Pre-mRNA选择性剪接对植物盐碱胁迫响应的调控机制

doi:10.11686/cyxb2025447

草业学报 ›› 2026, Vol. 35 ›› Issue (9): 236-246.DOI: 10.11686/cyxb2025447

• 综合评述 • 上一篇

Pre-mRNA选择性剪接对植物盐碱胁迫响应的调控机制

梁雪枫¹(), 方淑梅²^,³, 梁喜龙¹^,³()

^1.黑龙江八一农垦大学农学院，黑龙江大庆 163319
^2.黑龙江八一农垦大学生命科学技术学院，黑龙江大庆 163319
^3.黑龙江省植物生长调节剂工程技术研究中心，黑龙江大庆 163319

收稿日期:2025-11-04 修回日期:2025-12-15 出版日期:2026-09-20 发布日期:2026-07-27
通讯作者: 梁喜龙
作者简介:Corresponding author. E-mail： xilongliang@126.com
梁雪枫（1997-），女，黑龙江七台河人，在读博士。E-mail： liangxf1230@163.com
基金资助:
国家自然科学基金面上项目(32472238);国家自然科学基金面上项目(32272225);黑龙江省自然科学基金(LH2023C079)

Regulatory mechanisms of pre-mRNA alternative splicing in plant responses to saline-alkali stress

Xue-feng LIANG¹(), Shu-mei FANG²^,³, Xi-long LIANG¹^,³()

^1.College of Agriculture，Heilongjiang Bayi Agricultural University，Daqing 163319，China
^2.College of Life Science and Technology，Heilongjiang Bayi Agricultural University，Daqing 163319，China
^3.Heilongjiang Provincial Engineering and Technology Research Center for Plant Growth Regulators，Daqing 163319，China

Received:2025-11-04 Revised:2025-12-15 Online:2026-09-20 Published:2026-07-27
Contact: Xi-long LIANG

摘要/Abstract

摘要：

选择性剪接（AS）作为真核生物转录后调控的关键机制，通过对pre-mRNA进行差异化剪接，显著扩展蛋白质组多样性，在植物生长发育及逆境响应中发挥重要作用。本研究系统阐述了AS在植物盐碱胁迫响应中的调控作用与功能机制的研究进展。AS通过调控盐超敏感（SOS）、活性氧（ROS）及脱落酸（ABA）信号通路等，参与离子稳态维持、活性氧平衡、渗透调节等生理过程。探讨了AS在植物盐碱胁迫记忆的形成与跨代传递过程中的潜在作用。未来可进一步通过RNA-seq、RT-PCR、基因编辑及生物信息学等技术，探究AS在调控植物响应非生物胁迫方面的机制，进一步揭示AS在植物耐盐碱机制中的核心作用，从而为作物抗逆遗传改良提供新的理论依据与育种策略。

关键词: 选择性剪接, pre-mRNA, 盐碱胁迫, 植物响应, 分子调控

Abstract:

Alternative splicing （AS） is a critical post-transcriptional regulatory mechanism in eukaryotes， and significantly expands protein diversity through differential modification of pre-mRNAs， playing a key role in plant growth and development as well as stress responses. This study systematically reviews the research progress in defining the regulatory function and operational mechanisms of AS in plant responses to saline-alkali stress. It analyzes how AS participates in physiological processes such as ion homeostasis maintenance， reactive oxygen species （ROS） balance， and osmotic regulation by modulating signaling pathways including the salt overly sensitive （SOS）， ROS， and abscisic acid （ABA） pathway. This study investigated the potential role of AS in the formation and transgenerational transmission of saline-alkali stress memory in plants. Furthermore， AS reshapes splicing patterns of stress-related genes， thereby influencing the establishment and transgenerational transmission of stress memory in plants. Future research employing RNA-seq， RT-PCR， gene editing， and bioinformatics technologies can further explore the mechanisms by which the AS regulates plant responses to abiotic stress. This will reveal the key roles of AS in plant salt stress resistance mechanisms， providing new theoretical foundations and breeding strategies for genetic improvement of crop stress tolerance.

Key words: alternative splicing, pre-mRNA, saline-alkali stress, plant response, molecular regulation

梁雪枫, 方淑梅, 梁喜龙. Pre-mRNA选择性剪接对植物盐碱胁迫响应的调控机制[J]. 草业学报, 2026, 35(9): 236-246.

Xue-feng LIANG, Shu-mei FANG, Xi-long LIANG. Regulatory mechanisms of pre-mRNA alternative splicing in plant responses to saline-alkali stress[J]. Acta Prataculturae Sinica, 2026, 35(9): 236-246.

