning of virescent white leaf gene in rice (Oryza sativa L.)[D]. Nanjing: Nanjing Agricultural University, 2006. (in Chinese with English abstract)
[14]	陈达刚, 周新桥, 李丽君, 刘传光, 陈友订. 水稻叶厚性状的研究进展[J]. 农学学报, 2015, 5(11): 22-25.
	Chen D G, Zhou X Q, Li L J, Liu C G, Chen Y D. Research progress on rice (Oryza sativa L.) leaf thickness[J]. Journal of Agriculture, 2015, 5(11): 22-25. (in Chinese with English abstract)
[15]	高艳红, 吕川根, 王茂青, 王澎, 闫晓燕, 谢坤, 万建民. 水稻卷叶性状QTL的初步定位[J]. 江苏农业学报, 2007, 23(1): 5-10.
	Gao Y H, Lü C G, Wang M Q, Wang P, Yan X Y, Xie K, Wan J M. QTL mapping for rolled leaf gene in rice[J]. Jiangsu Journal of Agricultural Science, 2007, 23(1): 5-10. (in Chinese with English abstract)
[16]	Yoshida S. Fundamentals of Rice Crop Science[J]. Los Banos, the Philippines: International Rice Research Institute, 1981: 1-61.
[17]	范玉斌, 梁婉琪. 水稻叶极性发育分子机制研究进展[J]. 上海交通大学学报: 农业科学版, 2014, 32(1): 16-22.
	Fan Y B, Liang W Q. Research progress on the mechanism of leaf polarity establishment in rice[J]. Journal of Shanghai Jiaotong University: Agricultural Science, 2014, 32(1): 16-22. (in Chinese with English abstract)
[18]	Xiang J J, Zhang G H, Qian Q, Xue H W. SEMI-ROLLED LEAF1 encodes a putative GPI-anchored protein and modulates rice leaf rolling by regulating the formation of bulliform cells[J]. Plant Physiology, 2012, 159(4): 1488-1500.
[19]	Hibara K I, Obara M, Hayashida E, Abe M, Ishimaru T, Satoh H, Itoh J I, Nagato Y. The ADAXIALIZED LEAF1 gene functions in leaf and embryonic pattern formation in rice[J]. Developmental Biology, 2009, 334(2): 345-354.
[20]	Zhang T, You J, Zhang Y, Yao W Y, Chen W B, Duan Q N, Xiao W W, Ye L, Zhou Y, Sang X C, Ling Y H, He G H, Li Y F. LF1 regulates the lateral organs polarity development in rice[J]. The New Phytologist, 2021, 231(3): 1265-1277.
[21]	You J, Xiao W W, Zhou Y, Shen W Q, Ye L, Yu P, Yu G L, Duan Q N, Zhang X F, He Z F, Xiang Y, Sang X C, Li Y F, Zhao F M, Ling Y H, He G H, Zhang T. The APC/CTAD1-WIDE LEAF 1-NARROW LEAF 1 pathway controls leaf width in rice[J]. The Plant Cell, 2022, 34(11): 4313-4328.
[22]	姚栋萍. 水稻培矮64S直立叶基因的初步定位[D]. 长沙: 湖南农业大学, 2016.
	Yao D P. Preliminary mapping of the rice erect leaf gene from Peiai 64s[D]. Changsha: Hunan Agricultural University, 2016. (in Chinese with English abstract)
[23]	Zhao S Q, Hu J, Guo L B, Qian Q, Xue H W. Rice leaf inclination2, a VIN3-like protein, regulates leaf angle through modulating cell division of the collar[J]. Cell Research, 2010, 20(8): 935-947.
[24]	Tanaka A, Nakagawa H, Tomita C, Shimatani Z, Ohtake M, Nomura T, Jiang C J, Dubouzet J G, Kikuchi S, Sekimoto H, Yokota T, Asami T, Kamakura T, Mori M. BRASSINOSTEROID UPREGULATED1, encoding a Helix-Loop-Helix protein, is a novel gene involved in brassinosteroid signaling and controls bending of the lamina joint in rice[J]. Plant Physiology, 2009, 151(2): 669-680.
