基因可变剪接调控植物响应非生物胁迫研究进展

doi:10.11983/CBB24189

植物学报 ›› 2025, Vol. 60 ›› Issue (3): 435-448.DOI: 10.11983/CBB24189 cstr: 32102.14.CBB24189

基因可变剪接调控植物响应非生物胁迫研究进展

熊良林¹^,², 梁国鲁¹^,², 郭启高¹^,², 景丹龙¹^,²^,^*()

¹西南大学园艺园林学院, 长江上游农业生物安全与绿色生产教育部重点实验室, 重庆 400715
²西南大学含弘学院, 重庆 400715

收稿日期:2024-12-05 接受日期:2025-02-22 出版日期:2025-05-10 发布日期:2025-02-26
通讯作者: *景丹龙, 男, 博士, 副教授, 硕士生导师。主要从事果树资源评价、重要农艺性状和逆境响应的分子调控机制研究。近年来, 主持国家自然科学基金面上项目、国家重点研发计划项目子课题、国家自然科学青年基金、重庆市自然科学基金面上项目和中央高校基本业务费重点项目等项目; 指导国家级大学生创新创业训练计划项目2项。以第一作者和通讯作者在Plant Biotechnology Journal和Horticulture Research等国际学术期刊上发表SCI论文20余篇, 其中2篇论文入选ESI高被引论文。获授权国家发明专利15项。现任枇杷产业国家创新联盟副秘书长、重庆市科技特派员。荣获“梁希青年论文奖”二等奖(中国林学会)、园艺园林学院首届优秀青年教师。担任Forestry Research首届青年编委、《西北植物学报》和《南方农业学报》首届青年编委, Food Research International和Frontiers in Plant Science等国际期刊的审稿专家。E-mail: jingdanlong@swu.edu.cn
基金资助:
国家自然科学基金(32470386);国家自然科学基金(32102321);重庆市科技局项目(CSTB2022NSCQ-MSX1541);重庆市科技局项目(cstc2021jscx- gksbX0010);中央高校基本科研业务费-能力提升项目(SWU-KT22055)

Advances in the Regulation of Alternative Splicing of Genes in Plants in Response to Abiotic Stress

Xiong Lianglin¹^,², Liang Guolu¹^,², Guo Qigao¹^,², Jing Danlong¹^,²^,^*()

¹Key Laboratory of Agricultural Biosafety and Green Production in the Upper Reaches of the Yangtze River, Ministry of Education, College of Horticulture and Landscape Architecture of Southwest University, Chongqing 400715, China
²Hanhong College of Southwest University, Chongqing 400715, China

Received:2024-12-05 Accepted:2025-02-22 Online:2025-05-10 Published:2025-02-26
Contact: *E-mail: jingdanlong@swu.edu.cn

摘要/Abstract

摘要： 外界环境对植物生长发育产生至关重要的影响, 近年来频繁出现的极端气候严重威胁植物生长发育。明确植物抗逆调控机制对于保障植物生存和发育(特别是经济作物的产量)具有重要意义。基因可变剪接是一种重要的转录后调控机制, 对于植物基因功能的多样性与抗逆性均具有重要作用。目前已在不同植物中鉴定出多种抗逆相关基因可变剪接体, 并阐明部分基因可变剪接介导的植物抗逆调控机制, 有效奠定了植物抗逆研究的相关理论基础。因此, 挖掘和鉴定更多抗逆基因在非生物胁迫下的可变剪接调控机制对于植物抗逆研究具有重要意义。该文综述了植物基因可变剪接类型以及剪接机制, 重点阐述了非生物胁迫下相关基因可变剪接介导的植物抗逆研究进展, 展望了未来的研究方向。

