玉米籽粒淀粉生物合成及调控因素研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2024-0922

摘要/Abstract

摘要：

玉米作为一种在全球范围内广泛栽培的作物，不仅是人类饮食结构中不可或缺的粮食来源，而且在饲料产业及清洁能源领域同样发挥着重要作用。其籽粒中富含的淀粉，构成了玉米生物量的主体，直接决定了玉米的产量水平，是农业生产和食品加工中不可或缺的原料。然而，随着全球人口的不断增长、城市化进程的加速推进，以及气候变化带来的农业用地减少和自然灾害频发，全球粮食供应体系正面临着前所未有的挑战。玉米作为世界重要农作物之一，通过解析其淀粉生物合成途径培育高产优良品种变得十分重要。本文总结了参与玉米淀粉生物合成途径的关键酶及转运蛋白，并探讨这些因子在淀粉合成过程中的具体作用。同时，对合成途径中关键酶的突变体籽粒表型进行综述，分析基因突变对淀粉产量与品质的影响。此外，本文还总结了调控淀粉生物合成的各种因素，探讨如何通过优化这些因素来提高玉米的产量与品质。最后，本文展望了未来的研究方向，旨在为优质高产玉米的分子设计育种提供坚实的理论基础与实践指导。

关键词: 玉米, 胚乳, 淀粉, 淀粉合成酶, 调控因素

Abstract:

Maize, as a widely cultivated crop worldwide, is not only an indispensable food source in the human diet, but also plays an important role in the feed industry and clean energy. The starch rich in its kernels constitutes the main body of maize biomass and directly determines the yield level of maize. It is an indispensable raw material in agricultural production and food processing. However, with the continuous growth of the global population, the acceleration of urbanization, and the reduction of agricultural land and frequent natural disasters caused by climate change, the global food supply system is facing unprecedented challenges. As important crops in the world,it is very crtical to cultivate high-yield varieties by analyzing its starch biosynthesis pathway. This paper summarizes the key enzymes and transporters involved in the maize starch biosynthesis pathway and discusses the specific roles of these factors in the process of starch synthesis. Concurrently, this paper reviews the grain phenotypes of mutants of key enzymes in the synthesis pathway, and analyze the effects of gene mutations on starch yield and quality. In addition, this paper also summarizes the various factors regulating starch biosynthesis and discusses how to improve the yield and quality of maize by optimizing these factors. Finally, this paper looks forward to the future research direction, aiming to provide a solid theoretical basis and practical guidance for molecular design breeding of high quality and high yield maize.

Key words: maize, endosperm, starch, starch synthetase, regulatory factors

王涛, 胡社伟, 张宇, 邓文文, 尚春缘, 王婉艺. 玉米籽粒淀粉生物合成及调控因素研究进展[J]. 生物技术通报, 2025, 41(3): 1-13.

WANG Tao, HU She-wei, ZHANG Yu, DENG Wen-wen, SHANG Chun-yuan, WANG Wan-yi. Research Progress in Starch Biosynthesis and Regulatory Factors in Maize Kernel[J]. Biotechnology Bulletin, 2025, 41(3): 1-13.