[25]	Ning J, Zhang B C, Wang N L, Zhou Y H, Xiong L Z. Increased Leaf Angle1, a Raf-Like MAPKKK that interacts with a nuclear protein family, regulates mechanical tissue formation in the lamina joint of rice[J]. The Plant Cell, 2011, 23(12): 4334-4347.
[26]	Wu X R, Tang D, Li M, Wang K J, Cheng Z K. Loose Plant Architecture1, an INDETERMINATE DOMAIN protein involved in shoot gravitropism, regulates plant architecture in rice[J]. Plant Physiology, 2013, 161(1): 317-329.
[27]	Yasuno N, Takamure I, Kidou S, Tokuji Y, Ureshi A, Funabiki A, Ashikaga K, Yamanouchi U, Yano M, Kato K. Rice shoot branching requires an ATP-binding cassette subfamily G protein[J]. New Phytologist, 2009, 182(1): 91-101.
[28]	Nakagawa M, Shimamoto K, Kyozuka J. Overexpression of RCN1 and RCN2, rice TERMINAL FLOWER 1/CENTRORADIALIS homologs, confers delay of phase transition and altered panicle morphology in rice[J]. The Plant Journal, 2002, 29(6): 743-750.
[29]	Terao T, Nagata K, Morino K, Hirose T. A gene controlling the number of primary rachis branches also controls the vascular bundle formation and hence is responsible to increase the harvest index and grain yield in rice[J]. Theoretical and Applied Genetics, 2010, 120(5): 875-893.
[30]	Ikeda-Kawakatsu K, Maekawa M, Izawa T, Itoh JI, Nagato Y. ABERRANT PANICLE ORGANIZATION 2/RFL, the rice ortholog of Arabidopsis LEAFY, suppresses the transition from inflorescence meristem to floral meristem through interaction with APO1[J]. The Plant Journal, 2012, 69(1): 168-180.
[31]	Zeng X Q, Zhuang H, Cheng Q L, Tang J, Yang F Y, Huang M J, Wang Z Y, Li Z C, Zhu H H, Chen R, He G H, Li Y F. SB1 encoding Ring-Like Zinc-Finger protein regulates branch development as a transcription repressor[J]. Rice Science, 2021, 28(3): 243-256.
[32]	Zhang D B, Yuan Z. Molecular control of grass inflorescence development[J]. Annual Review of Plant Biology, 2014, 65(1): 553-578.
[33]	Pautler M, Tanaka W, Hirano H Y, Jackson D. Grass meristems: I. Shoot apical meristem maintenance, axillary meristem determinacy and the floral transition[J]. Plant and Cell Physiology, 2013, 54(3): 302-312.
[34]	Bradley D, Ratcliffe O, Vincent C, Carpenter R, Coen E. Inflorescence commitment and architecture in Arabidopsis[J]. Science, 1997, 275(5296): 80-83.
[35]	Ratcliffe O J, Bradley D J, Coen E S. Separation of shoot and floral identity in Arabidopsis[J]. Development, 1999, 126(6): 1109-1120.
[36]	Mimida N, Goto K, Kobayashi Y, Araki T, Ahn J H, Weigel D, Murata M, Motoyoshi F, Sakamoto W. Functional divergence of the TFL1-like gene family in Arabidopsis revealed by characterization of a novel homologue[J]. Genes to Cells, 2001, 6(4): 327-336.
[37]	Rao N N, Prasad K, Kumar P R, Vijayraghavan U. Distinct regulatory role for RFL, the rice LFY homolog, in determining flowering time and plant architecture[J]. Proceedings of the National Academy of Sciences of the United States of America, 2008, 105(9): 3646-3651.
[38]	Yoshida A, Sasao M, Yasuno N, Takagi K, Daimon Y, Chen R H, Yamazaki R, Tokunaga H, Kitaguchi Y, Sato Y, Nagamura Y, Ushijima T, Kumamaru T, Iida S, Maekawa M, Kyozuka J. TAWAWA1, a regulator of rice inflorescence architecture, functions through the suppression of meristem phase transition[J]. Proceedings of the National Academy of Sciences of the United States of America, 2013, 110(2): 767-772.
