作物叶绿体发育的温度调控研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2025-0756

摘要/Abstract

摘要：

叶绿体是植物进行光合作用的核心场所，作为植物细胞特有的半自主性细胞器，其发育过程受到植物内在生长发育信号和外界环境信号的双重调控。其中，环境温度是影响叶绿体发育的重要外界因素，它通过调控叶绿体膜系统、形态结构、质体分裂与分化等多个方面，最终影响叶绿体的发育及功能。本综述首先概述了叶绿体的结构、功能及发育过程对环境温度的响应机制；其次，通过系统梳理植物叶色突变体的研究进展，总结了叶绿体发育受温度调控的分子基础；并进一步从叶绿体基因的转录调控、转录后调控以及蛋白质合成与稳态三个层次，深入讨论了叶绿体发育对温度变化的应答机制。最后，从机制研究和生产实践两个角度，对未来如何利用叶色基因揭示叶绿体发育的温度响应机制提出了总结与展望。综上，本综述全面分析了环境温度变化对作物叶绿体发育及生理功能的影响，并进一步探讨温度对作物光合作用的潜在影响，以期为全球气候变化下的作物广适性高光效分子设计育种提供理论支持与实践指导。

关键词: 叶绿体发育, 温度, 质体基因表达, 共转录调控

Abstract:

Chloroplasts are the key organelles responsible for photosynthesis in plants. As semi-autonomous structures unique to plant cells, their development is regulated by both internal growth and developmental signals as well as external environmental cues. Among these factors, temperature plays a critical role in shaping chloroplast development. It influences various aspects, such as the chloroplast membrane system, morphology, plastid division, and differentiation, ultimately impacting their development and function. This review begins with a concise overview of the structure, function, and developmental processes of chloroplasts in response to temperature fluctuations. It then summarizes research progress on plant leaf color mutants, shedding light on the molecular basis of temperature regulation in chloroplast development. Additionally, the review explores the mechanisms of temperature regulation in chloroplasts across three levels: Transcriptional regulation of chloroplast genes, post-transcriptional regulation of chloroplast genes, and protein synthesis and homeostasis in chloroplast. Finally, it discusses future directions, offering insights into how leaf color-related genes can be leveraged to uncover the mechanisms underlying temperature-responsive chloroplast development, with applications in both research and agricultural practices. In conclusion, this review thoroughly examines how environmental temperature changes affect chloroplast development and physiological functions in crops, while also exploring the broader implications for photosynthesis. By addressing these impacts, it aims to provide theoretical insights and practical guidance for developing molecular breeding strategies that enhance crop adaptability and photosynthetic efficiency in the face of global climate change.

Key words: chloroplast development, temperature, plastid gene expression, co-transcriptional regulation

蒋韫博, 陈雪雪, 赵玉胜. 作物叶绿体发育的温度调控研究进展[J]. 生物技术通报, 2025, 41(10): 20-31.

JIANG Yun-bo, CHEN Xue-xue, ZHAO Yu-sheng. Research Progress in Temperature Regulation of Chloroplast Development in Crops[J]. Biotechnology Bulletin, 2025, 41(10): 20-31.

