细胞分裂素对植物生长发育的调控机理研究进展及其在水稻生产中的应用探讨

doi:10.16819/j.1001-7216.2018.8027

摘要/Abstract

摘要：

细胞分裂素(cytokinin, CTK)对植物的形态、生理及产量有重要调控作用,是调控氮素吸收、转运与代谢的主要因子之一。本文概述了氮素的吸收、转运、代谢以及CTK的代谢、转运和信号转导路径,重点阐述了CTK与氮素协作调控根-冠关系的生理机制,即反式玉米素(tZ)及其核苷(tZR)受氮素诱导在根中合成,并转运至地上部,调控地上部氮的转运及分布,影响氮代谢酶的生理特性,从而影响植株光合特性及产量;冠层中氮能够诱导异戊烯基腺嘌呤(iP)及其腺苷(iPR)的合成,并通过韧皮部转运至根系,抑制根系氮素吸收、转运,抑制根系形态建成。在此基础上,进一步论述了CTK在协调源-库关系及提高籽粒充实度方面的作用,分析了栽培措施对CTK生理代谢的影响及其与作物生长相关的机理。同时探讨了CTK上述功能应用于水稻大田生产时存在的问题,并对今后的研究方向提出了建议。

关键词: 细胞分裂素, 氮素, 作物产量

Abstract:

Cytokinins (CTKs) play important roles in regulating crop morphology, physiology, and yield. In the development of crops, CTKs are the main factors controlling the uptake, translocation, and metabolisms of nutrients, especially for nitrogen. In this paper, we summarized the uptake, translocation and metabolism of nitrogen, and synthesis, translocation and signaling of cytokinins, with a focus on the coordination of CTKs and nitrogen in regulating root-shoot relationships and their influences on crop agronomic traits. Generally, the synthesis of trans-zeatin (tZ) and tZ riboside (tZR) was up-regulated by nitrogen in roots, and they are translocated to shoots, regulating the portioning and metabolism of nitrogen, influencing the photosynthetic characteristics and yield. In shoots, the synthesis of N⁶-(Δ²-isopentenyl)- adenine (iP) and iP riboside (iPR) were up-regulated by nitrogen, and they are translocated to roots by phloem, reducing the uptake and translocation of nitrogen, influencing the root morphology. Based on current knowledge, we further discussed the role of CTKs in coordinating source-sink relationships and improving grain filling. We also analyzed the influence of cultivation practices on metabolism of CTKs and its correlation with crop growth. At the same time, we also discussed the existing problems in the application of CTKs in rice production systems. We hope that it could provide valuable information for high yielding and high efficiency rice production.

Key words: cytokinin, nitrogen, crop yield

中图分类号:

李志康, 严冬, 薛张逸, 顾逸彪, 李思嘉, 刘立军, 张耗, 王志琴, 杨建昌, 顾骏飞. 细胞分裂素对植物生长发育的调控机理研究进展及其在水稻生产中的应用探讨[J]. 中国水稻科学, 2018, 32(4): 311-324.

Zhikang LI, Dong YAN, Zhangyi XUE, Yibiao GU, Sijia LI, Lijun LIU, Hao ZHANG, Zhiqin WANG, Jianchang YANG, Junfei GU. Regulations of Plant Growth and Development by Cytokinins and Their Applications in Rice Production[J]. Chinese Journal OF Rice Science, 2018, 32(4): 311-324.

