水稻粒型基因克隆和调控机制研究进展

doi:10.16819/j.1001-7216.2018.7016

摘要/Abstract

摘要：

水稻粒型是影响其产量和品质的重要性状,阐明其遗传调控机理,有助于提高水稻单产和改良品质。水稻粒型性状主要包括粒长、粒宽、粒厚、长/宽比,属于数量性状,受胚、胚乳及母体植株等不同遗传体系的控制。随着水稻功能基因组学和重测序技术的快速发展,目前已经定位超过400个与水稻粒型相关的数量性状位点(QTL),并已克隆了60个水稻粒型基因,涉及植物激素、泛素-蛋白酶体通路、丝裂原活化蛋白激酶(MAPK)信号通路、G蛋白信号通路及表观修饰等多个调控通路。本文对水稻粒型基因克隆及其调控机制的研究进展进行了系统总结和梳理,并对这些基因在育种上的利用价值进行了评价。

关键词: 水稻, 粒型, 基因, 调控机制

Abstract:

Grain shape is an important trait that affects the yield and quality of rice, so it is necessary to understand the genetic regulation mechanism of grain shape to improve rice yield and quality. Grain shape is characterized by a combination of grain length, grain width, grain thickness, and grain length-to-width ratio, belonging to quantitative traits, which is controlled by different genetic systems, such as embryo, endosperm and maternal plant. With the rapid development of rice functional genomics and re-sequencing technology, more than 400 QTLs related to rice grain shape have been located at present, and at least 60 genes associated with rice grain traits have been identified. Some rice grain shape regulation pathways were identified, including phytohormones, the ubiquitin-proteasome pathway, the mitogen-activated protein kinase (MAPK) signaling pathway, epigenetic modification, the G protein signaling pathways. In this review, we systematically summarize and sort out the research progress of the cloning and functional analysis of rice grain shape genes, and evaluate the utilization value of rice grain shape genes in rice breeding for high yield and good quality.

Key words: rice(Oryza sativa L.), grain shape, gene, regulation of mechanism

中图分类号:

刘喜, 牟昌铃, 周春雷, 程治军, 江玲, 万建民. 水稻粒型基因克隆和调控机制研究进展[J]. 中国水稻科学, 2018, 32(1): 1-11.

Xi LIU, Changling MOU, Chunlei ZHOU, Zhijun CHENG, Ling JIANG, Jianmin WAN. Research Progress on Cloning and Regulation Mechanism of Rice Grain Shape Genes[J]. Chinese Journal OF Rice Science, 2018, 32(1): 1-11.