图/表 2

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突变体 Mutant	类型 Type	基因 Gene	通路 Pathway	蛋白功能 Protein function	物种 Species	温度 Temperature （℃）	参考文献 Reference
近等基因系Thermo-tolerance3^WYJ	-	TT3.2	-	未知功能的跨膜蛋白，定位于叶绿体类囊体	水稻	42	［20］
albino embryo and seedling	白化	AES	基因转录	PPR蛋白，RNA剪接	拟南芥	-	［22］
rice albino 1	白化	RA1	蛋白翻译	氨酰tRNA合成酶，蛋白合成相关	水稻	-	［23］
albino leaf 1	白化	AL1	核糖体合成	八肽重复蛋白，调节核糖体RNA和核糖体蛋白的稳态	水稻	-	［24］
albino leaf 2	白化	AL2	RNA加工	叶绿体I型和II型内含子剪接	水稻	-	［25］
CN19M06	白化返绿	TSCA1	-	定位于2A染色体长臂6.5 Mb的区间内，有111个注释基因	小麦	15	［26］
tic100^cue8	返绿	TIC100	蛋白稳态	叶绿体TIC蛋白，参与叶绿体蛋白质输入，影响逆行信号	拟南芥	-	［27］
white green leaf 2	白化返绿	WGL2	核糖体合成	定位于叶绿体的核糖体蛋白	水稻	-	［28］
thermo-sensitive virescent 2	白化返绿	TSV2	蛋白翻译	氨酰tRNA合成酶	水稻	20 24 28	［29］
virescent 2	返绿	V2	核苷酸代谢	鸟苷酸激酶	水稻	20	［30］
cde4	白化返绿	CDE4	RNA加工	PPR蛋白，叶绿体RNA剪切	水稻	20	［31］
delayed virescence 4	返绿	DV4	-	定位于7D染色体短臂1.46 Mb的区间内，有3个注释基因	小麦	-	［32-33］
ch1	褪绿	CAO	色素代谢	叶绿素合成	拟南芥	-	［34］
yellow leaf color 1	褪绿	YL1	-	定位于7D染色体8.75 Mb的区间内，有110个注释基因	小麦	-	［35］
chlorophyll	褪绿	TaCHLI-7D	色素代谢	镁螯合酶I亚基，叶绿素合成	小麦	-	［36］
wsl3	条纹	WSL3	基因转录	PPR蛋白，PEP活性	水稻	25	［38］
wlp2s， wlp2w	条纹	WLP2	基因转录	PEP转录相关	水稻	32	［39］
wsl4	条纹	WSL4	RNA加工	PPR蛋白，参与II型内含子剪接	水稻	20	［40］
young seedling stripe1	条纹	YSS1	基因转录	DUF3727家族蛋白，参与调控PEP活性	水稻	20	［41］
wsl5	条纹	WSL5	RNA加工	PPR蛋白，RNA编辑和剪接	水稻	20	［42］
wsl6	条纹	WSL6	核糖体合成	GTP结合蛋白，核糖体合成	水稻	20	［43］
wsl9	条纹	WSL9	RNA加工	HNH内切酶，编辑效率	水稻	20	［44］
young leaf white stripe	条纹	YLWS	RNA加工	PPR蛋白，参与II型内含子剪接	水稻	20	［45］
B03S	斑马纹	-	色素代谢	未知基因，可能与玉米黄素合成有关	水稻	-	［46］
brittle culm and zebra leaf 1	斑马纹	BZ1	糖代谢	UDP-半乳糖/葡萄糖差向异构酶	水稻	-	［47］
zebra leaf 15	斑马纹	Z15	-	受体激酶	水稻	18	［48］
white kernel mutant 3735	斑马纹	ZmPTOX1	色素代谢	类胡萝卜素合成	玉米	-	［49-50］
zebra7	斑马纹	ZB7	色素代谢	MEP途径	玉米	32	［51］
thermo-sensitive chlorophyll-deficient mutant 1	白化	TCM1	基因转录	PEP转录相关	水稻	20	［53］
tcd12	白化	TCD12/ SIG2A	基因转录	RNA聚合酶Sigma因子	水稻	23	［54］
Dular	白化	DUA1	基因转录 RNA加工	PPR蛋白，与SIG1和WSP1互作分别介导叶绿体基因表达和RNA编辑	水稻	18	［55-56］
low-temperature-sensitive	白化	OsPUS1	RNA加工	叶绿体核糖体RNA的假尿苷化修饰	水稻	20	［57］
temperature-sensitive chlorophyll-deficient 11	白化	TSCD11	蛋白翻译	氨酰tRNA合成酶，蛋白合成相关	水稻	35	［59］
tcd9	白化	TCD9	蛋白折叠蛋白稳态	伴侣蛋白，质体分裂	水稻	20	［60］
T-DNA插入突变体oshsp70cp1	褪绿	HSP70CP1	蛋白折叠蛋白稳态	蛋白转运与组装	水稻	37	［61］
yellow variegated1	不均匀的褪绿	FtsH5/VAR1	蛋白稳态	FtsH蛋白酶，光系统稳定性	拟南芥	-	［62］
yellow variegated2	不均匀的褪绿	FtsH2/VAR2	蛋白稳态	FtsH蛋白酶，光系统稳定性	拟南芥	8	［62-63］
thermosensitive 36	褪绿	FtsH11	蛋白稳态	FtsH蛋白酶，光反应蛋白复合体稳定性	拟南芥	30	［64］
SALK_033047	褪绿	FtsH11	蛋白稳态	FtsH蛋白酶，光反应蛋白复合体稳定性	拟南芥	26	［65］
T-DNA插入突变体Salk_028162C	白化	AT3G06730	-	水稻硫氧还蛋白WP2同源蛋白	拟南芥	22	［66］
white panicle2	白化	WP2	RNA加工	硫氧还蛋白，叶绿体RNA编辑	水稻	35	［66］
tcd5	白化返绿	TCD5	基因转录	氧化还原蛋白，单加氧酶，叶绿体基因表达	水稻	20	［74］
temperature-sensitive virescent	白化返绿	TSV	基因转录	氧化还原蛋白，与TrxZ相互作用，PEP转录相关	水稻	20	［75］
white stripe leaf	条纹	WSL	基因转录	PPR蛋白，PEP基因转录	水稻	20	［76］