图/表 3

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基因名称 Gene name		AGI 码 AGI code	蛋白名称 Protein name		互作效应 Interaction response
基因名称 Gene name		AGI 码 AGI code	蛋白名称 Protein name		反式玉米素型细胞分裂素 tZ-CTK	细胞分裂素 Cytokinin			硝态氮 Nitrate		铵态氮 Ammonium				氮肥施用 Nitrogen application
氮生理过程 Nitrogen physiological process
无机氮素吸收 Inorganic nitrogen acquisition
硝态氮 Nitrate
AtNRT1.1		At1g12110	双亲和硝酸盐转运蛋白1.1 Dual-affinity nitrate transporter 1.1			D [101]			U [7]
AtNRT1.2		At1g69850	低亲和硝酸盐转运蛋白1.2 Low-affinity nitrate transporter 1.2			D [101]
AtNRT2.1		At1g08090	高亲和转运蛋白2.1 High-affinity nitrate transporter 2.1			D [7,15,101]			U [7]
AtNRT2.2		At1g08100	高亲和硝酸盐转运蛋白2.2 High-affinity nitrate transporter 2.2			D [101]			U [7]
AtNRT2.3		At5g60780	高亲和硝酸盐转运蛋白 2.3 High-affinity nitrate transporter 2.3			D [7,15]
AtNRT2.4		At5g60770	高亲和硝酸盐转运蛋白 2.4 High-affinity nitrate transporter 2.4			D [101]			U [7]
AtNRT2.6		At3g45060	高亲和转运蛋白2.6 High-affinity nitrate transporter 2.6			D [7]
AtNAR2.1		At5g50200	类NAR蛋白 NAR-like protein			D [15]
OsNRT1.1b		Os10g40600	双亲和硝酸盐转运蛋白1.1b Dual-affinity nitrate transporter 1.1b						U [8]
OsNPF2.4		Os03g48180	低亲和硝酸盐转运蛋白2.4 Low-affinity nitrate transporter 2.4						U [8]
OsNRT2.1		Os02g02170	高亲和硝酸盐转运蛋白2.1 High-affinity nitrate transporter 2.1						U [8]		D [8]
OsNRT2.2		Os02g02190	高亲和硝酸盐转运蛋白2.2 High-affinity nitrate transporter 2.2						U [8]		D [8]
OsNRT2.3a		Os01g50820	高亲和硝酸盐转运蛋白2.3a High-affinity nitrate transporter 2.3a						U [8]		D [8]
OsNRT2.3b		Os01g50820	高亲和硝酸盐转运蛋白2.3b High-affinity nitrate transporter 2.3b						I [8]				I [8]
OsNRT2.4		Os01g36720	可能的高亲和硝酸盐转运蛋白2.4 Putative high-affinity nitrate transporter 2.4						U [8]		D [8]
OsNAR2.1		Os02g38230	类NAR蛋白 NAR-like protein						U [8]		D [8]
铵态氮 Ammonium
AtAMT1;1		At4g13510	双亲和铵态氮转运蛋白1;1 Dual-affinity amminion transporter			D [15]			D [7]
AtAMT1;2		At1g64780	铵态氮转运蛋白1;1 Ammonium transporter 1;2			D [15]
AtAMT1;3		At3g24290	铵态氮转运蛋白1;3 Ammonium transporter 1;3			D [15]			U [7]
AtAMT1;5		At3g24290	铵态氮转运蛋白1;5 Ammonium transporter 1;5			D [15]					U [8]
OsAMT1;1		Os04g43070	双亲和铵态氮转运蛋白1;1 Dual-affinity amminion transporter 1;1								U [8]				R [8]
OsAMT1;2		Os02g40710	铵态氮转运蛋白1;2 Ammonium transporter 1;2								U [8]
OsAMT1;3		Os02g40730	铵态氮转运蛋白1;3 Ammonium transporter 1;3								U [8]
OsAMT2;1		Os05g39240	低亲和铵态氮转运蛋白2;1 Low-affinity ammonium transporter 2;1						I [8]			I [8]
OsAMT3;1		Os01g65000	可能的铵态氮转运蛋白3;1 Putative ammonium transporter 3;1						IN [8]			IN [8]
氮素冠层分布 Nitrogen distribution in canopy
AtNRT1.3		At3g21670	低亲和硝酸盐转运蛋白1.3 Low-affinity nitrate transporter 1.3		U [15]
AtNRT1.4		At2g26690	低亲和硝酸盐转运蛋白1.4 Low-affinity nitrate transporter 1.4		U [15]