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酶/转运蛋白	同工酶	基因名称	基因号	玉米突变体表型
Enzymes/Transport proteins	Isoenzymes	Gene name	Gene ID	Phenotypes of maize mutants
蔗糖合成酶	SUS-sh1	Shrunken1（Sh1）	Zm00001d045042	籽粒冠部塌陷，淀粉含量显著降低^［10］
	SUS1	Sus1	Zm00001d047253	籽粒无明显缺陷表型、淀粉含量无显著变化，新鲜籽粒蔗糖含量累积^［10］
	SUS2	Sus2	Zm00001d029091	与sus1表型基本一致^［10-11］
ADP-葡萄糖焦磷酸化酶	AGPL3	Shrunken2 （Sh2）/Zmagpl3	Zm00001d044129	籽粒淀粉含量显著下降、蔗糖水平上升，籽粒胚乳严重皱缩、粒重降低^［12］
	AGPL2	Agplemzm/ZmagpL2	Zm00001d005546	/
	AGPL1	Agpllzm/ZmagpL1	Zm00001d019266	叶片淀粉含量显著下降；籽粒无明显缺陷表型、粒重略微降低^［13］
	AGPS2	Brittle2 （Bt2）/Zmagps2	Zm00001d050032	籽粒淀粉含量显著下降、蔗糖水平上升，籽粒胚乳严重皱缩、质地酥脆易碎、粒重降低^［14］
	AGPS3	Agpsemzm/Zmagps3	Zm00001d039131	籽粒胚淀粉含量显著下降、胚乳淀粉略微下降，籽粒无明显缺陷表型、粒重略微降低^［13］
	AGPS1	Agpslzm/Zmagps1	Zm00001d032385	叶片淀粉含量显著下降、蔗糖含量上升；籽粒无明显缺陷表型、粒重略微降低^［15］
ADPG转运蛋白	Brittle1	Brittle1 （Bt1）	Zm00001d015746	籽粒胚乳中淀粉含量严重下降，籽粒皱缩、多棱、粒重显著降低^［16-17］
淀粉合成酶	GBSSI	Waxy （Wx）	Zm00001d045462	籽粒支链淀粉几乎占总淀粉含量100%，籽粒胚乳不透明、无光泽、外观晦暗呈蜡质状^［18］
	GBSSIIa	ZmGBSSIIa	Zm00001d019479	/
	GBSSIIb	ZmGBSSIIb	Zm00001d027242	/
	SSI	SSI	Zm00001d045261	/
	SSIIa	Sugary2 （Su2）	Zm00001d037234	籽粒淀粉含量下降、直链淀粉上升10%-15%，籽粒胚乳半透明、呈玻璃状、无光泽^［19-20］
	SSIII	Dull1 （Du1）	Zm00001d000002	根据遗传背景的不同籽粒直链淀粉含量略微或显著上升，籽粒胚乳呈玻璃状、无光泽、外观钝化^［21-22］
淀粉分支酶	SBEI	Sbe1	Zm00001d014844	籽粒和叶片淀粉性质和含量几乎没有变化^［23-24］
	SBEIIa	Sbe2a	Zm00001d011301	叶片早期衰老，叶片淀粉累积、淀粉分支数减少、分支长度增加；籽粒淀粉性质和含量几乎没有变化^［25］
	SBEIIb	Amylose Extender （Ae）	Zm00001d016684	籽粒直链淀粉含量由20%上升至50%，籽粒胚乳透明呈玻璃状、体积变小、种皮皱缩^［26-27］
去淀粉分支酶	ISA1	Sugary1 （Su1）	Zm00001d049753	籽粒总淀粉和支链淀粉降低、植物糖原、多种单糖以及蔗糖含量积累，籽粒外观皱缩凹陷^［28-29］
	ISA2	Sugary4 （Su4）	Zm00001d038121	籽粒总淀粉与野生型基本一致，籽粒无明显缺陷表型^［30-31］
	ISA3	ISO3	Zm00001d020799	/
	PUL	Zpu1	Zm00001d004438	叶片和籽粒中淀粉分解代谢受阻，籽粒中淀粉性质和含量几乎没有变化，籽粒无明显缺陷表型^［32］
淀粉磷酸化酶	PHOL/PHO1	Shrunken4 （Sh4）	Zm00001d034074	籽粒总淀粉含量显著降低，籽粒胚乳皱缩且粉质^［8-9］
淀粉磷酸化酶	PHOH/PHO2	ZmPHOH/Chl3	Zm00001d042842	叶片细胞叶绿体中的瞬时淀粉累积，叶片褪绿、光合效率显著降低^［8-9］