suppressor of ospus1-1 10	白化	SOP10	RNA加工	PPR蛋白，调控线粒体RNA剪接和编辑	水稻	22	［77］
virescent-1	返绿	V1/NUS1	RNA降解	细菌转录终止抑制子，叶绿体RNA代谢	水稻	20	［78］
RNAi株系	褪绿	CRP1	RNA降解	mRNA稳定性	水稻	4	［79］
umpk	褪绿	UMPK	核苷酸代谢	尿苷5′-单磷酸激酶	水稻	6	［80］
virescent 3	返绿	V3	核苷酸代谢	核糖核苷酸还原酶大亚基，叶绿体DNA合成	水稻	20	［81］
wsl12	条纹	WSL12	核苷酸代谢	核苷二磷酸激酶	水稻	22 34	［82］
stripe and drooping leaf	条纹	SDL/ RNRS1	核苷酸代谢	核糖核苷酸还原酶小亚基	水稻	20	［83］
temperature-sensitive chlorophyll-deficient 3	白化	TCD3	核苷酸代谢	假尿苷合酶，叶绿体RNA代谢，RNA修饰	水稻	20	［84］
pale-green leaf 12	褪绿	PGL12	核糖体合成	PPR蛋白，叶绿体16S rRNA加工	水稻	20	［85］
tcd11	白化	TCD11	核糖体合成	叶绿体核糖体亚基	水稻	20	［86］
white leaf and panicles 1	返绿	WLP1	核糖体合成	核糖体L13蛋白	水稻	23	［87］
tcm5	白化	TCM5	蛋白稳态	蛋白酶，光合系统II保护	水稻	32	［88］
RNAi株系	褪绿	NOA1	蛋白翻译核糖体合成	YqeH同源蛋白，叶绿体中核糖体组装和蛋白翻译	水稻	22	［89-91］
chlorophyll deficient 1	褪绿	CDE1/GluRS	蛋白翻译	tRNA合酶，蛋白合成	水稻	32	［92］
XN1376B	白化返绿	-	-	-	小麦	-	［93］
Zm-murE-2	白化	MurE	基因转录	PEP活性，叶绿体逆行信号	玉米	-	［94］
ppr5-1	白化	PPR5	基因转录	PPR蛋白，叶绿体逆行信号	玉米	-	［94］

