植物细胞中RNA结合蛋白相分离现象研究进展

doi:10.16819/j.1001-7216.2026.241116

摘要/Abstract

摘要：

RNA结合蛋白(RBP)作为关键的RNA代谢调控因子，在真核生物的细胞发育中参与多种RNA加工事件。这些RBP能够与RNA结合形成核糖核蛋白复合物，并可通过相分离机制形成动态的无膜凝集体。本文首先综述了RBP和相分离的基本概念，进而深入解析了植物细胞中RBPs相分离的分子机制，重点阐明其在植物生长发育和对环境变化的响应中的重要作用。最后，展望了RBP相分离机制研究中面临的挑战及其在作物改良中的潜在应用价值。

关键词: RNA结合蛋白, 相分离, 凝集体, 发育, 抗逆性

Abstract:

RNA-binding proteins (RBPs) are key regulators of RNA metabolism and play crucial roles in various RNA processing events during cellular development in eukaryotes. They interact with RNA to form ribonucleoprotein complexes and can undergo phase separation, leading to the formation of dynamic, membrane-less condensates. This review begins by outlining the fundamental concepts of RBPs and phase separation. It then provides an in-depth analysis of the molecular mechanisms governing RBP-mediated phase separation in plant cells, highlighting its critical functions in plant growth, development, and responses to environmental stresses. Finally, the review discusses future challenges in researching RBP phase separation and its potential applications for crop improvement.

Key words: RNA-binding protein, phase separation, condensate, development, stress resilience

周华成, 包秀浩, 王晓乐, 鲁镇飞, 马荣荣, 陆永法, 王晓燕, 蔡克锋, 唐志明, 周卓奇, 陈志新, 马炜炜. 植物细胞中RNA结合蛋白相分离现象研究进展[J]. 中国水稻科学, 2026, 40(1): 27-36.

ZHOU Huacheng, BAO Xiuhao, WANG Xiaole, LU Zhenfei, MA Rongrong, LU Yongfa, WANG Xiaoyan, CAI Kefeng, TANG Zhiming, ZHOU Zhuoqi, CHEN Zhixin, MA Weiwei. Research Progress in the Phase Separation of RNA-binding Proteins in Plant Cells[J]. Chinese Journal OF Rice Science, 2026, 40(1): 27-36.