酶/转运蛋白	同工酶	基因名称	基因号	玉米突变体表型
Enzymes/Transport proteins	Isoenzymes	Gene name	Gene ID	Phenotypes of maize mutants
蔗糖合成酶	SUS-sh1	Shrunken1（Sh1）	Zm00001d045042	籽粒冠部塌陷，淀粉含量显著降低^［10］
	SUS1	Sus1	Zm00001d047253	籽粒无明显缺陷表型、淀粉含量无显著变化，新鲜籽粒蔗糖含量累积^［10］
	SUS2	Sus2	Zm00001d029091	与sus1表型基本一致^［10-11］
ADP-葡萄糖焦磷酸化酶	AGPL3	Shrunken2 （Sh2）/Zmagpl3	Zm00001d044129	籽粒淀粉含量显著下降、蔗糖水平上升，籽粒胚乳严重皱缩、粒重降低^［12］
	AGPL2	Agplemzm/ZmagpL2	Zm00001d005546	/
	AGPL1	Agpllzm/ZmagpL1	Zm00001d019266	叶片淀粉含量显著下降；籽粒无明显缺陷表型、粒重略微降低^［13］
	AGPS2	Brittle2 （Bt2）/Zmagps2	Zm00001d050032	籽粒淀粉含量显著下降、蔗糖水平上升，籽粒胚乳严重皱缩、质地酥脆易碎、粒重降低^［14］
	AGPS3	Agpsemzm/Zmagps3	Zm00001d039131	籽粒胚淀粉含量显著下降、胚乳淀粉略微下降，籽粒无明显缺陷表型、粒重略微降低^［13］
	AGPS1	Agpslzm/Zmagps1	Zm00001d032385	叶片淀粉含量显著下降、蔗糖含量上升；籽粒无明显缺陷表型、粒重略微降低^［15］
ADPG转运蛋白	Brittle1	Brittle1 （Bt1）	Zm00001d015746	籽粒胚乳中淀粉含量严重下降，籽粒皱缩、多棱、粒重显著降低^［16-17］
淀粉合成酶	GBSSI	Waxy （Wx）	Zm00001d045462	籽粒支链淀粉几乎占总淀粉含量100%，籽粒胚乳不透明、无光泽、外观晦暗呈蜡质状^［18］
	GBSSIIa	ZmGBSSIIa	Zm00001d019479	/
	GBSSIIb	ZmGBSSIIb	Zm00001d027242	/
	SSI	SSI	Zm00001d045261	/
	SSIIa	Sugary2 （Su2）	Zm00001d037234	籽粒淀粉含量下降、直链淀粉上升10%-15%，籽粒胚乳半透明、呈玻璃状、无光泽^［19-20］
	SSIII	Dull1 （Du1）	Zm00001d000002	根据遗传背景的不同籽粒直链淀粉含量略微或显著上升，籽粒胚乳呈玻璃状、无光泽、外观钝化^［21-22］
淀粉分支酶	SBEI	Sbe1	Zm00001d014844	籽粒和叶片淀粉性质和含量几乎没有变化^［23-24］
	SBEIIa	Sbe2a	Zm00001d011301	叶片早期衰老，叶片淀粉累积、淀粉分支数减少、分支长度增加；籽粒淀粉性质和含量几乎没有变化^［25］
	SBEIIb	Amylose Extender （Ae）	Zm00001d016684	籽粒直链淀粉含量由20%上升至50%，籽粒胚乳透明呈玻璃状、体积变小、种皮皱缩^［26-27］
去淀粉分支酶	ISA1	Sugary1 （Su1）	Zm00001d049753	籽粒总淀粉和支链淀粉降低、植物糖原、多种单糖以及蔗糖含量积累，籽粒外观皱缩凹陷^［28-29］
	ISA2	Sugary4 （Su4）	Zm00001d038121	籽粒总淀粉与野生型基本一致，籽粒无明显缺陷表型^［30-31］
	ISA3	ISO3	Zm00001d020799	/
	PUL	Zpu1	Zm00001d004438	叶片和籽粒中淀粉分解代谢受阻，籽粒中淀粉性质和含量几乎没有变化，籽粒无明显缺陷表型^［32］
淀粉磷酸化酶	PHOL/PHO1	Shrunken4 （Sh4）	Zm00001d034074	籽粒总淀粉含量显著降低，籽粒胚乳皱缩且粉质^［8-9］
淀粉磷酸化酶	PHOH/PHO2	ZmPHOH/Chl3	Zm00001d042842	叶片细胞叶绿体中的瞬时淀粉累积，叶片褪绿、光合效率显著降低^［8-9］

