水稻叶色调控机制及相关基因研究进展

doi:10.11983/CBB23055

摘要/Abstract

摘要： 植物光合作用主要依赖绿色叶片, 而叶片生长发育最直观的特征是叶色。目前已克隆200多个调控水稻(Oryza sativa)叶色的基因。水稻叶色调控机制复杂多样, 涉及多条调控途径, 包括光合色素的生物合成与降解、核-质信号转导和血红素的合成。此外, 温度和光照强度等外部环境也会影响水稻叶色的变化。该文从分子机制、叶色相关基因及环境因素等方面总结了水稻叶色遗传调控机制, 并提出该领域亟待解决的科学问题, 以期为水稻高光效育种及应用提供理论支撑。

关键词: 水稻, 叶色, 调控机制

Abstract: Photosynthesis of plants depends on the green leaves, and the most intuitive feature of leaf growth and development is its color. At present, more than 200 genes related to rice leaf color have been cloned. The regulatory mechanisms of rice leaf color are complex and diverse, which involve multiple regulatory pathways, including biosynthesis and degradation of photosynthetic pigments, pathway of nucleus-plastid signaling, and heme synthesis. In addition, external environment such as temperature and light intensity can also affect the color changes of rice leaves. Here we reviewed the molecular pathways of rice leaf color, environmental factors and leaf color related genes, as well as the genetic regulatory mechanisms of rice leaf color was revealed, which will provide theoretical basis for rice high photosynthetic breeding and application, and future research in addressing some scientific problems in this field.

Key words: rice, leaf color, regulatory mechanism

戴若惠, 钱心妤, 孙静蕾, 芦涛, 贾绮玮, 陆天麒, 路梅, 饶玉春. 水稻叶色调控机制及相关基因研究进展. 植物学报, 2023, 58(5): 799-812.

Dai Ruohui, Qian Xinyu, Sun Jinglei, Lu Tao, Jia Qiwei, Lu Tianqi, Lu Mei, Rao Yuchun. Research Progress on the Mechanisms of Leaf Color Regulation and Related Genes in Rice. Chinese Bulletin of Botany, 2023, 58(5): 799-812.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB23055

