光控植物免疫: 从光信号通路到免疫应答的调控网络

doi:10.11983/CBB25108

植物学报 ›› 2025, Vol. 60 ›› Issue (5): 786-803.DOI: 10.11983/CBB25108 cstr: 32102.14.CBB25108

光控植物免疫: 从光信号通路到免疫应答的调控网络

吴玉俊¹^,²^,^*(), 李英菊³, 罗巧玉¹, 马永贵¹^,²

¹青海师范大学生命科学学院, 青海省青藏高原药用动植物资源重点实验室, 西宁 810008
²青海师范大学青藏高原地表过程与生态保育教育部重点实验室, 西宁 810008
³兰州大学草地农业科技学院草种创新与草地农业生态系统全国重点实验室, 兰州 730020

收稿日期:2025-06-12 接受日期:2025-07-28 出版日期:2025-09-10 发布日期:2025-07-30
通讯作者: *吴玉俊, 理学博士, 兰州大学“萃英”博士后。现任青海师范大学副教授, 生物学硕士生导师。青海省“杰出青年基金”获得者, 中组部“西部之光”访问学者, 青海省“昆仑英才”。主要研究方向为植物抗病免疫信号转导及高原特色植物资源的开发利用。先后主持国家自然科学基金、博士后科学基金、青海省杰出青年基金、青海省重点研发与转化国际合作专项、甘肃省青年基金、中央高校博士后创新基金及教育部重点实验室开放课题等项目。作为第一作者、通讯作者及主要完成作者, 在Proc Natl Acad Sci USA、Cell Research、Plant Physiology和Plant Journal等国际高水平期刊发表论文20余篇。E-mail: gsyc_wyj@126.com
基金资助:
青海省自然科学基金(杰出青年基金)(2024-ZJ-907);甘肃省科学基金青年基金(22JR5RA526);青海师范大学中青年科研基金(2023QZR013)

Light-regulated Plant Immunity: The Regulatory Network From Light Signaling Pathways to Immune Responses

Wu Yujun¹^,²^,^*(), Li Yingju³, Luo Qiaoyu¹, Ma Yonggui¹^,²

¹Key Laboratory of Medicinal Animal and Plant Resources of Qinghai-Tibetan Plateau in Qinghai Province, School of Life Sciences, Qinghai Normal University, Xining 810008, China
²Ministry of Education Key Laboratory of Tibetan Plateau Land Surface Processes and Ecological Conservation, Qinghai Normal University, Xining 810008, China
³State Key Laboratory of Herbage Improvement and Grassland Agro-ecosystems, College of Pastoral Agriculture Science and Technology, Lanzhou University, Lanzhou 730020, China

Received:2025-06-12 Accepted:2025-07-28 Online:2025-09-10 Published:2025-07-30
Contact: *E-mail: gsyc_wyj@126.com

摘要/Abstract

摘要： 解析植物抗病免疫机制是抗病作物育种和国家粮食安全保障的重要科学基础。光受体作为植物感知环境信号的核心组分, 不仅参与植物生长发育的精细调控, 还在植物与病原菌互作中发挥关键信号枢纽作用。现有研究表明, 植物光受体通过直接或间接的作用方式, 依赖COP1/SPA复合体、HY5和PIFs等一系列光信号调控元件, 通过调控植物抗性防御基因的时空表达以及防御激素的合成代谢等多层级途径, 实现光信号与模式触发免疫(PTI)和效应子触发免疫(ETI)信号的精密整合, 使植物在有限的资源下巧妙地协同生长与免疫的平衡。近年来, 光信号与植物免疫系统的交互作用已成为植物生物学的研究热点, 其交互机制的解析也为未来抗病作物育种提供了新方向。该文聚焦于光受体整合调控植物免疫信号的分子机制, 重点综述了光受体介导的免疫启动机制及其与免疫激素信号的时空动态整合模式, 展望了光遗传学技术在解析光信号与免疫信号交互作用机制中的应用潜力, 旨在为基于光信号调控元件的抗病作物分子设计育种提供全新的理论依据和技术路径。