基因名称	基因号	转录因子家族	靶基因/潜在靶基因	启动子核心元件	参考文献
Gene name	Gene ID	Transcription factor family	Target genes/Potential target genes	The core element of promoter	References
ZmThx20	Zm00001d017420	GT-2 trihelix	PHOH， SBEIV， GBSSI， Isoamylase1	/	［90］
ZmTCP7	Zm00001d010730	TCP	Bt2， Bt1， SSI， SSIIa， SSIIIa， Sh2， SSIV	GAACCCCAC	［91］
ZmDof3	Zm00001d035651	DOF	Du1， Su2	AAAG	［92］
PBF	Zm00001d005100	DOF	PPDK1， PPDK2， SSIII， SSIIa， SBE	AAAG	［77］
ZmDOF36	Zm00001d029512	DOF	AGPS1a， AGPL1， GBSSI， SSIIa， ISA1， ISA3	/	［93］
ZmMADS1a	Zm00001d041781	MADS	AGPL1， AGPS1a， SBEIIb， GBSSI， ISA1， SSIIa	/	［94］
ZmES22	Zm00001d031620	MADS	SBEIIa， SBEIIb， GBSSI， GBSSII， AGPS， AGPL， ISA1， ISA2， ISA3， PUL， SSI， SSIIa， SSIIIa， SSIVa	/	［95］
ZmNAC126	Zm00001d005028	NAC	GBSSI， SSIIa， SSIV， ISA1， ISA2， Sh2， BT2， GBSSI， SSIIIa， BT1	CACG	［96］
ZmNAC34/ZmNAC128	Zm00001d040189	NAC	Bt2， Zpu1， SSI， SUS1， GBSSI， SBEIIb	ACGCAA	［82， 84］
ZmNAC130	Zm00001d008403	NAC	Bt2， Zpu1， SSI， SUS1， GBSSI， SBEIIb	ACGCAA	［82， 84］
ZmaNAC36	Zm00001d003052	NAC	AGPL2， AGPS2， SSI， GBSSIIb， SBEI	TTGCGTGTT	［85］
ZmABI4	Zm00001d038001	AP2/EREBP	SSI	CACCG	［97］
ZmEREB156	Zm00001d026447	AP2/EREBP	Sh2， SSIIIa	/	［98］
ZmEREB94	Zm00001d020043	AP2/EREBP	SSI， Sh2， GBSSI， Bt1， SBEI， SSIIa， SSIV， ISA1	/	［99］
ZmPLATZ2	Zm00001d029437	PLATZ	SSI， SSIIa， SSIIIa， SSIV， SSV， ISA1， ISA2， Sh2， Bt2， GBSSI	CAAAAAAA	［100］
ZmMYB14	Zm00001d021537	MYB	BT1， SSI， SSII， SBEI， GBSSI， Sh2， Bt2， ISA1	TAACTG	［86］
ZmMYB138	Zm00001d043131	MYB	BT2， Ae1， SSIIa	/	［89］
ZmMYB115	Zm00001d000361	MYB	Du1， Wx	/	［89］
ZmABI19	Zm00001d011712	B3 domain	O2， PBF， ZmbZIP22， ZmNAC130， O11	TGCATG	［88］
ZmbZIP29	Zm00001d034571	bZIP	O2	/	［101］
mEmBP-1	Zm00001d019166	bZIP	Sbe1	/	［102］
ZmbZIP91/ZmbZIP22	Zm00001d021191	bZIP	AGPS1， SSI， SSIIIa， ISA1， AGPL1a， SBEIIa， GBSSIa， ISA2， PHOL， SSIa	ACTCAT	［78， 103］
O2	Zm00001d018971	bZIP	PPDK1， PPDK2， SSIII， Sh1， SUS1， SUS2	ACGT	［10， 77］
O11	Zm00001d003677	bHLH	O2， PBF， ZmDof3	CACGTG	［87］
ZmICE1	Zm00001d042263	bHLH	SSIIa， GBSSI	CANNTG	［104］
NKD1	Zm00001d002654	IDD	O2	TTGTCGT	［105］
NKD2	Zm00001d026113	IDD	O2	/	［105］

