水稻花粉外壁孢粉素合成及运输相关基因研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2020-0421

摘要/Abstract

摘要：

花粉外壁是花粉正常发育的保障,可为小孢子建立结构支持从而保护其免受恶劣条件的影响,其主要由孢粉素组成。花粉外壁的正常发育是一个复杂的多基因表达调控过程,水稻中目前已分离和鉴定出一系列参与到花粉外壁合成特别是孢粉素代谢相关的基因。本文归纳总结了22个与水稻孢粉素合成及运输相关的基因及其功能、分子机制及调控网络,旨为进一步研究水稻雄性生殖发育分子机制提供理论依据。

关键词: 水稻, 花粉外壁, 孢粉素

Abstract:

The pollen exine is the guarantee for the normal pollen development,and the main component is sporpollen,which can establish the structural support for the microspore cytoplasm to protect it from harsh conditions. The normal development of the pollen exine is a complex process of multi-gene expression regulation,there are a large number of genes involved in the development of pollen exine,especially the metabolism of sporopollenin,have been isolated and identified in rice. This paper summarized 22 genes related to rice sporopollen synthesis and transport,as well as their functions,molecular mechanisms and regulatory networks,which is aimed to provide a theoretical basis for further research on the molecular mechanism of rice male reproductive development.

Key words: rice, pollen exine, sporopollen

王联红, 张秋云, 刘家林, 沈亚琦, 欧阳林娟, 贺浩华, 胡丽芳. 水稻花粉外壁孢粉素合成及运输相关基因研究进展[J]. 生物技术通报, 2020, 36(12): 170-177.

WANG Lian-hong, ZHANG Qiu-yun, LIU Jia-lin, SHEN Ya-qi, OUYANG Lin-juan, HE Hao-hua, HU Li-fang. Advances in the Study of Genes Related to Pollen Exine Sporopollen Synthesis and Transport in Rice[J]. Biotechnology Bulletin, 2020, 36(12): 170-177.

图/表 3

参考文献 41

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基因	编码蛋白	功能	文献
CYP703A3 Os08g0131100	链内羟化酶月桂酸（中链脂肪酸）	催化孢粉素合成中间产物C10至C14脂肪酸的链内羟基化	[6]
CYP704B2 Os03g0168600	v-羟基化脂肪酸	催化孢粉素合成中间产物C16至C18长链脂肪酸的ω-羟基化	[7]
GAMYB Os01g0812000	一种R2R3MYB转录因子	激活CYP703A3表达,间接调控孢粉素前体物质合成	[8]
DPW Os03g0167600	脂肪酰基载体蛋白还原酶	将棕榈酰ACP和辅酶A酯还原为脂肪醇,是孢粉素生物合成的初级脂肪醇	[9]
DPW2 Os01g0924933	细胞质定位的BAHD酰基转移酶	催化羟基化的孢粉素合成中间产物ω-羟基脂肪酸转化为羟基肉桂酰辅酶A	[10]
OsACOS12 Os04g0310800	酰基辅酶A合成酶	催化C18：1形成C18：1-CoA,参与孢粉质前体所需的脂酰辅酶A合成	[11]
OsPKS1 Os10g0484800	PKS III超家族蛋白	催化次生代谢物的合成。催化脂酰CoA缩合生成孢粉素合成所需的3-α-吡喃酮	[12]
OsPKS2 Os07g0411300	聚酮化合物合成酶	催化丙二酰CoA缩合生成孢粉素合成所需的4-α-吡喃酮	[13]
OsTKPR1 Os09g0493500	四肽α-吡喃酮还原酶	催化PKS产生的四酮的羰基还原为仲醇,是孢粉素前体合成必须物质	[14]
Wda1 Os10g0471100	一个富含组氨酸基序的蛋白质	还原超长链脂肪酸为各种所需要的脂肪醇类,影响孢粉素前体的合成	[15]
OsCER1 Os02g0621300	脂肪酸羟化酶	还原超长链脂肪酸（VLCFAs）为各种所需要的脂肪醇类,调控孢粉素前体的合成	[16]
OsNP1 Os10g0524500	一种假定的葡萄糖-甲醇-胆碱氧化还原酶	具有催化醇氧化成相应醛的生物化学活性。可催化脂肪酸中链羟基脱水生成环氧化物,对孢粉素合成产生影响	[17]
HTH1 Os04g0573100	一种假定的葡萄糖-甲醇-胆碱氧化还原酶	催化孢粉素合成中间产物长链α-,邻二羧酸脂肪酸的氧化	[18]
OsGPAT3 Os11g0679700	甘油-3-磷酸酰基转移酶	OsGPAT3直接或间接影响各种孢粉素合成代谢相关基因	[19]