[39]	Huang Y, Bai X F, Luo M F, Xing Y Z. Short Panicle 3 controls panicle architecture by upregulating APO2/RFL and increasing cytokinin content in rice[J]. Journal of Integrative Plant Biology, 2019, 61(9): 987-999.
[40]	Xue W Y, Xing Y Z, Weng X Y, Zhao Y, Tang W J, Wang L, Zhou H J, Yu S B, Xu C G, Li X H, Zhang Q F. Natural variation in Ghd7 is an important regulator of heading date and yield potential in rice[J]. Nature Genetics, 2008, 40(6): 761-767.
[41]	Wei X J, Xu J F, Guo H N, Jiang L, Chen S H, Yu C Y, Zhou Z L, Hu P S, Zhai H Q, Wan J M. DTH8 suppresses flowering in rice, influencing plant height and yield potential simultaneously[J]. Plant Physiology, 2010, 153(4): 1747-1758.
[42]	Yan W H, Wang P, Chen H X, Zhou H J, Li Q P, Wang C R, Ding Z H, Zhang Y S, Yu S B, Xing Y Z, Zhang Q F. A major QTL, Ghd8, plays pleiotropic roles in regulating grain productivity, plant height, and heading date in rice[J]. Molecular Plant, 2011, 4(2): 319-330.
[43]	Dai X D, Ding Y N, Tan L B, Fu Y C, Liu F X, Zhu Z F, Sun X Y, Sun X W, Gu P, Cai H W, Sun C Q. LHD1, an allele of DTH8/Ghd8, controls late heading date in common wild rice (Oryza rufipogon)[J]. Journal of Integrative Plant Biology, 2012, 54(10): 790-799.
[44]	Sheng P K, Wu F Q, Tan J J, Zhang H, Ma W W, Chen L P, Wang J C, Wang J, Zhu S S, Guo X P, Wang J L, Zhang X, Cheng Z J, Bao Y Q, Wu C Y, Liu X M, Wan J M. A CONSTANS-like transcriptional activator, OsCOL13, functions as a negative regulator of flowering downstream of OsphyB and upstream of Ehd1 in rice[J]. Plant Molecular Biology, 2016, 92(1-2): 209-222.
[45]	Li S B, Qian Q, Fu Z M, Zeng D L, Meng X B, Kyozuka J, Maekawa M, Zhu X D, Zhang J, Li J Y, Wang Y H. Short panicle1 encodes a putative PTR family transporter and determines rice panicle size[J]. The Plant Journal, 2009, 58(4): 592-605.
[46]	Jiang G H, Xiang Y H, Zhao J Y, Yin D D, Zhao X F, Zhu L H, Zhai W X. Regulation of inflorescence branch development in rice through a novel pathway involving the pentatricopeptide repeat protein sped1-D[J]. Genetics, 2014, 197(4): 1395-1407.
[47]	Qiao Y L, Piao R H, Shi J X, Lee S I, Jiang W Z, Kim B K, Lee J, Han L Z, Ma W B, Koh H J. Fine mapping and candidate gene analysis of dense and erect panicle 3, DEP3, which confers high grain yield in rice (Oryza sativa L.)[J]. Theoretical and Applied Genetics, 2011, 122(7): 1439-1449.
[48]	Piao R H, Jiang W Z, Ham T H, Choi M S, Qiao Y, Chu S H, Park J H, Woo M O, Jin Z, An G, Lee J, Koh H J. Map-based cloning of the ERECT PANICLE 3 gene in rice[J]. Theoretical and Applied Genetics, 2009, 119(8): 1497-1506.
[49]	Li M, Tang D, Wang K J, Wu X R, Lu L L, Yu H X, Gu M D, Yan C J, Cheng Z K. Mutations in the F-box gene LARGER PANICLE improve the panicle architecture and enhance the grain yield in rice[J]. Plant Biotechnology Journal, 2011, 9(9): 1002-1013.
