Towards Understanding the Genetic Regulations of Endosperm Development in Rice

doi:10.16819/j.1001-7216.2021.210307

Abstract

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

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

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.

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.

Figures/Tables 4

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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]