图/表 2

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物种 Species	基因 Gene	剪接变体/作用形式 Splicing isoform/form of action	功能 Function	参考文献 Reference
拟南芥 A. thaliana	AtSR45	产生SR45.1变体Produces SR45.1 isoform	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。The SR45.1 isoform controls ion homeostasis and confers salt tolerance by regulating salt overly sensitive （SOS） gene expression.	［44］
	AtSR45a	产生两种变体SR45a-1a和SR45a-1b Produces SR45a-1a and SR45a-1b isoform	剪接因子SR45a的两种变体都可以与CBP20共同作用，调控拟南芥多种盐胁迫核心因子核糖核酸加工过程。Two isoforms of the splicing factor SR45a can act synergistically with CBP20 to regulate the RNA processing of multiple core salt stress factors in A.thaliana.	［27］
	AtSR34b	调控靶基因剪接与mRNA稳定性Regulates splicing and mRNA stability of target genes	通过IRT1的剪接和稳定性促进IRT1蛋白积累Promotes IRT1 protein accumulation via regulating the splicing and stability of IRT1 mRNA	［45］
	AtSAD1	调控剪接效率Regulates splicing efficiency	提高盐胁迫响应基因的剪接效率Improves the splicing efficiency of salt stress response genes	［46］
	AtSKIP	参与剪接体组装Involves in the assembly of the isoform	调控全基因组范围内大量胁迫响应基因的选择性剪接Regulation of alternative splicing in numerous stress response gene across the entire genome	［47］
	AtSRAS1	产生SRAS1.1和SRAS1.2剪接变体Produces SRAS1.1 and SRAS1.2 isoforms	调控COP9信号复合体亚基5A（CSN5A）的蛋白平衡Regulates the protein balance of COP9 signalosome subunit 5A（CSN5A）	［48］
	AtRCD1	产生RCD1.1和RCD1.2剪接变体Produces RCD1.1 and RCD1.2 isoforms	RCD1的剪接变体可减少盐诱导的细胞死亡Splicing isoform of RCD1 reduce salt-induced cell death	［26］
水稻 O. sativa	OsNHX1	Pre-mRNA剪接产生3种变体Pre-mRNA splicing generates three isoforms	过表达异构体的品系耐盐性增强Lines overexpressing the isoform exhibit enhanced salt tolerance	［24］
	OsMAPK5	产生维持激酶活性的OsMAPK5a亚型Produces the OsMAPK5a isoform with retained kinase activity	OsMAPK5a可以增强对盐分胁迫的耐受性OsMAPK5a enhances tolerance to salt stress	［49］
	OsIM1	产生两种功能分化的剪接变体Produces two functionally differentiated isoforms	通过完整功能域维持叶绿体能量代谢稳态，又在胁迫持续时通过变体的降解或反馈调节实现能量分配优化。Maintains chloroplast energy metabolism homeostasis via intact domains； Optimizes energy allocation through isoform degradation or feedback regulation under prolonged stress.	［8］
大麦 H. vulgare	HvDRF1	产生两种含AP2结构域的功能变体Produces two functional isoforms containing the AP2 domain	两种剪接变体均通过AP2结构域发挥转录激活作用，响应盐碱胁迫的脱落酸核心信号通路。Both splicing isoforms exert transcriptional activation through the AP2 domain， responding to the core hormone signal pathway of abscisic acid in response to saline-alkali stress.	［15］
谷子 S. italica	SiCYP19	产生剪接变体Produces splicing isoform	剪接变体可以提高脯氨酸含量和促进活性氧清除能力。Splicing isoforms can increase proline content and promote reactive oxygen species （ROS） scavenging capacity.	［25］
胡杨 P. euphratica	PeuHKT1；3	产生PeuHKT1；3的可变3′剪接位点Generation of variable 3′splice sites for PeuHKT1；3	通过截短变体的降解与反馈调节实现能量分配，优化重塑离子选择性。Optimizes energy allocation and reshaping ion selectivity through degradation and feedback regulation of truncated isoforms.	［17］
大豆 G. max	GmPeNTL9	产生剪接变体Produces splicing isoform	剪接变体激活抗氧化清除系统Splicing isoform activates the antioxidant scavenging system	［18］
	GsSCL30a	GsSCL30a剪接因子对自身进行剪接The splicing factor GsSCL30a undergoes self-splices	GsSCL30a与GsSnRK1协同作用可增强植株的耐碱性GsSCL30a synergistically enhances plant alkaline tolerance with GsSnRK1	［19］
	GmAP2	GmAP2基因中微外显子的缺失Microexon deletion in the GmAP2 gene	AP2基因中微外显子的缺失增强了对盐碱的抗性Deletion of a microexon in the AP2 gene enhances resistance to saline-alkali stress	［20］
百慕大草Cynodon dactylon	CdDHN4	产生CdDHN4-L和CdDHN4-S Produces CdDHN4-L and CdDHN4-S	显著降低电解质渗漏和增强光合能力Significantly reduces electrolyte leakage and enhances photosynthetic capacity	［51］
盐芥T. salsuginea	TsHKT1	选择性剪接产生多种变体Alternative splicing generates multiple isoforms	形成互作复合体调控Na⁺/K⁺平衡Forms an interaction complex to regulate Na⁺/K⁺ balance	［16］