[50]	Gao X C, Liang W Q, Yin C S, Ji S M, Wang H M, Su X, Guo C, Kong H Z, Xue H W, Zhang D B. The SEPALLATA-like gene OsMADS34 is required for rice inflorescence and spikelet development[J]. Plant Physiology, 2010, 153(2): 728-740.
[51]	Kobayashi K, Maekawa M, Miyao A, Hirochika H, Kyozuka J. PANICLE PHYTOMER2 (PAP2), encoding a SEPALLATA subfamily MADS-box protein, positively controls spikelet meristem identity in rice[J]. Plant and Cell Physiology, 2010, 51(1): 47-57.
[52]	Zhuang H, Wang H L, Zhang T, Zeng X Q, Chen H, Wang Z W, Zhang J, Zheng H, Tang J, Ling Y H, Yang Z L, He G H, Li Y F. NONSTOP GLUMES1 encodes a C2H2 zinc finger protein that regulates spikelet development in rice[J]. The Plant Cell, 2020, 32(2): 392-413.
[53]	Zhang T, Li Y F, Ma L, Sang X C, Ling Y H, Wang Y T, Yu P, Zhuang H, Huang J Y, Wang N, Zhao F M, Zhang C W, Yang Z L, Fang L K, He G H. LATERAL FLORET 1 induced the three-florets spikelet in rice[J]. Proceedings of the National Academy of Sciences of the United States of America, 2017, 114(37): 9984-9989.
[54]	Ren D Y, Li Y F, Zhao F M, Sang X C, Shi J Q, Wang N, Guo S, Ling Y H, Zhang C W, Yang Z L, He G H. MULTI-FLORET SPIKELET1, which encodes an AP2/ERF protein, determines spikelet meristem fate and sterile lemma identity in rice[J]. Plant Physiology, 2013, 162(2): 872-884.
[55]	Li Y F, Zeng X Q, Li Y, Wang L, Zhuang H, Wang Y, Tang J, Wang H L, Xiong M, Yang F Y, Yuan X Z, He G H. MULTI-FLORET SPIKELET 2, a MYB transcription factor, determines spikelet meristem fate and floral organ identity in rice[J]. Plant Physiology, 2020, 184(2): 988-1003.
[56]	Wang Y, Zeng X Q, Lu L, Cheng Q L, Yang F Y, Huang M J, Xiong M, Li Y F. MULTI-FLORET SPIKELET 4 (MFS4) regulates spikelet development and grain size in rice[J]. Rice Science, 2021, 28(4): 344-357.
[57]	Zheng H, Zhang J, Zhuang H, Zeng X Q, Tang J, Wang H L, Chen H, Li Y, Ling Y H, He G H, Li Y F. mfs3) in rice (Oryza sativa L.)[J]. Journal of Integrative Agriculture, 2019, 18(12): 2673-2681.
[58]	Ren D Y, Li Y F, He G H, Qian Q. Multifloret spikelet improves rice yield[J]. New Phytologist, 2020, 225(6): 2301-2306.
[59]	徐建龙, 薛庆中, 罗利军, 黎志康. 水稻粒重及其相关性状的遗传解析[J]. 中国水稻科学, 2002, 16(1): 6-10.
	Xu J L, Xue Q Z, Luo L J, Li Z K. Genetic dissection of grain weight and its related traits in rice (Oryza sativa L.)[J]. Chinese Journal of Rice Science, 2002, 16(1): 6-10. (in Chinese with English abstract)
[60]	Li N, Xu R, Duan P G, Li Y H. Control of grain size in rice[J]. Plant Reproduction, 2018, 31(3): 237-251.
[61]	Li N, Xu R, Li Y H. Molecular networks of seed size control in plants[J]. Annual Review of Plant Biology, 2019, 70(1): 435-463.
[62]	Fan Y W, Li Y B. Molecular, cellular and Yin-Yang regulation of grain size and number in rice[J]. Molecular Breeding, 2019, 39(12): 1-25.