https://www.chinbullbotany.com/CN/Y2023/V58/I5/799

图/表 4

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调控途径	基因	表型	染色体定位	基因功能	参考文献
叶绿素	LYL1	黄化	2	编码叶绿体信号识别颗粒	Li et al., 2019
合成	CHLD	黄化	3	镁离子螯合酶D亚基	Zhang et al., 2015
	CHLI	黄绿叶	3	镁离子螯合酶I亚基	Zhang et al., 2015
	CHLH	白化	3	镁离子螯合酶H亚基	Zhou et al., 2012
	HemA	淡绿	10	谷氨酰-tRNA还原酶基因	Zeng et al., 2020
	YLC2	黄叶	3	亚铁血红素加氧酶基因	Mahawar and Shekhawat, 2018
	FdC2	黄绿叶	3	编码铁氧还蛋白类似物, 调节电子转移和叶绿	Zhao et al., 2015
	FdC2	黄绿叶	3	素含量	Zhao et al., 2015
	PORA	黄化	4	原叶绿素酸酯氧化还原酶A基因	Liu et al., 2016
	PORB	淡绿	10	原叶绿素酸酯氧化还原酶B基因	Liu et al., 2016
	YGL1	黄绿叶	5	叶绿素合成酶基因	Liu et al., 2016
	YGL2	黄叶	6	编码血红素加氧酶	Chen et al., 2013
	CAO1	淡绿	10	叶绿素a加氧酶基因	Yang et al., 2015
	CAO2	敲除无差异	10	叶绿素a加氧酶基因	Yang et al., 2015
	YL1	黄绿叶	2	编码光诱导叶绿体蛋白	Wang et al., 2017b
	V5A	黄叶	9	与OsPORA和OsPORB互作, 调控叶绿素合成	Liu et al., 2016
叶绿素	NOL	永绿	3	叶绿素b还原酶基因	孟祥州等, 2015
降解	DVR	黄绿叶	3	编码联乙烯还原酶	Wang et al., 2013
	SGRL	持绿	4	参与叶绿素分解	Rong et al., 2013
	VAL1	白化转绿	8	嘌呤核苷酸生物合成酶基因	Zhang et al., 2018
	DYE1	持绿	8	编码捕光复合体I亚基	Yamatani et al., 2018
类胡萝	PDS	白化	3	编码ABA合成	Fang et al., 2019
卜素	IspE	黄化转绿	1	参与异戊二烯生物合成	Chen et al., 2018
合成	MIT1	持绿	12	编码胡萝卜素异构酶	Liu et al., 2021a
	ZEBRA2	斑马叶	11	类胡萝卜素异构酶基因	Chai et al., 2011
核-质信	WLP2	白叶和白穗	1	编码质体编码的RNA聚合酶相关蛋白	Lv et al., 2017
号转导	WSL	白条纹叶	1	编码PPR蛋白, 参与叶绿体RNA剪接	Wang et al., 2017b
相关	WSL4	温敏条纹	2	编码PPR蛋白, 参与叶绿体RNA剪接	Wang et al., 2017b
	WSL9	白叶条纹	3	编码含HNH核酸内切酶结构域的蛋白质	Zhu et al., 2020b
	CAF1	白化致死	1	编码叶绿体RNA剪接因子	Zhang et al., 2019
	SLA4	白化	7	编码PPR蛋白, 参与RNA剪接	Lv et al., 2020
	WSP1	白条纹叶和白穗	4	叶绿体基因RNA编辑因子, 编码MORF家族蛋白	Zhang et al., 2017
	PPR6	白化致死	5	编码PPR蛋白, 参与叶绿体RNA编辑与剪接	Tang et al., 2017
	DUA1	低温白化	9	编码PPR蛋白, 参与叶绿体RNA编辑	Yap et al., 2015
	Z3	斑马叶	3	编码柠檬酸转运蛋白	Kim et al., 2018
叶绿体	ASL1	白化致死	1	编码叶绿体核糖体蛋白	Song et al., 2014
发育相	ALS3	白化致死	1	编码叶绿体靶向的PPR蛋白, 含有10个串联	Lin et al., 2018
关	ALS3	白化致死	1	的PPR基序	Lin et al., 2018
	WP3	幼苗白条纹	1	编码靶向线粒体的蛋白质	Li et al., 2018
	WSL6	白条纹叶	5	编码GTP结合蛋白	Sun et al., 2019
	TSV3	温敏白化	3	编码GTP结合蛋白	Lin et al., 2018
	WSL8	白条纹叶	5	脱氧核糖核酸激酶基因	Liu et al., 2020
	V1	温敏白化转绿	3	编码叶绿体定位蛋白NUS1, 参与叶绿体RNA	Kusumi et al., 2011
	V1	温敏白化转绿	3	代谢调节	Kusumi et al., 2011
	V3	白化转绿	6	编码核糖核酸还原酶大亚基	Inagaki et al., 2015
	V4	低温白化	7	编码PPR蛋白	Lv et al., 2020
	TCD5	温敏白化	5	单加氧酶基因	Wang et al., 2016a
	GARS	白条纹叶	8	编码一种催化嘌呤核苷酸生物合成第2步的酶	Cao et al., 2018
	YGL8	黄叶	1	编码尿嘧啶核苷酸激酶	Kong et al., 2016
	Rey(k2)	黄化转绿	10	磷酸果糖激酶B型碳水化合物激酶基因	Zhu et al., 2020a