关键词: 植物, 基因可变剪接, 调控机制, 非生物胁迫, 胁迫耐受性

Abstract: The external environment severely affects growth and development of plants. In recent years, the increasing extreme climates have posed a serious threat to the growth and development of plants. Understanding the regulatory mechanisms of plant stress tolerance is of great significance for ensuring the survival and development of plants (especially economic crops) and their yields. Alternative splicing is an important post-transcriptional regulatory mechanism and plays an important role in the diversity of plant gene functions and stress resistance. At present, a variety of alternative splicing variants of stress-resistant related genes have been identified in different plant species, and several regulatory mechanisms involved in alternative splicing have been elucidated, effectively advancing the relevant theoretical basis for plant stress resistance in plants. This paper reviews the types and splicing mechanisms of alternative splicing in plants, highlights the recent progress in alternative splicing regulation of plant responses to abiotic stress, and provides a prospect for the future direction of research on alternative splicing in plants.

Key words: plants, gene alternative splicing, regulatory mechanisms, abiotic stress, stress tolerance

熊良林, 梁国鲁, 郭启高, 景丹龙. 基因可变剪接调控植物响应非生物胁迫研究进展. 植物学报, 2025, 60(3): 435-448.

Xiong Lianglin, Liang Guolu, Guo Qigao, Jing Danlong. Advances in the Regulation of Alternative Splicing of Genes in Plants in Response to Abiotic Stress. Chinese Bulletin of Botany, 2025, 60(3): 435-448.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB24189

https://www.chinbullbotany.com/CN/Y2025/V60/I3/435

图/表 3

图1 非生物胁迫下可变剪接的主要类型(参考Laloum et al., 2018)

Figure 1 Main types of alternative splicing under abiotic stress (refer to Laloum et al., 2018)

图2 正常条件和不同程度热胁迫下, 富含丝氨酸/精氨酸(SR)蛋白质介导的前体mRNA调控机制(参考Ling et al., 2021) (A) 植物正常发育情况下进行的组成型剪接模式图(在正常条件下, 热响应基因(heat-responsive genes, HRG)未被激活或几乎不表达, 生长相关基因(genes responding for growth, GRG)正常表达。SR蛋白发生磷酸化(例如, 由剪接因子复合物(splicing factor complex, SFC)中的SR蛋白激酶(SR protein kinase, SRPK)进行磷酸化), 从而参与组成型剪接, 产生大量发育相关蛋白); (B) 植物在轻度热胁迫下可变剪接模式图(在轻度热胁迫(例如, 32°C持续6小时)下, 包括热诱导SR基因在内的HRGs在转录水平被激活, 磷酸化的SR蛋白参与HRGs的组成型剪接或可变剪接, 同时GRGs在转录水平上被抑制。部分来自GRGs的转录本片段也由磷酸化SR蛋白剪接, 产生不同功能的剪接变体。植物细胞通过该机制产生大量的热耐受蛋白进而响应热胁迫); (C) 植物在重度热胁迫下的可变剪接模式图(在重度热胁迫(例如, 45°C持续90分钟)下, 包括热诱导SR基因在内的HRGs在转录水平被激活。然而, 在重度热胁迫下SR蛋白去磷酸化(例如, 由SRPK缺乏引起), 导致功能性剪接因子短缺, 从而使HRGs和GRGs内含子保留的转录本在细胞核中积累, 最终导致功能性转录本数量显著减少。因此, 植物细胞中的热耐受蛋白和发育相关蛋白短缺)。ASF: 可变剪接片段; ×: 该过程被抑制。