[63]	Huang X Z, Qian Q, Liu Z B, Sun H Y, He S Y, Luo D, Xia G M, Chu C C, Li J Y, Fu X D. Natural variation at the DEP1 locus enhances grain yield in rice[J]. Nature Genetics, 2009, 41(4): 494-497.
[64]	Mao H L, Sun S Y, Yao J L, Wang C R, Yu S B, Xu C G, Li X H, Zhang Q F. Linking differential domain functions of the GS3 protein to natural variation of grain size in rice[J]. Proceedings of the National Academy of Sciences of the United States of America, 2010, 107(45): 19579-19584.
[65]	Takano-Kai N, Doi K, Yoshimura A. GS3 participates in stigma exsertion as well as seed length in rice[J]. Breeding Science, 2011, 61(3): 244-250.
[66]	Sun S Y, Wang L, Mao H L, Shao L, Li X H, Xiao J H, Ouyang Y D, Zhang Q F. A G-protein pathway determines grain size in rice[J]. Nature Communications, 2018, 9(1): 851.
[67]	Huang H X, Ye Y F, Song W Z, Li Q, Han R X, Wu C C, Wang S X, Yu J P, Liu X Y, Fu X D, Liu Q, Wu K. Modulating the C-terminus of DEP1 synergistically enhances grain quality and yield in rice[J]. Journal of Genetics and Genomics, 2022, 49(5): 506-509.
[68]	Yamamuro C, Ihara Y, Wu X, Noguchi T, Fujioka S, Takatsuto S, Ashikari M, Kitano H, Matsuok M. Loss of function of a rice brassinosteroid insensitive1 homolog prevents internode elongation and bending of the lamina joint[J]. The Plant Cell, 2000, 12(9): 1591-1606.
[69]	Morinaka Y, Sakamoto T, Inukai Y, Agetsuma M, Kitano H, Ashikari M, Matsuoka M. Morphological alteration caused by brassinosteroid insensitivity increases the biomass and grain production of rice[J]. Plant Physiology, 2006, 141(3): 924-931.
[70]	Ito Y, Takaya K, Kurata N. Expression of SERK family receptor-like protein kinase genes in rice[J]. Biochimica et Biophysica Acta, 2005, 1730(3): 253-258.
[71]	Li D, Wang L, Wang M, Xu Y Y, Luo W, Liu Y J, Xu Z H, Li J. Engineering OsBAK1 gene as a molecular tool to improve rice architecture for high yield[J]. Plant Biotechnology Journal, 2009, 7(8): 791-806.
[72]	Tong H N, Liu L C, Jin Y, Du L, Yin Y H, Qian Q, Zhu L H, Chu C C. DWARF AND LOW-TILLERING acts as a direct downstream target of a GSK3/SHAGGY-like kinase to mediate brassinosteroid responses in rice[J]. The Plant Cell, 2012, 24(6): 2562-2577.
[73]	Tong H N, Jin Y, Liu W B, Li F, Fang J, Yin Y H, Qian Q, Zhu L H, Chu C C. DWARF AND LOW- TILLERING, a new member of the GRAS family, plays positive roles in brassinosteroid signaling in rice[J]. The Plant Journal, 2009, 58(5): 803-816.
[74]	Sun L J, Li X J, Fu Y C, Zhu Z F, Tan L B, Liu F X, Sun X Y, Sun X W, Sun C Q. GS6, a member of the GRAS gene family, negatively regulates grain size in rice[J]. Journal of Integrative Plant Biology, 2013, 55(10): 938-949.
[75]	Song X J, Huang W, Shi M, Zhu M Z, Lin H X. A QTL for rice grain width and weight encodes a previously unknown RING-type E3 ubiquitin ligase[J]. Nature Genetics, 2007, 39(5): 623-630.
[76]	Hao J Q, Wang D K, Wu Y B, Huang K, Duan P G, Li N, Xu R, Zeng D L, Dong G J, Zhang B L, Zhang L M, Inzé D, Qian Q, Li Y H. The GW2-WG1-OsbZIP47 pathway controls grain size and weight in rice[J]. Molecular Plant, 2021, 14(8): 1266-1280.