关键词: 光受体, 光信号转导, 植物抗病, 免疫调控机制, 环境适应

Abstract: Unraveling the mechanisms of plant disease resistance and immunity is crucial for breeding disease-resistant crops and safeguarding national food security. Photoreceptors, which are central for perceiving environmental signals, not only fine-tune plant growth and development, but also serve as key signaling hubs in plant-pathogen interactions. Studies have demonstrated that photoreceptor interact directly or indirectly with the COP1/SPA complex, HY5, PIFs, and other light-signaling components. By regulating the spatiotemporal expression of resistance-related defense genes and controlling the synthesis and response networks of defense-related hormones, photoreceptors precisely integrate light signals with pattern-triggered immunity (PTI) and effector-triggered immunity (ETI), thereby balancing plant growth and immunity. In recent years, research into the interaction between light signaling and plant immune systems has become a hot topic in the field of plant biology. Elucidating these underlying mechanisms offers new directions for breeding disease-resistant crops. This paper focuses on the molecular mechanisms of plant disease resistance regulated by photoreceptors, particularly the immune activation mechanisms mediated by photoreceptors and their spatiotemporal integration with immune-related hormone signals. Additionally, it delves into the potential application of optogenetic technology in studying this interaction. The aim is to provide new theoretical and technical avenues for future molecular breeding of disease-resistant crops, which will be based on photoreceptor modification and signal transduction pathways.

Key words: photoreceptor, light signaling, plant disease resistance, immune regulation mechanism, environmental adaptation

吴玉俊, 李英菊, 罗巧玉, 马永贵. 光控植物免疫: 从光信号通路到免疫应答的调控网络. 植物学报, 2025, 60(5): 786-803.

Wu Yujun, Li Yingju, Luo Qiaoyu, Ma Yonggui. Light-regulated Plant Immunity: The Regulatory Network From Light Signaling Pathways to Immune Responses. Chinese Bulletin of Botany, 2025, 60(5): 786-803.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB25108

https://www.chinbullbotany.com/CN/Y2025/V60/I5/786

图/表 2

图1 光受体调控植物生长与免疫的分子模式 (A) 当植物感知红光刺激后, 细胞质中的Ca2+浓度瞬时升高, 同时光敏色素蛋白phyB发生构象重排并被激活。Ca2+信号激活的CPK随即与光活化的phyB互作, 通过磷酸化修饰促使其入核。入核的phyB进而介导下游PIF转录因子(TF)的降解, 由此调控生长与免疫相关基因的表达。在低R/FR比值的光环境或phyB缺失条件下, JAZ10蛋白稳定性增强, 同时促进DELLA蛋白及COP1介导的转录因子HY5降解, 从而抑制茉莉酸(JA)信号并激活赤霉素(GA)信号转导。最终, 通过调控生长发育以及抗性防御相关基因的表达, 实现红光诱导的光形态建成与抗病免疫的协同调控。(B) 在蓝光照射下, CRY1被激活后可诱导LURP1的转录, 为植物抵御病原体的攻击做好准备。同时蓝光介导的LURP1的棕榈酰基化修饰促使其从细胞质转移到质膜, 通过直接与细胞表面的免疫传感器复合物相互作用增强其功能, 进而调控植物的病原相关分子模式激发的免疫(PTI)响应。此外, 在蓝光作用下, 蓝光激活的CRYs家族蛋白通过增强水杨酸(SA)受体NPR1与PIF4的相互作用, 促进NPR1介导的PIF4泛素化降解, 进而协同调控下游生长与免疫相关基因的表达。其次, CRY2在蓝光下通过抑制COP1介导的泛素化降解途径, 维持NLR蛋白的稳定性, 从而确保植物效应蛋白激发的免疫(ETI)信号的正常激活。(C) 紫外光(UV-B)激活光受体UVR8后, 二聚体结构的UVB8解离为单体, 并促进单体化的UVR8在细胞核内积累, 使之能与下游的调节因子COP1作用, 进而解除其对HY5、PIF和BBX等转录因子活性的抑制, 最终影响下游生长与免疫响应基因的表达。同时, 单体化的UVB8可在细胞核内直接与BES1、WRKY、MYB和TCP4等转录因子结合, 通过抑制或增强它们的DNA结合能力, 正调控或负调控下游靶基因的表达, 从而协同平衡植物的光适应性生长与抗病免疫过程。箭头表示直接或间接激活作用, 钝线表示直接或间接抑制作用。