图/表 3

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基因 Gene
D2/SMG11	1	细胞色素P450加氧酶,参与BR生物合成	隐性突变体,小粒	[10-11]
D61/OsBRI1	1	BR受体激酶	隐性突变体,小粒	[12-15]
RAV6	2	B3 DNA结合结构域蛋白,介导油菜素内酯稳态,受表观遗传修饰调控	显性突变体,小粒	[16]
SDG725	2	H3K36 甲基转移酶,参与调节BR相关基因的表达	隐性突变体,小粒	[17-18]
SMG1	2	有丝分裂激活的蛋白激酶激酶4,可能作为MAPK通路和BR间连接因子	隐性突变体,小粒	[19]
TH1/AFD1	2	DUF640结构域的蛋白,影响细胞分裂和扩张	隐性突变体,小粒	[20-21]
FUWA	2	NHL结构域蛋白,限制细胞过度分裂	隐性突变体,小粒	[22]
GW2	2	环型E3泛素连接酶,负调节细胞分裂	隐性突变体,宽粒	[23-25]
OsGRF4/GS2/GL2	2	生长调控因子,与BR负调控因子GSK2互作	转基因过表达株,大粒	[26-28]
PGL2	2	非典型的不结合DNA的碱性螺旋-环-螺旋蛋白,与APG互作	转基因过表达株,长粒	[29]
BG1	3	未知蛋白,参与调节生长素的转运	隐性突变体,小粒	[30]
PGL1	3	非典型的不结合DNA的碱性螺旋-环-螺旋蛋白	转基因过表达株,长粒	[31]
RGB1	3	G蛋白β亚基,正向调控细胞增殖	隐性突变体,小粒	[32-33]
GS3	3	G蛋白的γ亚基,负调控粒重	转基因过表达株,长粒	[34-38]
OsPPKL1/GL3.1	3	蛋白磷酸酶,调控细胞周期蛋白T1;3	转基因干扰株,长粒	[39-41]
BRD1/OsBR6ox	3	细胞色素P450加氧酶,参与BR生物合成	隐性突变体,小粒	[42-43]
LTS1/OsNaPRT1	3	烟酸磷酸核糖转移酶,影响烟酰胺的含量	隐性突变体,小粒	[44]
TUD1	3	U-box家族的E3泛素连接酶,参与BR应答	隐性突变体,短粒	[45]
XIAO	4	LRR 激酶,调控BR信号传递和动态平衡以及细胞周期	隐性突变体,小粒	[46]
D11/CPB1	4	细胞色素P450加氧酶,参与BR生物合成	隐性突变体,小粒	[47-48]
An-1	4	bHLH转录因子,调控细胞分裂	隐性突变体,短粒	[49]
OsBSK3	4	BR信号激酶	转基因干扰株,小粒	[50]
Flo2	4	含TPR结构域蛋白	隐性突变体,小粒	[51]
GW5/qSW5	5	核蛋白,调节细胞分裂	转基因干扰株,宽粒	[52-53]
GS5	5	丝氨酸羧肽酶,与 BAK1互作,参与BR信号	转基因过表达株,大粒	[54-55]
GSK2	5	与拟南芥BIN2同源的类GSK3/SHAGGY激酶,BR信号负调控因子	转基因干扰株,大粒	[56]
OsCYP51G3	5	钝叶醇14α-脱甲基酶	转基因干扰株,短粒	[57]
OsLAC	5	漆酶蛋白,影响BR信号	转基因干扰株,大粒	[58]
SRS3	5	驱动蛋白13 基因家族成员,影响细胞纵向长度	隐性突变体,小圆粒	[59]