图/表 2

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名称 Name	结构特征 Structural characteristics	生理功能 Physiological function
SR蛋白 Serine/Arginine-rich protein	N端：含RRM结构域 N-terminus: containing an RRM domain C端：富含SR的结构域 C-terminus: containing an SR-rich domain	调控前体mRNA的剪接、转运及翻译，参与植物生长发育调控(如细胞增殖、开花时间)，增强高温、低温等非生物胁迫适应性 Regulates precursor mRNA splicing, transport, and translation; participates in plant growth and development regulation (e.g., cell proliferation, flowering time); enhances adaptation to abiotic stresses such as high and low temperatures
GR-RBP Glycine-rich RNA-binding protein	N端：含RRM或CSD结构域 N-terminus: containing RRM or CSD domain C端：存在富含甘氨酸区域(部分该类型蛋白的该区域间隔分布着CCHC型锌指基序) C-terminus: containing glycine-rich region(in some proteins, this region is interspersed with conserved CCHC-type zinc finger motifs)	促进RNA折叠，响应干旱、盐、低温胁迫；通过激素信号调控植物开花、生物钟及种子萌发等发育过程 Promotes RNA folding; responds to drought, salt, and low-temperature stresses; regulates flowering, circadian rhythm, and seed germination through hormone signaling
PPR蛋白 Pentatricopeptide repeat protein	P类：串联重复的35个氨基酸基序 P-type: tandem repeats of 35-amino acid motifs PLS类：由P基序(35 aa)、L基序(35~36 aa)和S基序 (31~32 aa)组成 PLS-type: composed of P motifs (35 aa), L motifs (35-36 aa), and S motifs (31-32 aa)	靶向线粒体和叶绿体，调控RNA编辑、剪接及稳定性，影响光合作用、呼吸作用、胚胎发育，并参与胁迫响应 Regulates mitochondria and chloroplasts; controls RNA editing, splicing, and stability; influences photosynthesis, respiration, embryogenesis, and participates in stress responses
DEAD-box RH DEAD-box RNA helicase	包含保守的DEAD基序及Q、I、III等结构域，有ATP 依赖的RNA解旋活性 Containing conserved DEAD motifs (Q, I, II, III, V, VI) with ATP-dependent RNA helicase activity	催化RNA二级结构解旋，参与核与细胞器RNA加工(如内含子剪接、rRNA成熟)，调节生长发育及低温、干旱胁迫耐受性 Catalyze RNA secondary structure unwinding; participate in post-transcriptional RNA processing (e.g., intron splicing, mRNA maturation); regulate growth and development, as well as tolerance to cold and drought stress
RNA分子伴侣 RNA molecular chaperone	结构多样，部分属于GR-RBP或DEAD-box家族，具有非特异性RNA结合能力 Structurally diverse; some belong to GR-RBP or DEAD- box families; exhibit nonspecific RNA-binding ability	通过纠正RNA错误折叠维持其功能，在低温等胁迫下保护RNA稳定性，促进叶绿体RNA代谢，提高植物逆境适应能力 Maintaining RNA function by correcting misfolding, protecting RNA stability under stresses such as low temperature, promoting chloroplast RNA metabolism, and enhancing plant stress adaptability
YTH蛋白 YT521-B homology domain-containing protein	含有保守的YTH结构域(约140个氨基酸)，与RRM具有结构上的相似性。能形成芳香族笼结构，与RNA上的 m⁶A甲基化修饰结合 Containing conserved YTH domain (approximately140 amino acids); specifically binding to m⁶A methylation on RNA	通过识别表观转录修饰的mRNA，调控RNA代谢，在生长发育(如叶片建成、根系发育、开花时间调控)及非生物胁迫响应(高温、盐胁迫、干旱等)中发挥关键作用 By recognizing epitranscriptomic modifications of mRNA, regulate RNA metabolism, maintain gene expression stability and homeostasis, and play key roles in plant adaptation under different environmental (heat, salt, drought, etc.) and biotic stress conditions