Figure 1 The molecular mechanism of photoreceptor in regulating plant growth and immunity (A) Upon perceiving red light, plants experience a temporary increase in cytosolic Ca2+ concentration. Simultaneously, the photoreceptor phyB undergoes conformational changes and becomes activated. Subsequently, Ca2+-activated CPK interacts with light-activated phyB, phosphorylating it to facilitate its nuclear entry. Once in the nucleus, phyB triggers the degradation of the transcription factor (TF) PIF4, thereby regulating the expression of genes related to growth and immunity. Conversely, under low R/FR light conditions or in the absence of phyB, the stability of the JAZ10 protein is enhanced and promotes the degradation of DELLA and the transcription factor HY5 via COP1-mediated processes. Consequently, jasmonic acid (JA) signaling is suppressed while gibberellin (GA) signaling is activated. Overall, these processes modulate the expression of growth-related and defense-related genes, enabling the co-regulation of plant morphogenesis and immune responses under red light. (B) Under blue light, activated CRY1 induces LURP1 transcription, priming plants to fend off pathogen assaults. Blue light also promotes LURP1 palmitoylation, redirecting it from the cytoplasm to the plasma membrane. As a result, modified LURP1 interacts with cell surface immune receptor complexes and regulates pattern-triggered immunity (PTI) responses. In addition, blue light-activated CRYs strengthen the interaction between the salicylic acid (SA) receptor NPR1 and PIF4. This facilitates PIF4 ubiquitination and degradation via NPR1, coordinating the expression of downstream growth- and immunity-related genes. Meanwhile, bathed in blue light, CRY2 stabilizes NLR proteins by inhibiting COP1-mediated ubiquitination, ensuring proper activation of effector-triggered immunity (ETI) signaling. (C) Upon activation by UV-B radiation, the UVR8 photoreceptor undergoes a dimer-to-monomer transition. These monomers then accumulate in the nucleus and bind to the regulatory factor COP1. This interaction inhibits COP1-mediated degradation of transcription factors such as HY5, PIF, and BBX, thereby regulating downstream genes involved in growth and immune responses. In addition, monomeric UVR8 can directly interact with transcription factors like BES1, WRKY, MYB, and TCP4 within the nucleus. By modulating their DNA-binding activities, UVR8 either enhances or suppresses the expression of target genes. This dual regulatory mechanism allows plants to coordinate and balance photoadaptive growth with disease resistance. Lines with arrows mean direct or indirect activation while blunt lines mean direct or indirect repression.

表1 光信号通路中下游元件在光信号及免疫信号转导中的作用

Table 1 The role of downstream components in the phototransduction pathway in light and immune signaling transduction

基因家族	蛋白类型	在光信号通路中的作用	在免疫信号通路中的作用	参考文献
COP1	E3泛素连接酶	负调控抑制光形态建成、下胚轴伸长及气孔运动, 参与开花调控	调控作用待明确影响抗病蛋白的稳定性, 参与脱落酸介导的气孔免疫	Gangappa and Kumar, 2018; Lim et al., 2018; Chen et al., 2021a; Gao et al., 2024a
PIF	转录因子	负调控抑制下胚轴伸长, 促进子叶张开, 调控开花及营养生长	负调控抑制防御相关基因的表达, 抑制气孔关闭及叶绿素积累	Gangappa et al., 2017; Zhang et al., 2024; Wang et al., 2025
HY5	转录因子	正调控促进光形态建成和下胚轴伸长, 诱导去黄化	正调控激活抗性防御基因的表达, 调控活性氧响应基因、水杨酸(SA)合成及响应基因以及WRKY转录因子等植物免疫相关元件的表达	Chen et al., 2013, 2021b; Chai et al., 2015; Yang et al., 2023
MYB	转录因子	负调控维持PIF蛋白稳定性, 抑制下胚轴伸长, 调控花青素生物合成	既能正调控也能负调控调控抗性基因的表达、木质素/类黄酮/角质层蜡的生物合成、多糖信号转导以及激素防御信号转导	Kim et al., 2020; He et al., 2020; Li et al., 2020; Yu et al., 2023
CBF	转录因子	负调控维持PIF蛋白稳定性, 激活花青素合成基因的表达	正调控激活抗氧化相关基因、抗病基因及植保素合成基因的表达	Shi et al., 2016; Li et al., 2024b
BBX	转录因子	既能正调控也能负调控促进子叶展开和下胚轴生长, 确保叶绿素合成与光系统组装的协调	既能正调控也能负调控调控气孔免疫与质外体免疫, 调控钙依赖蛋白激酶的活性, 影响模式触发免疫(PTI)、抗氧化基因及茉莉酸(JA)响应基因的表达	Luo et al., 2023; Ju et al., 2025; Xie et al., 2025
BES	转录因子	负调控调控植物激素信号转导, 影响下胚轴伸长	负调控调控胼胝质合酶基因和活性氧生成基因的表达, 参与JA介导的抗性防御反应	Kang et al., 2015; Shin et al., 2016; Xiong et al., 2022
TCP	转录因子	负调控抑制光形态建成相关蛋白的合成, 抑制PIF的转录激活活性	既能正调控也能负调控诱导抗病防御相关基因的表达, 参与效应子触发免疫(ETI)信号转导	Mukhtar et al., 2011; Weßling et al., 2014; Kim et al., 2014; Li et al., 2024a
WRKY	转录因子	正调控促进下胚轴伸长及叶片发育	既能正调控也能负调控参与抗病防御相关基因、SA及JA合成基因的表达, 参与抗病次级代谢产物的合成	Javed and Gao, 2023; Yu et al., 2024; Zhang et al., 2025
FHY3/FAR1	转录因子	正调控促进叶绿素生物合成与幼苗生长	负调控依赖于HEMB1, 调控活性SA的产生	Tang et al., 2012; Wang et al., 2016b