调控途径	基因	表型	染色体定位	基因功能	参考文献
叶绿素	LYL1	黄化	2	编码叶绿体信号识别颗粒	Li et al., 2019
合成	CHLD	黄化	3	镁离子螯合酶D亚基	Zhang et al., 2015
	CHLI	黄绿叶	3	镁离子螯合酶I亚基	Zhang et al., 2015
	CHLH	白化	3	镁离子螯合酶H亚基	Zhou et al., 2012
	HemA	淡绿	10	谷氨酰-tRNA还原酶基因	Zeng et al., 2020
	YLC2	黄叶	3	亚铁血红素加氧酶基因	Mahawar and Shekhawat, 2018
	FdC2	黄绿叶	3	编码铁氧还蛋白类似物, 调节电子转移和叶绿	Zhao et al., 2015
	FdC2	黄绿叶	3	素含量	Zhao et al., 2015
	PORA	黄化	4	原叶绿素酸酯氧化还原酶A基因	Liu et al., 2016
	PORB	淡绿	10	原叶绿素酸酯氧化还原酶B基因	Liu et al., 2016
	YGL1	黄绿叶	5	叶绿素合成酶基因	Liu et al., 2016
	YGL2	黄叶	6	编码血红素加氧酶	Chen et al., 2013
	CAO1	淡绿	10	叶绿素a加氧酶基因	Yang et al., 2015
	CAO2	敲除无差异	10	叶绿素a加氧酶基因	Yang et al., 2015
	YL1	黄绿叶	2	编码光诱导叶绿体蛋白	Wang et al., 2017b
	V5A	黄叶	9	与OsPORA和OsPORB互作, 调控叶绿素合成	Liu et al., 2016
叶绿素	NOL	永绿	3	叶绿素b还原酶基因	孟祥州等, 2015
降解	DVR	黄绿叶	3	编码联乙烯还原酶	Wang et al., 2013
	SGRL	持绿	4	参与叶绿素分解	Rong et al., 2013
	VAL1	白化转绿	8	嘌呤核苷酸生物合成酶基因	Zhang et al., 2018
	DYE1	持绿	8	编码捕光复合体I亚基	Yamatani et al., 2018
类胡萝	PDS	白化	3	编码ABA合成	Fang et al., 2019
卜素	IspE	黄化转绿	1	参与异戊二烯生物合成	Chen et al., 2018
合成	MIT1	持绿	12	编码胡萝卜素异构酶	Liu et al., 2021a
	ZEBRA2	斑马叶	11	类胡萝卜素异构酶基因	Chai et al., 2011
核-质信	WLP2	白叶和白穗	1	编码质体编码的RNA聚合酶相关蛋白	Lv et al., 2017
号转导	WSL	白条纹叶	1	编码PPR蛋白, 参与叶绿体RNA剪接	Wang et al., 2017b
相关	WSL4	温敏条纹	2	编码PPR蛋白, 参与叶绿体RNA剪接	Wang et al., 2017b
	WSL9	白叶条纹	3	编码含HNH核酸内切酶结构域的蛋白质	Zhu et al., 2020b
	CAF1	白化致死	1	编码叶绿体RNA剪接因子	Zhang et al., 2019
	SLA4	白化	7	编码PPR蛋白, 参与RNA剪接	Lv et al., 2020
	WSP1	白条纹叶和白穗	4	叶绿体基因RNA编辑因子, 编码MORF家族蛋白	Zhang et al., 2017
	PPR6	白化致死	5	编码PPR蛋白, 参与叶绿体RNA编辑与剪接	Tang et al., 2017
	DUA1	低温白化	9	编码PPR蛋白, 参与叶绿体RNA编辑	Yap et al., 2015
	Z3	斑马叶	3	编码柠檬酸转运蛋白	Kim et al., 2018
叶绿体	ASL1	白化致死	1	编码叶绿体核糖体蛋白	Song et al., 2014
发育相	ALS3	白化致死	1	编码叶绿体靶向的PPR蛋白, 含有10个串联	Lin et al., 2018
关	ALS3	白化致死	1	的PPR基序	Lin et al., 2018
	WP3	幼苗白条纹	1	编码靶向线粒体的蛋白质	Li et al., 2018
	WSL6	白条纹叶	5	编码GTP结合蛋白	Sun et al., 2019
	TSV3	温敏白化	3	编码GTP结合蛋白	Lin et al., 2018
	WSL8	白条纹叶	5	脱氧核糖核酸激酶基因	Liu et al., 2020
	V1	温敏白化转绿	3	编码叶绿体定位蛋白NUS1, 参与叶绿体RNA	Kusumi et al., 2011
	V1	温敏白化转绿	3	代谢调节	Kusumi et al., 2011
	V3	白化转绿	6	编码核糖核酸还原酶大亚基	Inagaki et al., 2015
	V4	低温白化	7	编码PPR蛋白	Lv et al., 2020
	TCD5	温敏白化	5	单加氧酶基因	Wang et al., 2016a
	GARS	白条纹叶	8	编码一种催化嘌呤核苷酸生物合成第2步的酶	Cao et al., 2018
	YGL8	黄叶	1	编码尿嘧啶核苷酸激酶	Kong et al., 2016
	Rey(k2)	黄化转绿	10	磷酸果糖激酶B型碳水化合物激酶基因	Zhu et al., 2020a