植物激素信号通路调控水稻粒型的分子机制

doi:10.13560/j.cnki.biotech.bull.1985.2023-0273

摘要/Abstract

摘要：

水稻是人类主要的粮食作物，如何有效提高其产量和品质是备受关注的重大科学问题。水稻籽粒大小是影响产量的主要因素之一，水稻籽粒发育调控的研究对利用分子设计育种提高产量、改善品质具有重要的指导意义。粒型由籽粒的长度、宽度和厚度共同决定，是受多基因调控的数量性状，是决定水稻产量和品质的关键因素之一。近年来，通过对水稻种子发育缺陷突变体的研究，发现了大量与粒型相关的数量性状位点（quantitative trait locus, QTL），一些相关基因也相继被克隆和鉴定，调控水稻粒型的复杂信号通路正在逐步阐明，其中一些基因涉及植物激素的合成、分解、运输，以及植物激素的信号转导途径。本文概述了水稻胚乳发育的基本过程，归纳了目前对胚乳发育过程中植物激素动态变化的整体认识，聚焦于控制水稻粒型的植物激素信号通路相关QTL和基因的研究现状，并对近年来取得较大进展的细胞分裂素、油菜素内酯、生长素、赤霉素、乙烯、茉莉酸和脱落酸相关通路与粒型调控的关系进行了总结和分析，进一步梳理了水稻粒型相关植物激素信号调控网络，旨在为鉴定和解析植物激素调控水稻粒型的分子机制提供参考，同时为水稻分子设计育种提供新的思路。

关键词: 水稻, 粒型, 基因, 植物激素信号

Abstract:

Rice is a major food crop for humans, and how to effectively improve its yield and quality is a major scientific concern. Rice grain size is one of the main factors affecting yield, and research on the regulation of rice grain development is an important guide for using molecular design breeding to improve yield and quality. Grain size is determined by a combination of the length, width and thickness of the grain; it is a quantitative trait regulated by multiple genes, and is one of the crucial determinants of rice yield and quality. In recent years, a large number of quantitative trait loci（QTLs）related to grain size have been identified through the study of mutants with seed development defects in rice, and some related genes have been cloned and identified, and the complex signalling pathways regulating grain size in rice are gradually being elucidated. Several studies have found that functional genes regulating rice seed development are involved in the synthesis, catabolism and transport of plant hormones as well as the signal transduction pathways of plant hormones. This review outlines the basic process of rice endosperm development, summarizes the overall the overall understanding of the dynamic changes of plant hormones during endosperm development, focuses on the current research status of QTLs related to phytohormone signalling pathways associated with rice grain size, summarizes and analyses the relationship between the pathways related to cytokinin, brassinosteroid, growth hormone, gibberellin, ethylene and jasmonic acid and grain size regulation, and further sorts out the regulation network of rice grain shape-related phytohormone signaling. It is aimed to provide a reference for identifying and analyzing the molecular mechanisms of phytohormone regulation of rice grain size, and to provide new ideas for rice molecular design breeding.

Key words: rice, grain size, gene, plant hormone signal

姚莎莎, 王晶晶, 王俊杰, 梁卫红. 植物激素信号通路调控水稻粒型的分子机制[J]. 生物技术通报, 2023, 39(8): 80-90.

YAO Sha-sha, WANG Jing-jing, WANG Jun-jie, LIANG Wei-hong. Molecular Mechanisms of Rice Grain Size Regulation Related to Plant Hormone Signaling Pathways[J]. Biotechnology Bulletin, 2023, 39(8): 80-90.