hsp21	褪绿	HSP21	基因转录	PEP功能，与pTAC5互作	拟南芥	30	［95］
RNAi株系	褪绿	pTAC5	基因转录	质体转录活性染色质蛋白	拟南芥	30	［95］
cp29a cp31a	褪绿	CP29ACP31A	RNA降解	RNA结合蛋白，RNA稳定性和加工	拟南芥	8	［96］
thermo-sensitive mutant in leafcolor 1	褪绿	DG1/TSL1	RNA加工	PPR蛋白，叶绿体RNA编辑和剪切，MORF2，DYW1，DYW2互作	拟南芥	29	［97］
T-DNA插入突变体rare1-1，rare1-2	褪绿	RARE1	RNA加工	PPR蛋白，通过RNA编辑影响脂肪酸合成	拟南芥	28	［98］
thermo-sensitive mutant in leafcolor 2	褪绿	FtsHi5/TSL2	蛋白稳态	叶绿体蛋白输入复合体成员	拟南芥	16	［99］
ggps1-2， ggps1-3	褪绿	GGPS1	色素代谢	叶绿素合成途径	拟南芥	27	［100］
gun5-1	褪绿	GUN5	色素代谢	镁离子螯合酶，质核信号穿梭	拟南芥	4	［101］
paleface1	褪绿	PFC1	核糖体合成	rRNA腺嘌呤去甲基化酶，核糖体合成	拟南芥	5	［102］
rbd1-1	褪绿	RBD1	核糖体合成	RNA结合蛋白，23S rRNA加工	拟南芥	4	［103］
rps1	白化	RPS1	核糖体合成	叶绿体核糖体蛋白	拟南芥	38	［104］
rps5	褪绿	RPS5	核糖体合成	叶绿体核糖体功能	拟南芥	4	［105］
crass-1， crass-2	褪绿	CRASS	蛋白翻译	核糖体相关蛋白，叶绿体蛋白翻译效率	拟南芥	4	［106］
svr3-1	褪绿	SVR3	蛋白翻译	翻译延伸因子	拟南芥	8	［107］