[77]	Hu X M, Qian Q, Xu T, Zhang Y E, Dong G J, Gao T, Xie Q, Xue Y B. The U-box E3 ubiquitin ligase TUD1 functions with a heterotrimeric G α subunit to regulate Brassinosteroid-mediated growth in rice[J]. PLOS Genetics, 2013, 9(3): e1003391.
[78]	Jiao Y Q, Wang Y H, Xue D W, Wang J, Yan M X, Liu G F, Dong G J, Zeng D L, Lu Z F, Zhu X D, Qian Q, Li J Y. Regulation of OsSPL14 by OsmiR156 defines ideal plant architecture in rice[J]. Nature Genetics, 2010, 42(6): 541-544.

基因 Gene	基因登录号 Gene ID	编码蛋白 Encoded protein	基因效应 Gene effect	文献 Reference
叶形 Leaf shape
SRL1	LOC_Os07g01240	糖基磷脂酰肌醇锚定蛋白	抑制泡状细胞形成与近轴面形成，特化叶片远轴面	[18]
ADL1	LOC_Os02g47970	半胱氨酸蛋白酶	促进表皮发育，维持叶片近轴和远轴	[19]
LF1	LOC_Os03g01890	HD-ZIP Ⅲ转录激活因子	突变体极性发育异常，叶片正面卷曲，外稃凹陷；影响生长素含量，调控水稻侧生器官极性发育	[20]
WL1		C2H2锌指转录因子	与水稻转录共抑制因子TPR结合下调窄叶基因NAL的表达，调控叶片的宽度	[21]
LC2	LOC_Os02g05840	类vernalization insensitive 3 蛋白	集中在叶片和叶鞘处表达，抑制维管束两侧泡状细胞增多；调控叶夹角大小	[23]
BU1	LOC_Os06g12210	HLH蛋白	正向调节水稻中的油菜素内酯含量；过表达增加叶片结合处弯曲角度，抑制表达的植株叶片恢复直立	[24]
ILA1	LOC_Os06g50920	丝裂原活化蛋白激酶	突变体中叶枕机械组织异常，影响叶倾角的形成	[25]
LPA1	LOC_Os03g13400	植物特有的结构域不确定 (ID) 转录抑制因子	突变体分蘖角和叶夹角增大	[26]
穗型 Panicle type
RCN1 RCN2	LOC_Os11g05470 LOC_Os02g32950	TFL1/CEN同源基因	过表达延长枝梗分生组织向花分生组织的转变时间，增加枝梗数和每穗粒数，决定枝梗(花序)形态	[28]
APO2/RFL	LOC_Os04g51000	拟南芥LFY的同源基因	维持枝梗 (花序) 分生组织命运；与APO1 相互作用共同延迟枝梗分生组织向小穗分生组织的转化；调控水稻枝梗数与每穗粒数	[30,37]
TAW1	LOC_Os10g33780	花序形态调控因子	调控枝梗分生组织，影响枝梗数与穗粒数	[38]
SP3		DOF转录激活因子	通过激活APO2/RFL基因调控枝梗分生组织；调控枝梗数与每穗粒数	[39]
SB1		SHI转录因子	等位突变体内枝梗分生组织分化延迟，枝梗数与小穗数增加；负向调节维持枝梗分生组织活性的基因	[31]
Ghd7	LOC_Os07g15770	CCT结构蛋白	影响抽穗期、株高与每穗粒数	[40]
Ghd8	LOC_Os08g07740	核因子Y的B亚基	上调控制分蘖和侧枝发生基因MOC1的表达，促进水稻分蘖和枝梗形成	[41⇓-43]