图/表 3

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植物激素 Plant hormone	基因 Gene	基因登录号 Accession No.	蛋白类型 Protein category	正/负调控 Positive（+）/ negative（-）regulator	参考文献 References
细胞分裂素 Cytokinin	OsSGL	LOC_Os02g04130	DUF1645蛋白	粒长（+）粒宽（-）	[16-17]
	DST	Os03g0786400	锌指转录因子	粒重（-）	[19]
	BG3/OsPUP4	Os01g0680200	嘌呤渗透酶	粒长（-）粒宽（-）粒重（-）	[20]
油菜素内酯Brassinolide	GS5	Os05g0158500	丝氨酸羧肽酶	粒宽（+）粒重（+）	[40]
	BRI1	Os01g0718300	受体激酶	粒宽（-）	[3]
	BAK1	Os03g0440900	BRI1相关激酶	粒长（+）粒宽（+）粒重（+）	[3]
	BZR1	Os07g0580500	BZR转录因子	粒长（+）粒宽（+）粒重（+）	[3]
	GW5	Os05g0187500	钙调素结合蛋白	粒宽（-）	[37-38]
	OFP19	Os05g0324600	OVATE家族蛋白	粒长（-）粒宽（+）	[33]
	GS2/OsGRF4	Os02g0701300	转录因子	粒长（+）粒宽（+）粒重（+）	[34]
	OsWRKY53	Os05g0343400	WRKY 转录因子	粒长（+）粒宽（+）	[30]
	GS9	Os09g0448500	转录激活因子	粒长（-）粒宽（+）	[35]
	DLT	Os06g0127600	GRAS家族蛋白	粒长（+）粒宽（-）粒重（-）	[29]
	D11	Os04g0469800	细胞色素P450蛋白	粒长（+）粒宽（+）	[24]
	D2	LOC_Os01g10040	细胞色素P450蛋白	粒长（+）粒宽（+）	[23]
	BRD1	OS03g0602300	生物合成酶	粒长（+）粒宽（+）	[21]
	BRD2	OS10g0397400	生物合成酶	粒长（+）粒宽（+）	[22]
生长素Auxin	OsARF4	Os01g0927600	生长素应答因子	粒长（-）粒宽（-）粒重（-）	[43]
	OsARF6	Os02g0164900	生长素应答因子	粒长（-）粒重（-）	[44]
	OsAUX3	Os05g0447200	生长素转运载体	粒长（-）粒重（-）	[44]
	Gnp4/LAX2	Os04g0396500	RAWUL蛋白	粒长（+）粒重（+）	[45]
	OsIAA3	Os12g0601400	Aux/IAA 蛋白	粒长（-）	[45]
	OsARF25	Os12g0613700	转录因子	粒长（+）粒宽（+）	[45]
	GSA1	Os03g0757500	UDP葡萄糖基转移酶	粒长（+）粒宽（+）粒重（+）	[48]
	TSG1	LOC_Os01g07500	色氨酸氨基转移酶	粒长（+）粒宽（+）粒重（+）	[47]
	DNR1	LOC_Os01g08270	生长素应答因子	粒重（+）	[46]
赤霉素Gibberellin	GW6	Os06g0266800	GAST 家族蛋白	粒长（+）粒宽（+）粒重（+）	[51]
	OsGASR9	Os07g0592000	GAST 家族蛋白	粒长（+）粒宽（+）粒重（+）	[52]
	OsWRKY36	Os04g0545000	WRKY转录因子	粒长（-）粒宽（-）粒重（-）	[53]
	SGD2	Os01g0643600	HD-Zip转录因子	粒长（+）粒宽（+）粒重（+）	[54]
	OsBC1	Os09g0510500	bHLH转录因子	粒长（+）粒宽（-）粒重（+）	[55]
	SNG1	Os01g0940100	己糖激酶样蛋白	粒长（+）粒宽（+）粒重（+）	[57]
乙烯Ethylene	OsERF115	Os08g0521600	ERF转录因子	粒宽（+）	[61]
茉莉酸Jasmonic acid	OsJAZ11	Os03g0180900	JAZ蛋白	粒长（+）粒宽（+）粒重（+）	[63-64]
脱落酸Abscisic acid	OsNCED3	Os03g0645900	9-顺式-环氧类胡萝卜素双加氧酶	粒长（+）粒宽（+）粒重（+）	[67]
其他调控因子Other regulators	RGB1	Os03g0669200	Gβ 亚基	粒长（+）	[72]
	qPE9-1/DEP1	Os09g0441900	Gγ亚基	粒长（+）	[74-75]
	qGL3/OsPPKL1	Os03g0646900	丝氨酸/苏氨酸磷酸酶	粒长（-）	[68-69]
	OsYUC11	Os12g0189500	黄素单加氧酶	粒重（+）	[73]

