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2024 Volume 3 Issue 6
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LIU Huidi, WANG Zhihui, TIAN Xiaowei. The Role of Photosynthesis in Plant-Pathogen Interactions[J]. PLANT HEALTH AND MEDICINE, 2024, 3(6): 32-41. doi: 10.13718/j.cnki.zwyx.2024.06.004
Citation: LIU Huidi, WANG Zhihui, TIAN Xiaowei. The Role of Photosynthesis in Plant-Pathogen Interactions[J]. PLANT HEALTH AND MEDICINE, 2024, 3(6): 32-41. doi: 10.13718/j.cnki.zwyx.2024.06.004
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The Role of Photosynthesis in Plant-Pathogen Interactions

  • 1.

    College of Horticulture and Landscape, Tianjin Agricultural University, Tianjin 300384, China

  • 2.

    Ankang Branch of Shaanxi Province Tobacco Company, Ankang Shanxi 725000, China

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  • Corresponding author: TIAN Xiaowei
  • Received Date: 14/04/2024
    Available Online: 25/12/2024

  • MSC: S432.44;S476

  • Throughout their growth cycles, plants face challenges from a variety of pathogens. To address these challenges, plants mobilize defense responses via their innate immune systems to counteract pathogen infections. Photosynthesis, as a crucial component in plants' defense against pathogenic microbial infections, not only provides essential energy and metabolites to support plant defense responses but also enhances plant resistance through various mechanisms such as signal transduction and gene expression regulation, ultimately influencing crop yield and quality. Comprehensive understanding of the effects of photosynthesis on plant-pathogen interactions is essential for prevention and control of disease and improving crop yield and quality. This paper comprehensively analyzes the effects of pathogen stress on the tissue structure, physiological, and biochemical processes associated with plant photosynthesis. This paper also briefly summarizes the factors influencing the photosynthesis, and the role of photosynthesis in disease prediction, also the linkage between photosynthesis and plant immunity, thereby offering a foundational reference for further research in plant photosynthesis under pathogen stress.

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The Role of Photosynthesis in Plant-Pathogen Interactions

    Corresponding author: TIAN Xiaowei
  • Received Date: 14/04/2024
  • Available Online: 25/12/2024

Abstract: 

Throughout their growth cycles, plants face challenges from a variety of pathogens. To address these challenges, plants mobilize defense responses via their innate immune systems to counteract pathogen infections. Photosynthesis, as a crucial component in plants' defense against pathogenic microbial infections, not only provides essential energy and metabolites to support plant defense responses but also enhances plant resistance through various mechanisms such as signal transduction and gene expression regulation, ultimately influencing crop yield and quality. Comprehensive understanding of the effects of photosynthesis on plant-pathogen interactions is essential for prevention and control of disease and improving crop yield and quality. This paper comprehensively analyzes the effects of pathogen stress on the tissue structure, physiological, and biochemical processes associated with plant photosynthesis. This paper also briefly summarizes the factors influencing the photosynthesis, and the role of photosynthesis in disease prediction, also the linkage between photosynthesis and plant immunity, thereby offering a foundational reference for further research in plant photosynthesis under pathogen stress.

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  • 植物在生长过程中不断受到外界生物胁迫(细菌、病毒、真菌、寄生植物、昆虫等)和非生物胁迫(干旱、低温、盐分、pH值变化等)的影响[1]. 这些外界压力迫使植物内部启动一系列复杂的生理和生化反应，涉及组织、细胞结构、功能和代谢等多个层面，以抵御胁迫造成的损害[2-7].

    植物病害作为限制全球农业生产的主要因素，严重威胁作物的健康和生长，进而影响作物的产量和品质. 光合作用作为植物生存的基础活动，在面对病原微生物侵袭时扮演着至关重要的角色. 当植物受到病原微生物侵染时，会通过调整气孔开闭状态以及合成特定代谢产物等方式来增强自身抵抗力. 这一过程中，参与光合作用的组织和细胞会发生一系列的变化，植物的光合能力也会发生改变[8]. 具体来说，病原微生物不仅破坏了叶绿体结构，还降低了气孔导度，增加了细胞间二氧化碳浓度，干扰了光合作用相关基因表达，从而降低了光合作用效率[9]. 此外，光合色素的降解进一步影响光能的捕获和转化效率，导致植物光合作用效率进一步下降[10]. 随着光合能力的变化，作物抵御病原微生物的能力也出现波动. 光合作用不仅是植物防御反应的重要能量来源，还可以调控防御信号分子合成，参与碳资源重新分配. 光合作用产物，如碳水化合物，在植物防御信号分子(如SA和JA)的合成中起着重要作用，能够参与调节植物的防御反应，提高自身的抗病能力[11].