表1 光信号通路中下游元件在光信号及免疫信号转导中的作用

Table 1 The role of downstream components in the phototransduction pathway in light and immune signaling transduction

基因家族	蛋白类型	在光信号通路中的作用	在免疫信号通路中的作用	参考文献
COP1	E3泛素连接酶	负调控抑制光形态建成、下胚轴伸长及气孔运动, 参与开花调控	调控作用待明确影响抗病蛋白的稳定性, 参与脱落酸介导的气孔免疫	Gangappa and Kumar, 2018; Lim et al., 2018; Chen et al., 2021a; Gao et al., 2024a
PIF	转录因子	负调控抑制下胚轴伸长, 促进子叶张开, 调控开花及营养生长	负调控抑制防御相关基因的表达, 抑制气孔关闭及叶绿素积累	Gangappa et al., 2017; Zhang et al., 2024; Wang et al., 2025
HY5	转录因子	正调控促进光形态建成和下胚轴伸长, 诱导去黄化	正调控激活抗性防御基因的表达, 调控活性氧响应基因、水杨酸(SA)合成及响应基因以及WRKY转录因子等植物免疫相关元件的表达	Chen et al., 2013, 2021b; Chai et al., 2015; Yang et al., 2023
MYB	转录因子	负调控维持PIF蛋白稳定性, 抑制下胚轴伸长, 调控花青素生物合成	既能正调控也能负调控调控抗性基因的表达、木质素/类黄酮/角质层蜡的生物合成、多糖信号转导以及激素防御信号转导	Kim et al., 2020; He et al., 2020; Li et al., 2020; Yu et al., 2023
CBF	转录因子	负调控维持PIF蛋白稳定性, 激活花青素合成基因的表达	正调控激活抗氧化相关基因、抗病基因及植保素合成基因的表达	Shi et al., 2016; Li et al., 2024b
BBX	转录因子	既能正调控也能负调控促进子叶展开和下胚轴生长, 确保叶绿素合成与光系统组装的协调	既能正调控也能负调控调控气孔免疫与质外体免疫, 调控钙依赖蛋白激酶的活性, 影响模式触发免疫(PTI)、抗氧化基因及茉莉酸(JA)响应基因的表达	Luo et al., 2023; Ju et al., 2025; Xie et al., 2025
BES	转录因子	负调控调控植物激素信号转导, 影响下胚轴伸长	负调控调控胼胝质合酶基因和活性氧生成基因的表达, 参与JA介导的抗性防御反应	Kang et al., 2015; Shin et al., 2016; Xiong et al., 2022
TCP	转录因子	负调控抑制光形态建成相关蛋白的合成, 抑制PIF的转录激活活性	既能正调控也能负调控诱导抗病防御相关基因的表达, 参与效应子触发免疫(ETI)信号转导	Mukhtar et al., 2011; Weßling et al., 2014; Kim et al., 2014; Li et al., 2024a
WRKY	转录因子	正调控促进下胚轴伸长及叶片发育	既能正调控也能负调控参与抗病防御相关基因、SA及JA合成基因的表达, 参与抗病次级代谢产物的合成	Javed and Gao, 2023; Yu et al., 2024; Zhang et al., 2025
FHY3/FAR1	转录因子	正调控促进叶绿素生物合成与幼苗生长	负调控依赖于HEMB1, 调控活性SA的产生	Tang et al., 2012; Wang et al., 2016b

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光控植物免疫: 从光信号通路到免疫应答的调控网络

Light-regulated Plant Immunity: The Regulatory Network From Light Signaling Pathways to Immune Responses

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