名称 Name	结构特征 Structural characteristics	生理功能 Physiological function
SR蛋白 Serine/Arginine-rich protein	N端：含RRM结构域 N-terminus: containing an RRM domain C端：富含SR的结构域 C-terminus: containing an SR-rich domain	调控前体mRNA的剪接、转运及翻译，参与植物生长发育调控(如细胞增殖、开花时间)，增强高温、低温等非生物胁迫适应性 Regulates precursor mRNA splicing, transport, and translation; participates in plant growth and development regulation (e.g., cell proliferation, flowering time); enhances adaptation to abiotic stresses such as high and low temperatures
GR-RBP Glycine-rich RNA-binding protein	N端：含RRM或CSD结构域 N-terminus: containing RRM or CSD domain C端：存在富含甘氨酸区域(部分该类型蛋白的该区域间隔分布着CCHC型锌指基序) C-terminus: containing glycine-rich region(in some proteins, this region is interspersed with conserved CCHC-type zinc finger motifs)	促进RNA折叠，响应干旱、盐、低温胁迫；通过激素信号调控植物开花、生物钟及种子萌发等发育过程 Promotes RNA folding; responds to drought, salt, and low-temperature stresses; regulates flowering, circadian rhythm, and seed germination through hormone signaling
PPR蛋白 Pentatricopeptide repeat protein	P类：串联重复的35个氨基酸基序 P-type: tandem repeats of 35-amino acid motifs PLS类：由P基序(35 aa)、L基序(35~36 aa)和S基序 (31~32 aa)组成 PLS-type: composed of P motifs (35 aa), L motifs (35-36 aa), and S motifs (31-32 aa)	靶向线粒体和叶绿体，调控RNA编辑、剪接及稳定性，影响光合作用、呼吸作用、胚胎发育，并参与胁迫响应 Regulates mitochondria and chloroplasts; controls RNA editing, splicing, and stability; influences photosynthesis, respiration, embryogenesis, and participates in stress responses
DEAD-box RH DEAD-box RNA helicase	包含保守的DEAD基序及Q、I、III等结构域，有ATP 依赖的RNA解旋活性 Containing conserved DEAD motifs (Q, I, II, III, V, VI) with ATP-dependent RNA helicase activity	催化RNA二级结构解旋，参与核与细胞器RNA加工(如内含子剪接、rRNA成熟)，调节生长发育及低温、干旱胁迫耐受性 Catalyze RNA secondary structure unwinding; participate in post-transcriptional RNA processing (e.g., intron splicing, mRNA maturation); regulate growth and development, as well as tolerance to cold and drought stress
RNA分子伴侣 RNA molecular chaperone	结构多样，部分属于GR-RBP或DEAD-box家族，具有非特异性RNA结合能力 Structurally diverse; some belong to GR-RBP or DEAD- box families; exhibit nonspecific RNA-binding ability	通过纠正RNA错误折叠维持其功能，在低温等胁迫下保护RNA稳定性，促进叶绿体RNA代谢，提高植物逆境适应能力 Maintaining RNA function by correcting misfolding, protecting RNA stability under stresses such as low temperature, promoting chloroplast RNA metabolism, and enhancing plant stress adaptability
YTH蛋白 YT521-B homology domain-containing protein	含有保守的YTH结构域(约140个氨基酸)，与RRM具有结构上的相似性。能形成芳香族笼结构，与RNA上的 m⁶A甲基化修饰结合 Containing conserved YTH domain (approximately140 amino acids); specifically binding to m⁶A methylation on RNA	通过识别表观转录修饰的mRNA，调控RNA代谢，在生长发育(如叶片建成、根系发育、开花时间调控)及非生物胁迫响应(高温、盐胁迫、干旱等)中发挥关键作用 By recognizing epitranscriptomic modifications of mRNA, regulate RNA metabolism, maintain gene expression stability and homeostasis, and play key roles in plant adaptation under different environmental (heat, salt, drought, etc.) and biotic stress conditions

生物学过程 Biological process	名称 Name	基因号 Gene ID	RNA结合域 RBD	RNA代谢类型 RNA metabolism type	物种 Species	参考文献 Reference
开花时间 Flowering time	SR45	AT1G16610	RRM	RNA剪接 RNA splicing	拟南芥Arabidopsis	[33]
	HRLP	AT2G44710	RRM	RNA剪接 RNA splicing	拟南芥Arabidopsis	[33]
	UBA2C	AT3G15010	RRM	转录 Transcription	拟南芥Arabidopsis	[34]
	FCA	AT4G16280	RRM	聚腺苷酸化Polyadenylation	拟南芥Arabidopsis	[35]
	SSF	AT2G47310	RRM	转录 Transcription	拟南芥Arabidopsis	[36]
	CPSF30L	AT1G30460	YTH	聚腺苷酸化Polyadenylation	拟南芥Arabidopsis	[37]
	EHD6	OS02G0517531	RRM	mRNA隔离 mRNA sequestration	水稻Rice	[38]
	YTH07	OS04G0608800	YTH	mRNA隔离 mRNA sequestration	水稻Rice	[38]
叶片发育 Leaf development	SERRATE	AT2G27100	ZnF C2H2	miRNA加工 miRNA processing	拟南芥Arabidopsis	[39]
芳香物质形成 Aroma formation	SlYTH2	SOLYC01G028860	YTH	转录抑制 Transcriptional repression	番茄Tomato	[40]
高温胁迫应答 Heat stress response	RBGD2/4	AT2G33410/ AT2G33410	RRM	mRNA转运 mRNA transport	拟南芥Arabidopsis	[41]
	AGO1	AT1G48410	PAZ PIWI	miRNA的保护 miRNA protection	拟南芥Arabidopsis	[42]
	GRP7	AT2G21660	RRM	转录Transcription	拟南芥Arabidopsis	[43]
渗透压胁迫应答Osmotic stress response	DCP5	AT1G26110	nil	mRNA隔离 mRNA sequestration	拟南芥Arabidopsis	[44]
盐胁迫应答 Salt stress response	ECT8	AT1G79270	YTH	mRNA降解 mRNA degradation	拟南芥Arabidopsis	[45]
干旱胁迫应答 Drought stress response	DRG9	OS09G0421700	ZnF	mRNA隔离 mRNA sequestration	水稻Rice	[46]
干旱胁迫应答 Drought stress response	SiYTH1	Seita.9G534000	YTH	转录Transcription	小米Foxtail millet	[47]
植物免疫应答 Plant immune response	RH6/8/12/20	AT2G45810/ AT3G61240/ AT4G00660/ AT1G55150	DEAD RRM	miRNA加工 miRNA processing	拟南芥Arabidopsis	[48,49]
植物免疫应答 Plant immune response	ECT1	AT3G03950	YTH	mRNA隔离 mRNA sequestration	拟南芥Arabidopsis	[50]