Figure 2 The mechanism of precursor mRNA regulation mediated by serine/arginine-rich (SR) proteins under normal conditions and different degrees of heat stress (refer to Ling et al., 2021) (A) Schematic diagram of constitutive splicing in plants under normal development (under normal conditions, heat-responsive genes (HRG) are not activated or are barely expressed, while growth-related genes (GRGs) are actively transcribed. SR proteins are efficiently phosphorylated (e.g., phosphorylated by SR protein kinase (SRPK) in the splicing factor complex (SFC)), thus participating in constitutive splicing and generating many development-related proteins); (B) Schematic diagram of alternative splicing in plants under mild heat stress (under mild heat stress (e.g., 32°C for 6 hours), HRGs including heat-inducible SR genes are transcriptionally activated. Phosphorylated SR proteins participate in constitutive or alternative splicing of HRGs. GRGs are suppressed at the transcriptional level, but their pre-mRNAs may still be alternative spliced by phosphorylated SR proteins to produce functionally distinct isoforms. Through this mechanism, plant cells produce a large number of heat-tolerant proteins to cope with heat stress); (C) Schematic diagram of alternative splicing in plants under severe heat stress (under severe heat stress (e.g., 45°C for 90 minutes), HRGs including heat-inducible SR genes are activated at the transcriptional level. However, under severe heat stress, the dephosphorylation of SR proteins (e.g., caused by the deficiency of SRPK) leads to a deficiency of active splicing factors, resulting in the accumulation of intron-retained transcripts of HRGs and GRGs in the nucleus, and leads to an extremely low number of functional transcripts. Therefore, there is a shortage of heat-tolerant proteins and development-related proteins in plant cells). ASF: Alternative spliced fragment; ×: The process is inhibited.

表1 不同植物响应非生物胁迫过程中涉及可变剪接的相关基因

Table 1 Alternative splicing-related genes involved in regulating plant responses to abiotic stress in different species

物种	基因	功能	胁迫类型	参考文献
拟南芥(Arabidopsis thaliana)	AtHsfA2	维持热休克蛋白相关基因的表达水平	热胁迫	Charng et al., 2007
	AtSIZ1	促进热休克蛋白相关基因表达	热胁迫	Ishida et al., 2012
	AtIRE1	通过调节转录因子bZIP60前体mRNA剪接上调BIP3蛋白的表达	热胁迫	Ling et al., 2021
	AtZIFL1	产生剪接变体, 靶向叶片气孔保卫细胞的质膜并介导耐旱性	干旱胁迫	Remy et al., 2013
	AtSPCH	降低气孔密度, 减少水分蒸发	干旱胁迫	Wang et al., 2024a
	AtSR	使剪接因子丰度和活性发生动态变化, 影响靶基因表达	冷胁迫	Syed et al., 2012
	AtLUC7	参与冷胁迫条件下内含子的剪接调节	冷胁迫	de Francisco Amorim et al., 2018
	AtSR45	SR45.1亚型通过调节SOS基因表达以控制离子稳态和赋予耐盐性	盐胁迫	Albaqami et al., 2019
	AtSR34b	通过IRT1的剪接和稳定性促进IRT1蛋白积累	盐胁迫	Zhang et al., 2014
	AtSAD1	提高盐胁迫响应基因的剪接效率	盐胁迫	Cui et al., 2014
	AtSKIP	参与剪接位点的识别或切割	盐胁迫	Feng et al., 2015
	AtRCD1	RCD1的剪接变体可减少盐诱导的细胞死亡	盐胁迫	Hong et al., 2023
水稻(Oryza sativa)	OsIRE1	调节热胁迫响应基因OsbZIP74的前体mRNA剪接	热胁迫	Ling et al., 2021
小麦(Triticum aestivum)	TaHSFA6e	TaHSFA6e的剪接变体可增强下游热休克蛋白基因的转录活性	热胁迫	Wen et al., 2023
毛白杨(Populus tomentosa)	PtoRSZ21	通过调节PtoATG2b的剪接影响植物的水分利用效率和抗逆能力	干旱胁迫	Huang et al., 2024
玉米(Zea mays)	ZmPP2C26	ZmPP2C26的2个剪接变体负向调节干旱耐受性	干旱胁迫	Lu et al., 2022
茶树(Camellia sinensis)	CsWRKY21	CsWRKY21的剪接变体抑制ABA分解相关基因的表达	冷胁迫	Mi et al., 2024
谷子(Setaria italica)	SiCYP19	剪接变体可以提高脯氨酸含量和促进活性氧清除	盐胁迫	Zhang et al., 2024c
胡杨(Populus euphratica)	PeuHKT1;3	产生PeuHKT1;3a变体, 改变离子选择性	盐胁迫	Lv et al., 2024
大豆(Glycine max)	GmPeNTL9	剪接变体激活抗氧化清除系统	盐胁迫	Liu et al., 2024
	GmSnRK1.1	调节转录因子的活性和稳定性	盐胁迫	Liu et al., 2023