OsCOL13	LOC_Os07g47140	类CONSTANS转录激活因子	延迟抽穗期，增加穗长，提高每穗粒数	[44]
SP1	LOC_Os11g12740	多肽转运家族蛋白	影响水稻穗轴的伸长和穗的大小，决定每穗粒数	[45]
LF1	LOC_Os03g01890	HD-ZIP III转录激活因子	激活分生组织维持基因OSH1异位表达，起始侧生花分生组织形成；理论上为培育“三花小穗”水稻品种实现水稻增产提供了证据	[53]
MFS1~4	LOC_Os05g41760 LOC_Os04g47890	AP2/ERF的转录因子	正向调控水稻小穗分生组织向花分生组织的转变	[54⇓⇓-57]
粒型 Grain type
GS3		由232个氨基酸组成的跨膜蛋白	在翻译水平影响粒型；与DEP1或GGC2竞争性结合Gβ，缩短粒长	[65-66]
DEP1	LOC_Os09g26999	G蛋白γ亚基	控制水稻产量性状；功能获得性突变体植株表现出短穗、直立穗、密穗等产量性状；调控粒长和粒重	[67]
GGC2		异三聚体G蛋白γ亚基	调控粒长，与DEP1对粒长的正调控有加性作用	[66]
OsBRI1	LOC_Os01g52050	BR信号受体	在调控水稻粒型中起多种作用；影响细胞伸长和增殖，调控器官发育	[68-69]
OsBAK1	LOC_Os08g07760	SERK家族类受体蛋白激酶；BR信号受体激酶	影响株高、叶夹角、粒型和抗病性	[70-71]
GSK2	LOC_Os05g11730	GSK3/SHAGGY激酶	负调控BR应答基因的表达，参与水稻株型相关性状的调控	[72]
DLT	LOC_Os06g03710	由617个氨基酸组成的植物特有的GRAS家族蛋白	作用于BR应答通路下游，正向介导BR应答；影响细胞增殖及细胞数目，影响颖壳发育，负调控籽粒大小	[72,74]
GW2	LOC_Os02g14720	E3泛素连接酶	负调控细胞分裂，影响水稻的粒宽和粒重；促进泛素转移，特异识别应降解的底物，影响外稃细胞分裂，调控粒重	[75]
WG1/OsGRX8	LOC_Os02g30850	CC型谷氧还蛋白OsGRX8	调控籽粒大小，是粒宽方向细胞增殖所必需；与GW2互作并将其泛素化，从而调控WG1的稳定性	[76]
TUD1	LOC_Os03g13010	U-box家族的E3泛素连接酶	与异三聚体G蛋白α亚基D1互作调控油菜素内酯介导水稻生长，进而影响株高、粒型等相关性状	[77]

基因 Gene	基因登录号 Gene ID	编码蛋白 Encoded protein	基因效应 Gene effect	文献 Reference
叶形 Leaf shape
SRL1	LOC_Os07g01240	糖基磷脂酰肌醇锚定蛋白	抑制泡状细胞形成与近轴面形成，特化叶片远轴面	[18]
ADL1	LOC_Os02g47970	半胱氨酸蛋白酶	促进表皮发育，维持叶片近轴和远轴	[19]
LF1	LOC_Os03g01890	HD-ZIP Ⅲ转录激活因子	突变体极性发育异常，叶片正面卷曲，外稃凹陷；影响生长素含量，调控水稻侧生器官极性发育	[20]
WL1		C2H2锌指转录因子	与水稻转录共抑制因子TPR结合下调窄叶基因NAL的表达，调控叶片的宽度	[21]
LC2	LOC_Os02g05840	类vernalization insensitive 3 蛋白	集中在叶片和叶鞘处表达，抑制维管束两侧泡状细胞增多；调控叶夹角大小	[23]
BU1	LOC_Os06g12210	HLH蛋白	正向调节水稻中的油菜素内酯含量；过表达增加叶片结合处弯曲角度，抑制表达的植株叶片恢复直立	[24]
ILA1	LOC_Os06g50920	丝裂原活化蛋白激酶	突变体中叶枕机械组织异常，影响叶倾角的形成	[25]