AtNRT1.7		At1g69870	低亲和硝酸盐转运蛋白1.7 Low-affinity nitrate transporter 1.7		U [15]
AtNRT2.7		At5g14570	高亲和硝酸盐转运蛋白2.7 High-affinity nitrate transporter 2.7		U [15]
基因名称 Gene name	AGI 码 AGI code			蛋白名称 Protein name			互作效应 Interaction response
基因名称 Gene name	AGI 码 AGI code			蛋白名称 Protein name			反式玉米素型细胞分裂素 tZ-CTK	细胞分裂素 Cytokinis		硝态氮 Nitrate				铵态氮 Ammonium		氮肥施用 Nitrogen application
氮代谢 Nitrogen metabolism
硝态氮还原 Nitrate reduction
NIA1	At1g77760			硝酸还原酶1 Nitrate reductase 1				U [7]		U [7]
NIA2	At1g37130			硝酸还原酶2 Nitrate reductase 2						U [7]
NII	At2g15620			亚硝酸还原酶 Nitrite reductase						U [7]
辅因子 Cofactor
CNX2	At2g31955			钼喋呤合成酶 Molybdopterin synthetase						U [7]
UPM	At5g40850			硫-腺苷甲硫氨酸尿卟啉原III S-adenosylmethionine uroporphyrinogen III						U [7]
氨同化 Ammonia assimilation
GLN1;1	At5g37600			谷氨酰胺合成酶1;1 Glutamine synthetase 1;1				D [7]		D [7]
GLN1;2	At1g66200			谷氨酰胺合成酶1;2 Glutamine synthetase 1;2				D [7]		U [7]
GLN2	At5g35630			谷氨酰胺合成酶2 Glutamine synthetase 2						U [7]
GLT1	At5g53460			NADH-谷氨酸合成酶1 NADH-glutamate synthase 1						U [7]
氨基酸代谢 Amino acid metabolism
GDH1	At5g18170			谷氨酸脱氢酶1 Glutamate dehydrogenase 1				U [7]
GDH2	At5g07440			谷氨酸脱氢酶2 Glutamate dehydrogenase 2				U [7]
ASP1	At2g30970			天冬氨酸转氨酶1 Aspartase aminotransferase 1				D [7]		U [7]
ASN1	At3g47340			天冬酰胺合成酶1 Asparagine synthetase 1						U [7]
ASN2	At5g65010			天冬酰胺合成酶2 Asparagine synthetase 2				U [7]		U [7]
细胞分裂素生理过程 Cytokinin physiological process
细胞分裂素合成 Cytokinins biosynthesis
AtIPT3	At3g63110			异戊烯基转移酶3 Isopentenyltransferase 3						U [7]
OsIPT4	Os03g0810100			异戊烯基转移酶4 Isopentenyltransferase 4												U [91]
OsIPT5	Os07g0211700			异戊烯基转移酶5 Isopentenyltransferase 5												U [91]
OsIPT7	Os05g0551700			异戊烯基转移酶5 Isopentenyltransferase 7												U [91]
OsIPT8	Os01g0688300			异戊烯基转移酶8 Isopentenyltransferase 8												U [91]
CYP735A2	At1g67110			细胞分裂素反式羟化酶 Cytokinin trans-hydroxylase				U [7]		U [7]
细胞分裂素代谢 Cytokinins metabolism
CKX4	At4g29740			细胞分裂素氧化酶4 Cytokinin oxidase 4				U [7]		U [7]
CKX5	At1g75450			细胞分裂素氧化酶5 Cytokinin oxidase 5				U [7]
转录调控 Transcriptional control
ARR3	At1g59940			A型反应调控因子3 Type-A response regulator 3				U [7]		U [7]
ARR4	At1g10470			A型反应调控因子4 Type-A response regulator 4				U [7]
ARR5	At3g48100			A型反应调控因子5 Type-A response regulator 5				U [7]		U [7]
ARR6	At5g62920			A型反应调控因子6 Type-A response regulator 6				U [7]		U [7]
ARR7	At1g19050			A型反应调控因子7 Type-A response regulator 7				U [7]		U [7]
ARR8	At2g41310			A型反应调控因子8 Type-A response regulator 8				U [7]		U [7]
ARR9	At3g57040			A型反应调控因子9 Type-A response regulator 9				U [7]		U [7]
ARR15	At1g74890			A型反应调控因子15 Type-A response regulator 15				U [7]		U [7]
ARR16	At1g74890			A型反应调控因子16 Type-A response regulator 16				U [7]