基因名称	基因号	转录因子家族	靶基因/潜在靶基因	启动子核心元件	参考文献
Gene name	Gene ID	Transcription factor family	Target genes/Potential target genes	The core element of promoter	References
ZmThx20	Zm00001d017420	GT-2 trihelix	PHOH， SBEIV， GBSSI， Isoamylase1	/	［90］
ZmTCP7	Zm00001d010730	TCP	Bt2， Bt1， SSI， SSIIa， SSIIIa， Sh2， SSIV	GAACCCCAC	［91］
ZmDof3	Zm00001d035651	DOF	Du1， Su2	AAAG	［92］
PBF	Zm00001d005100	DOF	PPDK1， PPDK2， SSIII， SSIIa， SBE	AAAG	［77］
ZmDOF36	Zm00001d029512	DOF	AGPS1a， AGPL1， GBSSI， SSIIa， ISA1， ISA3	/	［93］
ZmMADS1a	Zm00001d041781	MADS	AGPL1， AGPS1a， SBEIIb， GBSSI， ISA1， SSIIa	/	［94］
ZmES22	Zm00001d031620	MADS	SBEIIa， SBEIIb， GBSSI， GBSSII， AGPS， AGPL， ISA1， ISA2， ISA3， PUL， SSI， SSIIa， SSIIIa， SSIVa	/	［95］
ZmNAC126	Zm00001d005028	NAC	GBSSI， SSIIa， SSIV， ISA1， ISA2， Sh2， BT2， GBSSI， SSIIIa， BT1	CACG	［96］
ZmNAC34/ZmNAC128	Zm00001d040189	NAC	Bt2， Zpu1， SSI， SUS1， GBSSI， SBEIIb	ACGCAA	［82， 84］
ZmNAC130	Zm00001d008403	NAC	Bt2， Zpu1， SSI， SUS1， GBSSI， SBEIIb	ACGCAA	［82， 84］
ZmaNAC36	Zm00001d003052	NAC	AGPL2， AGPS2， SSI， GBSSIIb， SBEI	TTGCGTGTT	［85］
ZmABI4	Zm00001d038001	AP2/EREBP	SSI	CACCG	［97］
ZmEREB156	Zm00001d026447	AP2/EREBP	Sh2， SSIIIa	/	［98］
ZmEREB94	Zm00001d020043	AP2/EREBP	SSI， Sh2， GBSSI， Bt1， SBEI， SSIIa， SSIV， ISA1	/	［99］
ZmPLATZ2	Zm00001d029437	PLATZ	SSI， SSIIa， SSIIIa， SSIV， SSV， ISA1， ISA2， Sh2， Bt2， GBSSI	CAAAAAAA	［100］
ZmMYB14	Zm00001d021537	MYB	BT1， SSI， SSII， SBEI， GBSSI， Sh2， Bt2， ISA1	TAACTG	［86］
ZmMYB138	Zm00001d043131	MYB	BT2， Ae1， SSIIa	/	［89］
ZmMYB115	Zm00001d000361	MYB	Du1， Wx	/	［89］
ZmABI19	Zm00001d011712	B3 domain	O2， PBF， ZmbZIP22， ZmNAC130， O11	TGCATG	［88］
ZmbZIP29	Zm00001d034571	bZIP	O2	/	［101］
mEmBP-1	Zm00001d019166	bZIP	Sbe1	/	［102］
ZmbZIP91/ZmbZIP22	Zm00001d021191	bZIP	AGPS1， SSI， SSIIIa， ISA1， AGPL1a， SBEIIa， GBSSIa， ISA2， PHOL， SSIa	ACTCAT	［78， 103］
O2	Zm00001d018971	bZIP	PPDK1， PPDK2， SSIII， Sh1， SUS1， SUS2	ACGT	［10， 77］
O11	Zm00001d003677	bHLH	O2， PBF， ZmDof3	CACGTG	［87］
ZmICE1	Zm00001d042263	bHLH	SSIIa， GBSSI	CANNTG	［104］
NKD1	Zm00001d002654	IDD	O2	TTGTCGT	［105］
NKD2	Zm00001d026113	IDD	O2	/	［105］

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