    作物产量和品质的提升依赖于光合效率的增强. 因此，深入研究植物如何在病原微生物胁迫条件下调节其光合作用过程，探索有效的干预措施以提高作物抗病性，对于提高农作物产量和改善品质具有极其重要的意义. 这不仅有助于揭示植物与病原微生物互相作用的分子机制，还能为开发新型绿色农业技术提供科学依据.

1.   植物-病原微生物互作对光合作用的影响

  • 光合作用在植物与病害的互作中扮演着重要角色. 光合过程中，植物通过多种方式参与免疫防御反应[12]. 当植物受到病原微生物侵染时，其叶片的组织结构和叶绿素含量会发生变化，导致光合作用能力减弱. 同时，这种变化还会干扰气孔的正常调节功能，使得植物难以吸收足够的二氧化碳进行光合活动，从而直接影响植物的生长[13]. 病原微生物的侵染也会引起植物的氧化反应，诱导植物产生防御相关代谢产物，加剧植物对病害的应激反应，进一步影响其光合能力[14]. 为应对病原微生物的侵染，植物也会启动一系列的抗病机制. 例如，叶绿体内多种信号分子的合成、叶绿素含量的变化、光系统中心的调整以及光能量耗散方式的改变，都是植物抵御病原微生物的重要手段[15]. 此外，植物通过调整气孔的大小和数量，能够减轻病原微生物对其光合作用造成的负面影响，并尽可能优化光合作用条件以提高整体的光合速率.

    • 1.1.   病原微生物对光合作用的影响

    1.1.1.   对叶绿体结构的影响

  • 叶绿体是植物进行光合作用的核心器官，也是植物防御的主战场. 病原微生物通过攻击叶绿体，干扰其正常功能，从而降低植物的光合效率和免疫能力. 这种攻击会对叶绿体的结构和数量造成不同程度的影响[16]. 叶绿体结构变化包括：①叶绿体整体数量减少，叶绿素含量降低，且出现叶绿体聚集现象. ②叶绿体膨大、变形. ③膜结构异常，如出现外周囊泡、细胞质内陷、质膜增生、破裂等. ④叶绿体内部成分的变化，如间质小泡或空泡、膜间囊变大、淀粉粒增多，嗜锇体增多. ⑤叶绿体内部结构异常，如颗粒层消失，类囊体扭曲、松动或扩张，基质消失. 不同病原微生物对叶绿体结构的影响如表 1所示.

    • 1.1.2.   对光化学反应的影响

  • 病原微生物侵染寄主植物后，主要通过两个途径影响光化学反应. 一是病原微生物及其代谢产物通过干扰寄主植物的光合磷酸化过程，二是降低叶绿素含量，导致光化学反应活性降低. 例如，胡运高等[26]在研究稻瘟病粗毒素对水稻光系统Ⅱ的影响时，发现随着毒素浓度的增高，其对水稻光系统的影响作用增强，光合磷酸化作用下降，叶绿素含量降低，显著抑制了水稻的光化学活性. 而TMV病毒侵染甘蓝后，光化学活性却呈增加趋势，直到21~28 d后，光化学活性才受到抑制，迅速降低.

    • 1.1.3.   对植物水分代谢的影响

  • 水分胁迫会造成光抑制，给植物带来不利影响. 另外，水分胁迫也会导致气孔关闭，可能会使光合作用速率下降[27]. Wang等[28]在研究镰刀菌对黄瓜的影响时发现，感染尖孢镰刀菌后，黄瓜植株的吸水性和茎部导水能力明显下降，蒸腾速率和气孔导度显著降低，叶片细胞膜受损. 通过对蒸腾速率和导水能力与光化学活性进行相关性分析，发现它们之间存在显著相关性. 这表明水分代谢会影响植物的光合作用，水分利用率的降低也会导致光合作用下降[29].

    • 1.1.4.   对植物碳代谢的影响

  • 植物叶片作为光合作用的主要器官，可以将光能转化为化学能，维持植物正常的生长繁殖. 然而，由于叶片表面积大，容易受到病原微生物的侵染，导致叶片的代谢过程发生变化. CO2的同化是光合作用过程的重要组成部分，需要在叶绿体中进行，且涉及多种酶的参与. 病原微生物侵染叶片时往往会破坏叶绿体结构和参与光合作用酶的活性，从而抑制CO2的固定. 如受玉米黑粉病感染的玉米叶片组织的C4过程中会受到抑制，新陈代谢的过程会受到阻碍[30]. 感染叶片的葡萄糖与蔗糖的比值会下降[31]. 其中病原微生物侵染主要对宿主植物吸收CO2的速率具有影响，但不同病原微生物对宿主植物的影响程度及进程有所差别. 如大麦黄矮病毒(BYDV)侵染大麦后，植物对CO2的吸收速率明显降低[32]，而在侵染灰葡萄孢的番茄叶片中光合速率会上升[33].