表1 不同植物响应非生物胁迫过程中涉及可变剪接的相关基因

Table 1 Alternative splicing-related genes involved in regulating plant responses to abiotic stress in different species

物种	基因	功能	胁迫类型	参考文献
拟南芥(Arabidopsis thaliana)	AtHsfA2	维持热休克蛋白相关基因的表达水平	热胁迫	Charng et al., 2007
	AtSIZ1	促进热休克蛋白相关基因表达	热胁迫	Ishida et al., 2012
	AtIRE1	通过调节转录因子bZIP60前体mRNA剪接上调BIP3蛋白的表达	热胁迫	Ling et al., 2021
	AtZIFL1	产生剪接变体, 靶向叶片气孔保卫细胞的质膜并介导耐旱性	干旱胁迫	Remy et al., 2013
	AtSPCH	降低气孔密度, 减少水分蒸发	干旱胁迫	Wang et al., 2024a
	AtSR	使剪接因子丰度和活性发生动态变化, 影响靶基因表达	冷胁迫	Syed et al., 2012
	AtLUC7	参与冷胁迫条件下内含子的剪接调节	冷胁迫	de Francisco Amorim et al., 2018
	AtSR45	SR45.1亚型通过调节SOS基因表达以控制离子稳态和赋予耐盐性	盐胁迫	Albaqami et al., 2019
	AtSR34b	通过IRT1的剪接和稳定性促进IRT1蛋白积累	盐胁迫	Zhang et al., 2014
	AtSAD1	提高盐胁迫响应基因的剪接效率	盐胁迫	Cui et al., 2014
	AtSKIP	参与剪接位点的识别或切割	盐胁迫	Feng et al., 2015
	AtRCD1	RCD1的剪接变体可减少盐诱导的细胞死亡	盐胁迫	Hong et al., 2023
水稻(Oryza sativa)	OsIRE1	调节热胁迫响应基因OsbZIP74的前体mRNA剪接	热胁迫	Ling et al., 2021
小麦(Triticum aestivum)	TaHSFA6e	TaHSFA6e的剪接变体可增强下游热休克蛋白基因的转录活性	热胁迫	Wen et al., 2023
毛白杨(Populus tomentosa)	PtoRSZ21	通过调节PtoATG2b的剪接影响植物的水分利用效率和抗逆能力	干旱胁迫	Huang et al., 2024
玉米(Zea mays)	ZmPP2C26	ZmPP2C26的2个剪接变体负向调节干旱耐受性	干旱胁迫	Lu et al., 2022
茶树(Camellia sinensis)	CsWRKY21	CsWRKY21的剪接变体抑制ABA分解相关基因的表达	冷胁迫	Mi et al., 2024
谷子(Setaria italica)	SiCYP19	剪接变体可以提高脯氨酸含量和促进活性氧清除	盐胁迫	Zhang et al., 2024c
胡杨(Populus euphratica)	PeuHKT1;3	产生PeuHKT1;3a变体, 改变离子选择性	盐胁迫	Lv et al., 2024
大豆(Glycine max)	GmPeNTL9	剪接变体激活抗氧化清除系统	盐胁迫	Liu et al., 2024
	GmSnRK1.1	调节转录因子的活性和稳定性	盐胁迫	Liu et al., 2023

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基因可变剪接调控植物响应非生物胁迫研究进展

Advances in the Regulation of Alternative Splicing of Genes in Plants in Response to Abiotic Stress

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