OsPPKL2	5	含有Kelch重复域的蛋白磷酸酶,正向调控粒长	转基因过表达株,长粒	[41]
APG	5	bHLH蛋白,PGL1拮抗因子	转基因干扰株,长粒	[29]
D1	5	G蛋白亚基,调控细胞分裂	隐性突变体,小粒	[60-63]
TGW6	6	IAA-葡萄糖水解酶,水解IAA-葡萄糖成IAA和葡萄糖	转基因干扰株,大粒	[64]
GW6a	6	组蛋白乙酰转移酶,增加细胞数和加速灌浆速率,增大颖壳	转基因干扰株,小粒	[65]
GS6/DLT	6	GRAS基因家族,BR信号基因	隐性突变体,小粒	[66-67]
BU1	6	螺旋-环-螺旋蛋白,BR信号基因	转基因过表达株,大粒	[68]
OsARF19	6	生长素响应因子	转基因过表达株,瘪粒	[69]
HGW	6	泛素相关结构域蛋白,可能直接通过GIF1 控制水稻籽粒和质量	隐性突变体,细长粒	[70]
OsMAPK6/DSG1	6	有丝分裂激活的蛋白激酶,影响细胞增殖以及BR信号和稳态	隐性突变体,小粒	[71]
GL7/GW7/SLG7	7	TONNEAU1募集基序蛋白	转基因过表达株,长粒	[72-74]
GLW7	7	SPL家族转录因子	转基因过表达株,长粒	[75]
OsBZR1	7	BR信号因子	转基因过表达株,大粒	[76]
BG2/GE	7	细胞色素P450加氧酶,促进细胞增殖	显性突变体,大粒	[77-78]
SRS1/EP2	7	未知蛋白	隐性突变体,小圆粒	[79-80]
SLG	8	类BAHD酰基转移酶,调控BR的稳态	半显性突变体,细长粒	[81]
BAK1	8	BR信号受体BRI1的激酶,BR共受体	转基因过表达株,小粒	[82]
qGW8/OsSPL16	8	含SBP结构域的转录因子,结合GW7启动子,抑制其表达	转基因过表达株,细长粒	[83]
OsFIE1	8	多梳蛋白抑制复合体的类Esc核心元件,参与H3K27me3介导的基因抑制过程	显性突变体,小粒	[84]
OsFEI2	8	具有特异的组蛋白H3甲基转移酶活性,负责组蛋白H3第27位赖氨酸上三甲基化的形成	隐性突变体,小粒	[85]
GAD1	8	一个表皮模式因子类蛋白EPFL1,促进细胞分裂	转基因干扰株,短粒	[86]
GDD1	9	驱动蛋白4家族基因,控制水稻细胞周期进程和细胞壁的属性	隐性突变体,小粒	[87]
SG1	9	未知蛋白,与BR相关	转基因干扰株,短粒	[88]
DEP1/DN1/qNGR9	9	G蛋白γ亚基	隐性突变体,小粒	[89-93]
BRD2	10	拟南芥DIM1/DWF1同源基因,参与BR生物合成	隐性突变体,小粒	[94]
SRS5/TID1	11	微管蛋白	隐性突变体,小圆粒	[95-96]
OsGIF1	11	GRF互作因子	转基因过表达株,大粒	[27]
OsPPKL3	12	含有Kelch重复域的蛋白磷酸酶,负向调控粒长	转基因过表达株,短粒	[41]
miR1848		调控靶基因OsCYP51G3	转基因过表达株,短粒	[57]
miR397		调控靶基因OsLAC	转基因过表达株,大粒	[58]
miR396		调控靶基因GS2	转基因过表达株,小粒	[27]