基因名称 Gene name		AGI 码 AGI code	蛋白名称 Protein name		互作效应 Interaction response
基因名称 Gene name		AGI 码 AGI code	蛋白名称 Protein name		反式玉米素型细胞分裂素 tZ-CTK	细胞分裂素 Cytokinin			硝态氮 Nitrate		铵态氮 Ammonium				氮肥施用 Nitrogen application
氮生理过程 Nitrogen physiological process
无机氮素吸收 Inorganic nitrogen acquisition
硝态氮 Nitrate
AtNRT1.1		At1g12110	双亲和硝酸盐转运蛋白1.1 Dual-affinity nitrate transporter 1.1			D [101]			U [7]
AtNRT1.2		At1g69850	低亲和硝酸盐转运蛋白1.2 Low-affinity nitrate transporter 1.2			D [101]
AtNRT2.1		At1g08090	高亲和转运蛋白2.1 High-affinity nitrate transporter 2.1			D [7,15,101]			U [7]
AtNRT2.2		At1g08100	高亲和硝酸盐转运蛋白2.2 High-affinity nitrate transporter 2.2			D [101]			U [7]
AtNRT2.3		At5g60780	高亲和硝酸盐转运蛋白 2.3 High-affinity nitrate transporter 2.3			D [7,15]
AtNRT2.4		At5g60770	高亲和硝酸盐转运蛋白 2.4 High-affinity nitrate transporter 2.4			D [101]			U [7]
AtNRT2.6		At3g45060	高亲和转运蛋白2.6 High-affinity nitrate transporter 2.6			D [7]
AtNAR2.1		At5g50200	类NAR蛋白 NAR-like protein			D [15]
OsNRT1.1b		Os10g40600	双亲和硝酸盐转运蛋白1.1b Dual-affinity nitrate transporter 1.1b						U [8]
OsNPF2.4		Os03g48180	低亲和硝酸盐转运蛋白2.4 Low-affinity nitrate transporter 2.4						U [8]
OsNRT2.1		Os02g02170	高亲和硝酸盐转运蛋白2.1 High-affinity nitrate transporter 2.1						U [8]		D [8]
OsNRT2.2		Os02g02190	高亲和硝酸盐转运蛋白2.2 High-affinity nitrate transporter 2.2						U [8]		D [8]
OsNRT2.3a		Os01g50820	高亲和硝酸盐转运蛋白2.3a High-affinity nitrate transporter 2.3a						U [8]		D [8]
OsNRT2.3b		Os01g50820	高亲和硝酸盐转运蛋白2.3b High-affinity nitrate transporter 2.3b						I [8]				I [8]
OsNRT2.4		Os01g36720	可能的高亲和硝酸盐转运蛋白2.4 Putative high-affinity nitrate transporter 2.4						U [8]		D [8]
OsNAR2.1		Os02g38230	类NAR蛋白 NAR-like protein						U [8]		D [8]
铵态氮 Ammonium
AtAMT1;1		At4g13510	双亲和铵态氮转运蛋白1;1 Dual-affinity amminion transporter			D [15]			D [7]
AtAMT1;2		At1g64780	铵态氮转运蛋白1;1 Ammonium transporter 1;2			D [15]
AtAMT1;3		At3g24290	铵态氮转运蛋白1;3 Ammonium transporter 1;3			D [15]			U [7]
AtAMT1;5		At3g24290	铵态氮转运蛋白1;5 Ammonium transporter 1;5			D [15]					U [8]
OsAMT1;1		Os04g43070	双亲和铵态氮转运蛋白1;1 Dual-affinity amminion transporter 1;1								U [8]				R [8]
OsAMT1;2		Os02g40710	铵态氮转运蛋白1;2 Ammonium transporter 1;2								U [8]
OsAMT1;3		Os02g40730	铵态氮转运蛋白1;3 Ammonium transporter 1;3								U [8]
OsAMT2;1		Os05g39240	低亲和铵态氮转运蛋白2;1 Low-affinity ammonium transporter 2;1						I [8]			I [8]
OsAMT3;1		Os01g65000	可能的铵态氮转运蛋白3;1 Putative ammonium transporter 3;1						IN [8]			IN [8]
氮素冠层分布 Nitrogen distribution in canopy
AtNRT1.3		At3g21670	低亲和硝酸盐转运蛋白1.3 Low-affinity nitrate transporter 1.3		U [15]
AtNRT1.4		At2g26690	低亲和硝酸盐转运蛋白1.4 Low-affinity nitrate transporter 1.4		U [15]
AtNRT1.7		At1g69870	低亲和硝酸盐转运蛋白1.7 Low-affinity nitrate transporter 1.7		U [15]