植物激素 Plant hormone	基因 Gene	基因登录号 Accession No.	蛋白类型 Protein category	正/负调控 Positive（+）/ negative（-）regulator	参考文献 References
细胞分裂素 Cytokinin	OsSGL	LOC_Os02g04130	DUF1645蛋白	粒长（+）粒宽（-）	[16-17]
	DST	Os03g0786400	锌指转录因子	粒重（-）	[19]
	BG3/OsPUP4	Os01g0680200	嘌呤渗透酶	粒长（-）粒宽（-）粒重（-）	[20]
油菜素内酯Brassinolide	GS5	Os05g0158500	丝氨酸羧肽酶	粒宽（+）粒重（+）	[40]
	BRI1	Os01g0718300	受体激酶	粒宽（-）	[3]
	BAK1	Os03g0440900	BRI1相关激酶	粒长（+）粒宽（+）粒重（+）	[3]
	BZR1	Os07g0580500	BZR转录因子	粒长（+）粒宽（+）粒重（+）	[3]
	GW5	Os05g0187500	钙调素结合蛋白	粒宽（-）	[37-38]
	OFP19	Os05g0324600	OVATE家族蛋白	粒长（-）粒宽（+）	[33]
	GS2/OsGRF4	Os02g0701300	转录因子	粒长（+）粒宽（+）粒重（+）	[34]
	OsWRKY53	Os05g0343400	WRKY 转录因子	粒长（+）粒宽（+）	[30]
	GS9	Os09g0448500	转录激活因子	粒长（-）粒宽（+）	[35]
	DLT	Os06g0127600	GRAS家族蛋白	粒长（+）粒宽（-）粒重（-）	[29]
	D11	Os04g0469800	细胞色素P450蛋白	粒长（+）粒宽（+）	[24]
	D2	LOC_Os01g10040	细胞色素P450蛋白	粒长（+）粒宽（+）	[23]
	BRD1	OS03g0602300	生物合成酶	粒长（+）粒宽（+）	[21]
	BRD2	OS10g0397400	生物合成酶	粒长（+）粒宽（+）	[22]
生长素Auxin	OsARF4	Os01g0927600	生长素应答因子	粒长（-）粒宽（-）粒重（-）	[43]
	OsARF6	Os02g0164900	生长素应答因子	粒长（-）粒重（-）	[44]
	OsAUX3	Os05g0447200	生长素转运载体	粒长（-）粒重（-）	[44]
	Gnp4/LAX2	Os04g0396500	RAWUL蛋白	粒长（+）粒重（+）	[45]
	OsIAA3	Os12g0601400	Aux/IAA 蛋白	粒长（-）	[45]
	OsARF25	Os12g0613700	转录因子	粒长（+）粒宽（+）	[45]
	GSA1	Os03g0757500	UDP葡萄糖基转移酶	粒长（+）粒宽（+）粒重（+）	[48]
	TSG1	LOC_Os01g07500	色氨酸氨基转移酶	粒长（+）粒宽（+）粒重（+）	[47]
	DNR1	LOC_Os01g08270	生长素应答因子	粒重（+）	[46]
赤霉素Gibberellin	GW6	Os06g0266800	GAST 家族蛋白	粒长（+）粒宽（+）粒重（+）	[51]
	OsGASR9	Os07g0592000	GAST 家族蛋白	粒长（+）粒宽（+）粒重（+）	[52]
	OsWRKY36	Os04g0545000	WRKY转录因子	粒长（-）粒宽（-）粒重（-）	[53]
	SGD2	Os01g0643600	HD-Zip转录因子	粒长（+）粒宽（+）粒重（+）	[54]
	OsBC1	Os09g0510500	bHLH转录因子	粒长（+）粒宽（-）粒重（+）	[55]
	SNG1	Os01g0940100	己糖激酶样蛋白	粒长（+）粒宽（+）粒重（+）	[57]
乙烯Ethylene	OsERF115	Os08g0521600	ERF转录因子	粒宽（+）	[61]
茉莉酸Jasmonic acid	OsJAZ11	Os03g0180900	JAZ蛋白	粒长（+）粒宽（+）粒重（+）	[63-64]
脱落酸Abscisic acid	OsNCED3	Os03g0645900	9-顺式-环氧类胡萝卜素双加氧酶	粒长（+）粒宽（+）粒重（+）	[67]
其他调控因子Other regulators	RGB1	Os03g0669200	Gβ 亚基	粒长（+）	[72]
	qPE9-1/DEP1	Os09g0441900	Gγ亚基	粒长（+）	[74-75]
	qGL3/OsPPKL1	Os03g0646900	丝氨酸/苏氨酸磷酸酶	粒长（-）	[68-69]
	OsYUC11	Os12g0189500	黄素单加氧酶	粒重（+）	[73]