生物学过程 Biological process	名称 Name	基因号 Gene ID	RNA结合域 RBD	RNA代谢类型 RNA metabolism type	物种 Species	参考文献 Reference
开花时间 Flowering time	SR45	AT1G16610	RRM	RNA剪接 RNA splicing	拟南芥Arabidopsis	[33]
	HRLP	AT2G44710	RRM	RNA剪接 RNA splicing	拟南芥Arabidopsis	[33]
	UBA2C	AT3G15010	RRM	转录 Transcription	拟南芥Arabidopsis	[34]
	FCA	AT4G16280	RRM	聚腺苷酸化Polyadenylation	拟南芥Arabidopsis	[35]
	SSF	AT2G47310	RRM	转录 Transcription	拟南芥Arabidopsis	[36]
	CPSF30L	AT1G30460	YTH	聚腺苷酸化Polyadenylation	拟南芥Arabidopsis	[37]
	EHD6	OS02G0517531	RRM	mRNA隔离 mRNA sequestration	水稻Rice	[38]
	YTH07	OS04G0608800	YTH	mRNA隔离 mRNA sequestration	水稻Rice	[38]
叶片发育 Leaf development	SERRATE	AT2G27100	ZnF C2H2	miRNA加工 miRNA processing	拟南芥Arabidopsis	[39]
芳香物质形成 Aroma formation	SlYTH2	SOLYC01G028860	YTH	转录抑制 Transcriptional repression	番茄Tomato	[40]
高温胁迫应答 Heat stress response	RBGD2/4	AT2G33410/ AT2G33410	RRM	mRNA转运 mRNA transport	拟南芥Arabidopsis	[41]
	AGO1	AT1G48410	PAZ PIWI	miRNA的保护 miRNA protection	拟南芥Arabidopsis	[42]
	GRP7	AT2G21660	RRM	转录Transcription	拟南芥Arabidopsis	[43]
渗透压胁迫应答Osmotic stress response	DCP5	AT1G26110	nil	mRNA隔离 mRNA sequestration	拟南芥Arabidopsis	[44]
盐胁迫应答 Salt stress response	ECT8	AT1G79270	YTH	mRNA降解 mRNA degradation	拟南芥Arabidopsis	[45]
干旱胁迫应答 Drought stress response	DRG9	OS09G0421700	ZnF	mRNA隔离 mRNA sequestration	水稻Rice	[46]
干旱胁迫应答 Drought stress response	SiYTH1	Seita.9G534000	YTH	转录Transcription	小米Foxtail millet	[47]
植物免疫应答 Plant immune response	RH6/8/12/20	AT2G45810/ AT3G61240/ AT4G00660/ AT1G55150	DEAD RRM	miRNA加工 miRNA processing	拟南芥Arabidopsis	[48,49]
植物免疫应答 Plant immune response	ECT1	AT3G03950	YTH	mRNA隔离 mRNA sequestration	拟南芥Arabidopsis	[50]