基因	编码蛋白	功能	文献
CYP703A3 Os08g0131100	链内羟化酶月桂酸（中链脂肪酸）	催化孢粉素合成中间产物C10至C14脂肪酸的链内羟基化	[6]
CYP704B2 Os03g0168600	v-羟基化脂肪酸	催化孢粉素合成中间产物C16至C18长链脂肪酸的ω-羟基化	[7]
GAMYB Os01g0812000	一种R2R3MYB转录因子	激活CYP703A3表达,间接调控孢粉素前体物质合成	[8]
DPW Os03g0167600	脂肪酰基载体蛋白还原酶	将棕榈酰ACP和辅酶A酯还原为脂肪醇,是孢粉素生物合成的初级脂肪醇	[9]
DPW2 Os01g0924933	细胞质定位的BAHD酰基转移酶	催化羟基化的孢粉素合成中间产物ω-羟基脂肪酸转化为羟基肉桂酰辅酶A	[10]
OsACOS12 Os04g0310800	酰基辅酶A合成酶	催化C18：1形成C18：1-CoA,参与孢粉质前体所需的脂酰辅酶A合成	[11]
OsPKS1 Os10g0484800	PKS III超家族蛋白	催化次生代谢物的合成。催化脂酰CoA缩合生成孢粉素合成所需的3-α-吡喃酮	[12]
OsPKS2 Os07g0411300	聚酮化合物合成酶	催化丙二酰CoA缩合生成孢粉素合成所需的4-α-吡喃酮	[13]
OsTKPR1 Os09g0493500	四肽α-吡喃酮还原酶	催化PKS产生的四酮的羰基还原为仲醇,是孢粉素前体合成必须物质	[14]
Wda1 Os10g0471100	一个富含组氨酸基序的蛋白质	还原超长链脂肪酸为各种所需要的脂肪醇类,影响孢粉素前体的合成	[15]
OsCER1 Os02g0621300	脂肪酸羟化酶	还原超长链脂肪酸（VLCFAs）为各种所需要的脂肪醇类,调控孢粉素前体的合成	[16]
OsNP1 Os10g0524500	一种假定的葡萄糖-甲醇-胆碱氧化还原酶	具有催化醇氧化成相应醛的生物化学活性。可催化脂肪酸中链羟基脱水生成环氧化物,对孢粉素合成产生影响	[17]
HTH1 Os04g0573100	一种假定的葡萄糖-甲醇-胆碱氧化还原酶	催化孢粉素合成中间产物长链α-,邻二羧酸脂肪酸的氧化	[18]
OsGPAT3 Os11g0679700	甘油-3-磷酸酰基转移酶	OsGPAT3直接或间接影响各种孢粉素合成代谢相关基因	[19]

基因	编码蛋白	功能	文献
OsABCG3 Os01g0836600	ABC转运蛋白	ABC转运蛋白是孢粉素前体跨膜转运所必需,该基因利用ATP衍生的能量,将脂质分子跨膜运往花粉壁	[20]
OsABCG15 Os06g0607700	ABC转运蛋白	ABC转运蛋白是孢粉素前体跨膜转运所必需,该基因利用ATP衍生的能量,运输孢粉素前体至花药室	[21]
OsABCG26 Os10g0494300	ABC转运蛋白	ABC转运蛋白是孢粉素前体跨膜转运所必需,该基因利用ATP衍生的能量,运输孢粉素前体至花粉壁	[22]
PTC1 Os09g0449000	PHD-finger蛋白	作为具有转录激活活性的转录因子起作用,靶向一些参与孢粉素前体生物合成和转运的基因	[23]
OsEMF2b Os09g0306800	是Zeste抑制子,编码核心的PcG蛋白	介导OsLFL1和OsMADS4的H3K27me3沉积,可调控PTC1和GAMYB的表达介导孢粉素前体的合成	[24]
OsLecRK-S.7 Os02g0459600	一种质膜定位的豆荚凝集素受体激酶,编码LecRLK蛋白	该基因高表达,导致突变体花粉中外壁增厚,影响突变体花药中孢粉素运输	[25]
DPW3 Os02g0100700	一种α整合素样蛋白	催化羟基化的孢粉素合成中间产物脂肪酸转化为脂肪醇	[27]
OsC6/Osg6B Os11g0582500	脂质转移蛋白	具有脂质结合活性,参与孢粉素前体的转运	[28]

基因	编码蛋白	功能	文献
OsABCG3 Os01g0836600	ABC转运蛋白	ABC转运蛋白是孢粉素前体跨膜转运所必需,该基因利用ATP衍生的能量,将脂质分子跨膜运往花粉壁	[20]
OsABCG15 Os06g0607700	ABC转运蛋白	ABC转运蛋白是孢粉素前体跨膜转运所必需,该基因利用ATP衍生的能量,运输孢粉素前体至花药室	[21]
OsABCG26 Os10g0494300	ABC转运蛋白	ABC转运蛋白是孢粉素前体跨膜转运所必需,该基因利用ATP衍生的能量,运输孢粉素前体至花粉壁	[22]
PTC1 Os09g0449000	PHD-finger蛋白	作为具有转录激活活性的转录因子起作用,靶向一些参与孢粉素前体生物合成和转运的基因	[23]
OsEMF2b Os09g0306800	是Zeste抑制子,编码核心的PcG蛋白	介导OsLFL1和OsMADS4的H3K27me3沉积,可调控PTC1和GAMYB的表达介导孢粉素前体的合成	[24]
OsLecRK-S.7 Os02g0459600	一种质膜定位的豆荚凝集素受体激酶,编码LecRLK蛋白	该基因高表达,导致突变体花粉中外壁增厚,影响突变体花药中孢粉素运输	[25]
DPW3 Os02g0100700	一种α整合素样蛋白	催化羟基化的孢粉素合成中间产物脂肪酸转化为脂肪醇	[27]
OsC6/Osg6B Os11g0582500	脂质转移蛋白	具有脂质结合活性,参与孢粉素前体的转运	[28]