物种 Species	基因 Gene	剪接变体/作用形式 Splicing isoform/form of action	功能 Function	参考文献 Reference
拟南芥 A. thaliana	AtSR45	产生SR45.1变体Produces SR45.1 isoform	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。The SR45.1 isoform controls ion homeostasis and confers salt tolerance by regulating salt overly sensitive （SOS） gene expression.	［44］
	AtSR45a	产生两种变体SR45a-1a和SR45a-1b Produces SR45a-1a and SR45a-1b isoform	剪接因子SR45a的两种变体都可以与CBP20共同作用，调控拟南芥多种盐胁迫核心因子核糖核酸加工过程。Two isoforms of the splicing factor SR45a can act synergistically with CBP20 to regulate the RNA processing of multiple core salt stress factors in A.thaliana.	［27］
	AtSR34b	调控靶基因剪接与mRNA稳定性Regulates splicing and mRNA stability of target genes	通过IRT1的剪接和稳定性促进IRT1蛋白积累Promotes IRT1 protein accumulation via regulating the splicing and stability of IRT1 mRNA	［45］
	AtSAD1	调控剪接效率Regulates splicing efficiency	提高盐胁迫响应基因的剪接效率Improves the splicing efficiency of salt stress response genes	［46］
	AtSKIP	参与剪接体组装Involves in the assembly of the isoform	调控全基因组范围内大量胁迫响应基因的选择性剪接Regulation of alternative splicing in numerous stress response gene across the entire genome	［47］
	AtSRAS1	产生SRAS1.1和SRAS1.2剪接变体Produces SRAS1.1 and SRAS1.2 isoforms	调控COP9信号复合体亚基5A（CSN5A）的蛋白平衡Regulates the protein balance of COP9 signalosome subunit 5A（CSN5A）	［48］
	AtRCD1	产生RCD1.1和RCD1.2剪接变体Produces RCD1.1 and RCD1.2 isoforms	RCD1的剪接变体可减少盐诱导的细胞死亡Splicing isoform of RCD1 reduce salt-induced cell death	［26］
水稻 O. sativa	OsNHX1	Pre-mRNA剪接产生3种变体Pre-mRNA splicing generates three isoforms	过表达异构体的品系耐盐性增强Lines overexpressing the isoform exhibit enhanced salt tolerance	［24］
	OsMAPK5	产生维持激酶活性的OsMAPK5a亚型Produces the OsMAPK5a isoform with retained kinase activity	OsMAPK5a可以增强对盐分胁迫的耐受性OsMAPK5a enhances tolerance to salt stress	［49］
	OsIM1	产生两种功能分化的剪接变体Produces two functionally differentiated isoforms	通过完整功能域维持叶绿体能量代谢稳态，又在胁迫持续时通过变体的降解或反馈调节实现能量分配优化。Maintains chloroplast energy metabolism homeostasis via intact domains； Optimizes energy allocation through isoform degradation or feedback regulation under prolonged stress.	［8］
大麦 H. vulgare	HvDRF1	产生两种含AP2结构域的功能变体Produces two functional isoforms containing the AP2 domain	两种剪接变体均通过AP2结构域发挥转录激活作用，响应盐碱胁迫的脱落酸核心信号通路。Both splicing isoforms exert transcriptional activation through the AP2 domain， responding to the core hormone signal pathway of abscisic acid in response to saline-alkali stress.	［15］
谷子 S. italica	SiCYP19	产生剪接变体Produces splicing isoform	剪接变体可以提高脯氨酸含量和促进活性氧清除能力。Splicing isoforms can increase proline content and promote reactive oxygen species （ROS） scavenging capacity.	［25］
胡杨 P. euphratica	PeuHKT1；3	产生PeuHKT1；3的可变3′剪接位点Generation of variable 3′splice sites for PeuHKT1；3	通过截短变体的降解与反馈调节实现能量分配，优化重塑离子选择性。Optimizes energy allocation and reshaping ion selectivity through degradation and feedback regulation of truncated isoforms.	［17］
大豆 G. max	GmPeNTL9	产生剪接变体Produces splicing isoform	剪接变体激活抗氧化清除系统Splicing isoform activates the antioxidant scavenging system	［18］
	GsSCL30a	GsSCL30a剪接因子对自身进行剪接The splicing factor GsSCL30a undergoes self-splices	GsSCL30a与GsSnRK1协同作用可增强植株的耐碱性GsSCL30a synergistically enhances plant alkaline tolerance with GsSnRK1	［19］
	GmAP2	GmAP2基因中微外显子的缺失Microexon deletion in the GmAP2 gene	AP2基因中微外显子的缺失增强了对盐碱的抗性Deletion of a microexon in the AP2 gene enhances resistance to saline-alkali stress	［20］
百慕大草Cynodon dactylon	CdDHN4	产生CdDHN4-L和CdDHN4-S Produces CdDHN4-L and CdDHN4-S	显著降低电解质渗漏和增强光合能力Significantly reduces electrolyte leakage and enhances photosynthetic capacity	［51］
盐芥T. salsuginea	TsHKT1	选择性剪接产生多种变体Alternative splicing generates multiple isoforms	形成互作复合体调控Na⁺/K⁺平衡Forms an interaction complex to regulate Na⁺/K⁺ balance	［16］

Pre-mRNA选择性剪接对植物盐碱胁迫响应的调控机制

Regulatory mechanisms of pre-mRNA alternative splicing in plant responses to saline-alkali stress

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