突变体 Mutant	类型 Type	基因 Gene	通路 Pathway	蛋白功能 Protein function	物种 Species	温度 Temperature （℃）	参考文献 Reference
近等基因系Thermo-tolerance3^WYJ	-	TT3.2	-	未知功能的跨膜蛋白，定位于叶绿体类囊体	水稻	42	［20］
albino embryo and seedling	白化	AES	基因转录	PPR蛋白，RNA剪接	拟南芥	-	［22］
rice albino 1	白化	RA1	蛋白翻译	氨酰tRNA合成酶，蛋白合成相关	水稻	-	［23］
albino leaf 1	白化	AL1	核糖体合成	八肽重复蛋白，调节核糖体RNA和核糖体蛋白的稳态	水稻	-	［24］
albino leaf 2	白化	AL2	RNA加工	叶绿体I型和II型内含子剪接	水稻	-	［25］
CN19M06	白化返绿	TSCA1	-	定位于2A染色体长臂6.5 Mb的区间内，有111个注释基因	小麦	15	［26］
tic100^cue8	返绿	TIC100	蛋白稳态	叶绿体TIC蛋白，参与叶绿体蛋白质输入，影响逆行信号	拟南芥	-	［27］
white green leaf 2	白化返绿	WGL2	核糖体合成	定位于叶绿体的核糖体蛋白	水稻	-	［28］
thermo-sensitive virescent 2	白化返绿	TSV2	蛋白翻译	氨酰tRNA合成酶	水稻	20 24 28	［29］
virescent 2	返绿	V2	核苷酸代谢	鸟苷酸激酶	水稻	20	［30］
cde4	白化返绿	CDE4	RNA加工	PPR蛋白，叶绿体RNA剪切	水稻	20	［31］
delayed virescence 4	返绿	DV4	-	定位于7D染色体短臂1.46 Mb的区间内，有3个注释基因	小麦	-	［32-33］
ch1	褪绿	CAO	色素代谢	叶绿素合成	拟南芥	-	［34］
yellow leaf color 1	褪绿	YL1	-	定位于7D染色体8.75 Mb的区间内，有110个注释基因	小麦	-	［35］
chlorophyll	褪绿	TaCHLI-7D	色素代谢	镁螯合酶I亚基，叶绿素合成	小麦	-	［36］
wsl3	条纹	WSL3	基因转录	PPR蛋白，PEP活性	水稻	25	［38］
wlp2s， wlp2w	条纹	WLP2	基因转录	PEP转录相关	水稻	32	［39］
wsl4	条纹	WSL4	RNA加工	PPR蛋白，参与II型内含子剪接	水稻	20	［40］
young seedling stripe1	条纹	YSS1	基因转录	DUF3727家族蛋白，参与调控PEP活性	水稻	20	［41］
wsl5	条纹	WSL5	RNA加工	PPR蛋白，RNA编辑和剪接	水稻	20	［42］
wsl6	条纹	WSL6	核糖体合成	GTP结合蛋白，核糖体合成	水稻	20	［43］
wsl9	条纹	WSL9	RNA加工	HNH内切酶，编辑效率	水稻	20	［44］
young leaf white stripe	条纹	YLWS	RNA加工	PPR蛋白，参与II型内含子剪接	水稻	20	［45］
B03S	斑马纹	-	色素代谢	未知基因，可能与玉米黄素合成有关	水稻	-	［46］
brittle culm and zebra leaf 1	斑马纹	BZ1	糖代谢	UDP-半乳糖/葡萄糖差向异构酶	水稻	-	［47］
zebra leaf 15	斑马纹	Z15	-	受体激酶	水稻	18	［48］
white kernel mutant 3735	斑马纹	ZmPTOX1	色素代谢	类胡萝卜素合成	玉米	-	［49-50］
zebra7	斑马纹	ZB7	色素代谢	MEP途径	玉米	32	［51］
thermo-sensitive chlorophyll-deficient mutant 1	白化	TCM1	基因转录	PEP转录相关	水稻	20	［53］
tcd12	白化	TCD12/ SIG2A	基因转录	RNA聚合酶Sigma因子	水稻	23	［54］
Dular	白化	DUA1	基因转录 RNA加工	PPR蛋白，与SIG1和WSP1互作分别介导叶绿体基因表达和RNA编辑	水稻	18	［55-56］
low-temperature-sensitive	白化	OsPUS1	RNA加工	叶绿体核糖体RNA的假尿苷化修饰	水稻	20	［57］
temperature-sensitive chlorophyll-deficient 11	白化	TSCD11	蛋白翻译	氨酰tRNA合成酶，蛋白合成相关	水稻	35	［59］
tcd9	白化	TCD9	蛋白折叠蛋白稳态	伴侣蛋白，质体分裂	水稻	20	［60］
T-DNA插入突变体oshsp70cp1	褪绿	HSP70CP1	蛋白折叠蛋白稳态	蛋白转运与组装	水稻	37	［61］
yellow variegated1	不均匀的褪绿	FtsH5/VAR1	蛋白稳态	FtsH蛋白酶，光系统稳定性	拟南芥	-	［62］
yellow variegated2	不均匀的褪绿	FtsH2/VAR2	蛋白稳态	FtsH蛋白酶，光系统稳定性	拟南芥	8	［62-63］
thermosensitive 36	褪绿	FtsH11	蛋白稳态	FtsH蛋白酶，光反应蛋白复合体稳定性	拟南芥	30	［64］
SALK_033047	褪绿	FtsH11	蛋白稳态	FtsH蛋白酶，光反应蛋白复合体稳定性	拟南芥	26	［65］
T-DNA插入突变体Salk_028162C	白化	AT3G06730	-	水稻硫氧还蛋白WP2同源蛋白	拟南芥	22	［66］
white panicle2	白化	WP2	RNA加工	硫氧还蛋白，叶绿体RNA编辑	水稻	35	［66］
tcd5	白化返绿	TCD5	基因转录	氧化还原蛋白，单加氧酶，叶绿体基因表达	水稻	20	［74］
temperature-sensitive virescent	白化返绿	TSV	基因转录	氧化还原蛋白，与TrxZ相互作用，PEP转录相关	水稻	20	［75］
white stripe leaf	条纹	WSL	基因转录	PPR蛋白，PEP基因转录	水稻	20	［76］