基因 Gene
D2/SMG11	1	细胞色素P450加氧酶,参与BR生物合成	隐性突变体,小粒	[10-11]
D61/OsBRI1	1	BR受体激酶	隐性突变体,小粒	[12-15]
RAV6	2	B3 DNA结合结构域蛋白,介导油菜素内酯稳态,受表观遗传修饰调控	显性突变体,小粒	[16]
SDG725	2	H3K36 甲基转移酶,参与调节BR相关基因的表达	隐性突变体,小粒	[17-18]
SMG1	2	有丝分裂激活的蛋白激酶激酶4,可能作为MAPK通路和BR间连接因子	隐性突变体,小粒	[19]
TH1/AFD1	2	DUF640结构域的蛋白,影响细胞分裂和扩张	隐性突变体,小粒	[20-21]
FUWA	2	NHL结构域蛋白,限制细胞过度分裂	隐性突变体,小粒	[22]
GW2	2	环型E3泛素连接酶,负调节细胞分裂	隐性突变体,宽粒	[23-25]
OsGRF4/GS2/GL2	2	生长调控因子,与BR负调控因子GSK2互作	转基因过表达株,大粒	[26-28]
PGL2	2	非典型的不结合DNA的碱性螺旋-环-螺旋蛋白,与APG互作	转基因过表达株,长粒	[29]
BG1	3	未知蛋白,参与调节生长素的转运	隐性突变体,小粒	[30]
PGL1	3	非典型的不结合DNA的碱性螺旋-环-螺旋蛋白	转基因过表达株,长粒	[31]
RGB1	3	G蛋白β亚基,正向调控细胞增殖	隐性突变体,小粒	[32-33]
GS3	3	G蛋白的γ亚基,负调控粒重	转基因过表达株,长粒	[34-38]
OsPPKL1/GL3.1	3	蛋白磷酸酶,调控细胞周期蛋白T1;3	转基因干扰株,长粒	[39-41]
BRD1/OsBR6ox	3	细胞色素P450加氧酶,参与BR生物合成	隐性突变体,小粒	[42-43]
LTS1/OsNaPRT1	3	烟酸磷酸核糖转移酶,影响烟酰胺的含量	隐性突变体,小粒	[44]
TUD1	3	U-box家族的E3泛素连接酶,参与BR应答	隐性突变体,短粒	[45]
XIAO	4	LRR 激酶,调控BR信号传递和动态平衡以及细胞周期	隐性突变体,小粒	[46]
D11/CPB1	4	细胞色素P450加氧酶,参与BR生物合成	隐性突变体,小粒	[47-48]
An-1	4	bHLH转录因子,调控细胞分裂	隐性突变体,短粒	[49]
OsBSK3	4	BR信号激酶	转基因干扰株,小粒	[50]
Flo2	4	含TPR结构域蛋白	隐性突变体,小粒	[51]
GW5/qSW5	5	核蛋白,调节细胞分裂	转基因干扰株,宽粒	[52-53]
GS5	5	丝氨酸羧肽酶,与 BAK1互作,参与BR信号	转基因过表达株,大粒	[54-55]
GSK2	5	与拟南芥BIN2同源的类GSK3/SHAGGY激酶,BR信号负调控因子	转基因干扰株,大粒	[56]
OsCYP51G3	5	钝叶醇14α-脱甲基酶	转基因干扰株,短粒	[57]
OsLAC	5	漆酶蛋白,影响BR信号	转基因干扰株,大粒	[58]
SRS3	5	驱动蛋白13 基因家族成员,影响细胞纵向长度	隐性突变体,小圆粒	[59]
OsPPKL2	5	含有Kelch重复域的蛋白磷酸酶,正向调控粒长	转基因过表达株,长粒	[41]
APG	5	bHLH蛋白,PGL1拮抗因子	转基因干扰株,长粒	[29]
D1	5	G蛋白亚基,调控细胞分裂	隐性突变体,小粒	[60-63]
TGW6	6	IAA-葡萄糖水解酶,水解IAA-葡萄糖成IAA和葡萄糖	转基因干扰株,大粒	[64]
GW6a	6	组蛋白乙酰转移酶,增加细胞数和加速灌浆速率,增大颖壳	转基因干扰株,小粒	[65]
GS6/DLT	6	GRAS基因家族,BR信号基因	隐性突变体,小粒	[66-67]
BU1	6	螺旋-环-螺旋蛋白,BR信号基因	转基因过表达株,大粒	[68]
OsARF19	6	生长素响应因子	转基因过表达株,瘪粒	[69]
HGW	6	泛素相关结构域蛋白,可能直接通过GIF1 控制水稻籽粒和质量	隐性突变体,细长粒	[70]
OsMAPK6/DSG1	6	有丝分裂激活的蛋白激酶,影响细胞增殖以及BR信号和稳态	隐性突变体,小粒	[71]
GL7/GW7/SLG7	7	TONNEAU1募集基序蛋白	转基因过表达株,长粒	[72-74]
GLW7	7	SPL家族转录因子	转基因过表达株,长粒	[75]
OsBZR1	7	BR信号因子	转基因过表达株,大粒	[76]
BG2/GE	7	细胞色素P450加氧酶,促进细胞增殖	显性突变体,大粒	[77-78]
SRS1/EP2	7	未知蛋白	隐性突变体,小圆粒	[79-80]
SLG	8	类BAHD酰基转移酶,调控BR的稳态	半显性突变体,细长粒	[81]
BAK1	8	BR信号受体BRI1的激酶,BR共受体	转基因过表达株,小粒	[82]
qGW8/OsSPL16	8	含SBP结构域的转录因子,结合GW7启动子,抑制其表达	转基因过表达株,细长粒	[83]
OsFIE1	8	多梳蛋白抑制复合体的类Esc核心元件,参与H3K27me3介导的基因抑制过程	显性突变体,小粒	[84]
OsFEI2	8	具有特异的组蛋白H3甲基转移酶活性,负责组蛋白H3第27位赖氨酸上三甲基化的形成	隐性突变体,小粒	[85]
GAD1	8	一个表皮模式因子类蛋白EPFL1,促进细胞分裂	转基因干扰株,短粒	[86]
GDD1	9	驱动蛋白4家族基因,控制水稻细胞周期进程和细胞壁的属性	隐性突变体,小粒	[87]
SG1	9	未知蛋白,与BR相关	转基因干扰株,短粒	[88]
DEP1/DN1/qNGR9	9	G蛋白γ亚基	隐性突变体,小粒	[89-93]
BRD2	10	拟南芥DIM1/DWF1同源基因,参与BR生物合成	隐性突变体,小粒	[94]
SRS5/TID1	11	微管蛋白	隐性突变体,小圆粒	[95-96]
OsGIF1	11	GRF互作因子	转基因过表达株,大粒	[27]
OsPPKL3	12	含有Kelch重复域的蛋白磷酸酶,负向调控粒长	转基因过表达株,短粒	[41]
miR1848		调控靶基因OsCYP51G3	转基因过表达株,短粒	[57]
miR397		调控靶基因OsLAC	转基因过表达株,大粒	[58]
miR396		调控靶基因GS2	转基因过表达株,小粒	[27]