LPA1	LOC_Os03g13400	植物特有的结构域不确定 (ID) 转录抑制因子	突变体分蘖角和叶夹角增大	[26]
穗型 Panicle type
RCN1 RCN2	LOC_Os11g05470 LOC_Os02g32950	TFL1/CEN同源基因	过表达延长枝梗分生组织向花分生组织的转变时间，增加枝梗数和每穗粒数，决定枝梗(花序)形态	[28]
APO2/RFL	LOC_Os04g51000	拟南芥LFY的同源基因	维持枝梗 (花序) 分生组织命运；与APO1 相互作用共同延迟枝梗分生组织向小穗分生组织的转化；调控水稻枝梗数与每穗粒数	[30,37]
TAW1	LOC_Os10g33780	花序形态调控因子	调控枝梗分生组织，影响枝梗数与穗粒数	[38]
SP3		DOF转录激活因子	通过激活APO2/RFL基因调控枝梗分生组织；调控枝梗数与每穗粒数	[39]
SB1		SHI转录因子	等位突变体内枝梗分生组织分化延迟，枝梗数与小穗数增加；负向调节维持枝梗分生组织活性的基因	[31]
Ghd7	LOC_Os07g15770	CCT结构蛋白	影响抽穗期、株高与每穗粒数	[40]
Ghd8	LOC_Os08g07740	核因子Y的B亚基	上调控制分蘖和侧枝发生基因MOC1的表达，促进水稻分蘖和枝梗形成	[41⇓-43]
OsCOL13	LOC_Os07g47140	类CONSTANS转录激活因子	延迟抽穗期，增加穗长，提高每穗粒数	[44]
SP1	LOC_Os11g12740	多肽转运家族蛋白	影响水稻穗轴的伸长和穗的大小，决定每穗粒数	[45]
LF1	LOC_Os03g01890	HD-ZIP III转录激活因子	激活分生组织维持基因OSH1异位表达，起始侧生花分生组织形成；理论上为培育“三花小穗”水稻品种实现水稻增产提供了证据	[53]
MFS1~4	LOC_Os05g41760 LOC_Os04g47890	AP2/ERF的转录因子	正向调控水稻小穗分生组织向花分生组织的转变	[54⇓⇓-57]
粒型 Grain type
GS3		由232个氨基酸组成的跨膜蛋白	在翻译水平影响粒型；与DEP1或GGC2竞争性结合Gβ，缩短粒长	[65-66]
DEP1	LOC_Os09g26999	G蛋白γ亚基	控制水稻产量性状；功能获得性突变体植株表现出短穗、直立穗、密穗等产量性状；调控粒长和粒重	[67]
GGC2		异三聚体G蛋白γ亚基	调控粒长，与DEP1对粒长的正调控有加性作用	[66]
OsBRI1	LOC_Os01g52050	BR信号受体	在调控水稻粒型中起多种作用；影响细胞伸长和增殖，调控器官发育	[68-69]
OsBAK1	LOC_Os08g07760	SERK家族类受体蛋白激酶；BR信号受体激酶	影响株高、叶夹角、粒型和抗病性	[70-71]
GSK2	LOC_Os05g11730	GSK3/SHAGGY激酶	负调控BR应答基因的表达，参与水稻株型相关性状的调控	[72]
DLT	LOC_Os06g03710	由617个氨基酸组成的植物特有的GRAS家族蛋白	作用于BR应答通路下游，正向介导BR应答；影响细胞增殖及细胞数目，影响颖壳发育，负调控籽粒大小	[72,74]
GW2	LOC_Os02g14720	E3泛素连接酶	负调控细胞分裂，影响水稻的粒宽和粒重；促进泛素转移，特异识别应降解的底物，影响外稃细胞分裂，调控粒重	[75]
WG1/OsGRX8	LOC_Os02g30850	CC型谷氧还蛋白OsGRX8	调控籽粒大小，是粒宽方向细胞增殖所必需；与GW2互作并将其泛素化，从而调控WG1的稳定性	[76]
TUD1	LOC_Os03g13010	U-box家族的E3泛素连接酶	与异三聚体G蛋白α亚基D1互作调控油菜素内酯介导水稻生长，进而影响株高、粒型等相关性状	[77]