suppressor of ospus1-1 10	白化	SOP10	RNA加工	PPR蛋白，调控线粒体RNA剪接和编辑	水稻	22	［77］
virescent-1	返绿	V1/NUS1	RNA降解	细菌转录终止抑制子，叶绿体RNA代谢	水稻	20	［78］
RNAi株系	褪绿	CRP1	RNA降解	mRNA稳定性	水稻	4	［79］
umpk	褪绿	UMPK	核苷酸代谢	尿苷5′-单磷酸激酶	水稻	6	［80］
virescent 3	返绿	V3	核苷酸代谢	核糖核苷酸还原酶大亚基，叶绿体DNA合成	水稻	20	［81］
wsl12	条纹	WSL12	核苷酸代谢	核苷二磷酸激酶	水稻	22 34	［82］
stripe and drooping leaf	条纹	SDL/ RNRS1	核苷酸代谢	核糖核苷酸还原酶小亚基	水稻	20	［83］
temperature-sensitive chlorophyll-deficient 3	白化	TCD3	核苷酸代谢	假尿苷合酶，叶绿体RNA代谢，RNA修饰	水稻	20	［84］
pale-green leaf 12	褪绿	PGL12	核糖体合成	PPR蛋白，叶绿体16S rRNA加工	水稻	20	［85］
tcd11	白化	TCD11	核糖体合成	叶绿体核糖体亚基	水稻	20	［86］
white leaf and panicles 1	返绿	WLP1	核糖体合成	核糖体L13蛋白	水稻	23	［87］
tcm5	白化	TCM5	蛋白稳态	蛋白酶，光合系统II保护	水稻	32	［88］
RNAi株系	褪绿	NOA1	蛋白翻译核糖体合成	YqeH同源蛋白，叶绿体中核糖体组装和蛋白翻译	水稻	22	［89-91］
chlorophyll deficient 1	褪绿	CDE1/GluRS	蛋白翻译	tRNA合酶，蛋白合成	水稻	32	［92］
XN1376B	白化返绿	-	-	-	小麦	-	［93］
Zm-murE-2	白化	MurE	基因转录	PEP活性，叶绿体逆行信号	玉米	-	［94］
ppr5-1	白化	PPR5	基因转录	PPR蛋白，叶绿体逆行信号	玉米	-	［94］

hsp21	褪绿	HSP21	基因转录	PEP功能，与pTAC5互作	拟南芥	30	［95］
RNAi株系	褪绿	pTAC5	基因转录	质体转录活性染色质蛋白	拟南芥	30	［95］
cp29a cp31a	褪绿	CP29ACP31A	RNA降解	RNA结合蛋白，RNA稳定性和加工	拟南芥	8	［96］
thermo-sensitive mutant in leafcolor 1	褪绿	DG1/TSL1	RNA加工	PPR蛋白，叶绿体RNA编辑和剪切，MORF2，DYW1，DYW2互作	拟南芥	29	［97］
T-DNA插入突变体rare1-1，rare1-2	褪绿	RARE1	RNA加工	PPR蛋白，通过RNA编辑影响脂肪酸合成	拟南芥	28	［98］
thermo-sensitive mutant in leafcolor 2	褪绿	FtsHi5/TSL2	蛋白稳态	叶绿体蛋白输入复合体成员	拟南芥	16	［99］
ggps1-2， ggps1-3	褪绿	GGPS1	色素代谢	叶绿素合成途径	拟南芥	27	［100］
gun5-1	褪绿	GUN5	色素代谢	镁离子螯合酶，质核信号穿梭	拟南芥	4	［101］
paleface1	褪绿	PFC1	核糖体合成	rRNA腺嘌呤去甲基化酶，核糖体合成	拟南芥	5	［102］
rbd1-1	褪绿	RBD1	核糖体合成	RNA结合蛋白，23S rRNA加工	拟南芥	4	［103］
rps1	白化	RPS1	核糖体合成	叶绿体核糖体蛋白	拟南芥	38	［104］
rps5	褪绿	RPS5	核糖体合成	叶绿体核糖体功能	拟南芥	4	［105］
crass-1， crass-2	褪绿	CRASS	蛋白翻译	核糖体相关蛋白，叶绿体蛋白翻译效率	拟南芥	4	［106］
svr3-1	褪绿	SVR3	蛋白翻译	翻译延伸因子	拟南芥	8	［107］