AtNRT2.7		At5g14570	高亲和硝酸盐转运蛋白2.7 High-affinity nitrate transporter 2.7		U [15]
基因名称 Gene name	AGI 码 AGI code			蛋白名称 Protein name			互作效应 Interaction response
基因名称 Gene name	AGI 码 AGI code			蛋白名称 Protein name			反式玉米素型细胞分裂素 tZ-CTK	细胞分裂素 Cytokinis		硝态氮 Nitrate				铵态氮 Ammonium		氮肥施用 Nitrogen application
氮代谢 Nitrogen metabolism
硝态氮还原 Nitrate reduction
NIA1	At1g77760			硝酸还原酶1 Nitrate reductase 1				U [7]		U [7]
NIA2	At1g37130			硝酸还原酶2 Nitrate reductase 2						U [7]
NII	At2g15620			亚硝酸还原酶 Nitrite reductase						U [7]
辅因子 Cofactor
CNX2	At2g31955			钼喋呤合成酶 Molybdopterin synthetase						U [7]
UPM	At5g40850			硫-腺苷甲硫氨酸尿卟啉原III S-adenosylmethionine uroporphyrinogen III						U [7]
氨同化 Ammonia assimilation
GLN1;1	At5g37600			谷氨酰胺合成酶1;1 Glutamine synthetase 1;1				D [7]		D [7]
GLN1;2	At1g66200			谷氨酰胺合成酶1;2 Glutamine synthetase 1;2				D [7]		U [7]
GLN2	At5g35630			谷氨酰胺合成酶2 Glutamine synthetase 2						U [7]
GLT1	At5g53460			NADH-谷氨酸合成酶1 NADH-glutamate synthase 1						U [7]
氨基酸代谢 Amino acid metabolism
GDH1	At5g18170			谷氨酸脱氢酶1 Glutamate dehydrogenase 1				U [7]
GDH2	At5g07440			谷氨酸脱氢酶2 Glutamate dehydrogenase 2				U [7]
ASP1	At2g30970			天冬氨酸转氨酶1 Aspartase aminotransferase 1				D [7]		U [7]
ASN1	At3g47340			天冬酰胺合成酶1 Asparagine synthetase 1						U [7]
ASN2	At5g65010			天冬酰胺合成酶2 Asparagine synthetase 2				U [7]		U [7]
细胞分裂素生理过程 Cytokinin physiological process
细胞分裂素合成 Cytokinins biosynthesis
AtIPT3	At3g63110			异戊烯基转移酶3 Isopentenyltransferase 3						U [7]
OsIPT4	Os03g0810100			异戊烯基转移酶4 Isopentenyltransferase 4												U [91]
OsIPT5	Os07g0211700			异戊烯基转移酶5 Isopentenyltransferase 5												U [91]
OsIPT7	Os05g0551700			异戊烯基转移酶5 Isopentenyltransferase 7												U [91]
OsIPT8	Os01g0688300			异戊烯基转移酶8 Isopentenyltransferase 8												U [91]
CYP735A2	At1g67110			细胞分裂素反式羟化酶 Cytokinin trans-hydroxylase				U [7]		U [7]
细胞分裂素代谢 Cytokinins metabolism
CKX4	At4g29740			细胞分裂素氧化酶4 Cytokinin oxidase 4				U [7]		U [7]
CKX5	At1g75450			细胞分裂素氧化酶5 Cytokinin oxidase 5				U [7]
转录调控 Transcriptional control
ARR3	At1g59940			A型反应调控因子3 Type-A response regulator 3				U [7]		U [7]
ARR4	At1g10470			A型反应调控因子4 Type-A response regulator 4				U [7]
ARR5	At3g48100			A型反应调控因子5 Type-A response regulator 5				U [7]		U [7]
ARR6	At5g62920			A型反应调控因子6 Type-A response regulator 6				U [7]		U [7]
ARR7	At1g19050			A型反应调控因子7 Type-A response regulator 7				U [7]		U [7]
ARR8	At2g41310			A型反应调控因子8 Type-A response regulator 8				U [7]		U [7]
ARR9	At3g57040			A型反应调控因子9 Type-A response regulator 9				U [7]		U [7]
ARR15	At1g74890			A型反应调控因子15 Type-A response regulator 15				U [7]		U [7]
ARR16	At1g74890			A型反应调控因子16 Type-A response regulator 16				U [7]