    • 1.1.5.   气孔导度

  • 保卫细胞是植物调节气体交换和蒸腾作用的特化表皮细胞，保卫细胞间的空隙就是气孔. 在以往的植物-病原互作研究中，气孔主要被作为病原微生物侵染植物的入口开展研究. 近些年的研究发现，气孔作为植物固有的免疫系统的一部分，在抵御病原侵染中发挥着积极作用[34]. Melotto等[35]研究发现在细菌侵染植物时，气孔关闭会降低病害的严重程度. 在植物受到病原微生物侵染时，叶片气孔的形态和密度都会发生一系列的适应性变化[36-38]. 气孔随着不同病原微生物的侵染，呈现开放和关闭两种状态，间接影响光合效率[39-40]. 当植物受到病原微生物侵染，气孔关闭，阻碍环境中的CO2进入叶片，向叶绿体的羧化部位扩散，叶片细胞间的CO2浓度下降，影响碳同化，从而使光合速率下降，光合产物的累积及运输减少[13]. 气孔的打开或关闭取决于保护细胞的膨胀或松弛，由ROS、NO、NADPH、H2O2和Ca2+等多种信号调节[41]. 其中部分参与气孔调节的信号也参与着光合作用过程[42].

    植物受到病原微生物侵染时会诱发植物气孔不均匀关闭的现象. 在进行气孔导度测定中，往往会出现气孔导度和光合速率变化趋向相似的情况，但此时光合速率降低的因素不一定是气孔导度降低的结果，也有可能是非气孔限制因素. 有学者认为，只有当胞间CO2浓度降低、气孔限制值增大时，才可以得出光合速率降低是由于气孔导度降低引起的结论[43]. 部分学者认为，限制光合作用的气孔因素和非气孔因素不是相互独立的，而是在动态变化之中，关于气孔与光合作用之间的关系还需要进一步研究[44].

    • 1.2.   光合作用在病原微生物对植物侵染中的作用

    1.2.1.   植物免疫

  • 光合作用是植物生长发育的关键一环，免疫防御是植物应对复杂生长环境的关键. 已有多项研究表明，植物的光合作用与免疫防御之间相互关联. 光合作用可以影响细胞内的多个信号通路，为免疫防御提供物质和能量，而免疫防御过程也会对光合作用产生反馈作用[45]. 在免疫防御的过程中光合部位会发生一系列的变化，其中叶绿体作为光合作用的主要场所，不仅可以与其他细胞器传递信息，也是植物-病原微生物互作的关键战场. 在病原微生物的侵染下，通过诱导活性氧(ROS)产生和调节激素变化等方式参与植物的防御过程[46]. 参与植物免疫的光合作用因素如表 2所示.

    • 1.2.2.   病原微生物定殖

  • 叶绿体作为光合作用的主要器官，在植物免疫中起着重要作用，相应的，叶绿体也成为病原微生物的攻击目标. 不同的病原微生物通过破坏叶绿体组织等方式影响其功能，以促使病原微生物在寄主植物中的定殖. 叶绿体本身是来自其细菌祖先、病毒和宿主植物的各种起源成分的嵌合体. 越来越多的研究表明，叶绿体可以与病原微生物相互作用，涉及病原微生物复制、移动、症状和植物防御[62]. 病原微生物蛋白与叶绿体蛋白间的相互作用如表 3所示.

2.   结论与讨论

  • 光合作用在调节病原微生物侵染和植物免疫等生物过程中扮演着至关重要的角色. 保证植物的光合作用免受病原微生物效应物和植物毒素的影响，是提高作物产量与品质的关键策略之一，也是当前亟待深入研究的领域. 因此，研究者们需要进一步识别更多参与病原微生物侵染及病原微生物防御反应中的叶绿体因子，阐明它们在植物-病原微生物相互作用中的具体功能及其作用机制. 此外，还需探究病原微生物如何调控自身及其相关信号分子(如与叶绿体相关的)的表达，从而影响宿主植物对感染的反应；同时了解这些致病因子或植物防御信号是如何在不同细胞器间传递的. 通过上述研究，可以为开发新的抗病策略奠定基础，即通过调控宿主内特定因子的表达水平来有效控制病害发生，进而为农作物在面对生物胁迫时实现高产、优质的目标提供绿色解决方案.

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