番茄菌根形成过程中抗氧化酶活性的变化及内源茉莉酸的调节作用
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摘要
丛枝菌根（Arbuscular mycorrhiza，简称AM）是自然界最广泛存在的植物与真菌互利互惠共生体，80%以上的陆生植物可借助丛枝菌根真菌（AM fungi, AMF）从土壤中吸收矿物质（尤其是在缺磷状态下），以改善营养状况、提高对生物和非生物胁迫的耐受性以及加强物质和能量循环，改善自然生态系统。因此，有效地提高植物与AMF共生效率具有重要的科学意义和指导生产实践的价值。然而，达到这样目标的前提条件是必需充分认识菌根形成的生物学机理。由于AMF的纯培养十分困难，加之对寄主侵染的低频率以及缺乏同步性，使得对AM形成机理的认识仍很有限。近年来，不断增加的证据表明植物激素参与AM共生过程的各个主要阶段，如寄主对AMF效应剂的感受、信号转导、基因表达、蛋白质修饰等。其中茉莉酸（jasmonic acid, JA）及其衍生物直接参与AM的形成。然而，现有的研究大多数均采用外施JA的方法（如叶面喷洒或灌根），或者是测定植物内源JA的含量，而利用反向遗传学方法（即利用JA生物合成或信号感受缺失突变体）研究JA在AM形成中的作用则鲜见报道或是刚刚起步，对一些科学问题的认识仍十分有限。
我们以番茄野生型植株及其茉莉酸缺失的突变体spr2为材料，采用嫁接技术及人工接种菌根真菌，在实验室控制的条件下，初步研究了番茄内源JA在植物根与菌根真菌共生过程中的作用机理，重点关注JA水平的作用以及地上部分JA对菌根形成的系统性作用。检测的指标包括抗氧化酶超氧化物岐化酶（SOD）、过氧化物酶（POD）和过氧化氢酶（CAT）及其同工酶活性、可溶性糖含量以及脯氨酸水平等。结果显示：
1.内源JA生物合成缺失突变体spr2的AM定殖水平低于野生型的AM定殖水平，表明JA对AM的形成具有一定促进作用。
2.地上部分的JA对AM形成具有一定的系统性促进作用。
3.菌根真菌侵染初期，AM定殖水平高的野生型植株SOD和CAT活性低于未接种的对照，表明菌根形成初期真菌抑制抗氧化酶活性是其侵染的条件之一。
4. AM定殖水平高的野生型植株MDA含量高于定殖水平低的突变体，表明AM定殖引发了植株氧化胁迫。
5.野生型植株AM定殖后（接种60天后）的可溶性糖和脯氨酸含量高于JA缺失突变体，也高于未接种的对照，此外，AM形成效率高的嫁接植株可溶性糖和脯氨酸含量高于AM形成效率低的嫁接，表明AM的形成有利于这些可溶性物质的合成与积累。
Arbuscular mycorrhiza (AM) is the most widespread terrestrial symbiosis and is formedby at least80%of land plant species with fungi that belong to Glomeromycota. The AMhyphal network supply plants with nutrients (predominantly phosphate) and water, in turnobtain carbohydrates from plants. Therefore, AM symbiosis contributes significantly toglobal phosphate and carbon cycling and influences primary productivity in terrestrialecosystems. In addition, mycorrhizal can enhance markedly its host plant resistance topathogens and abiotic stresses. However, so far, the understanding of the mechanism of AMformation still remains limited due to the difficulty to cultivate the pure AM fungi, coupledwith the low frequency infection for the host and the lack of a synchronicity. Fortunately,recent studies demonstrate that several new plant hormones are involved directly in the AMformation, such as strigolactones and jasmonic acid (JA). In the case of JA, however, eventhough there are several lines of evidence including our previous study have shown thatendogenous JA or exogenous application of JA to plants play a crucial role in the symbiosisbetween hosts and AM fungi, the molecular mechanism by which JA regulates AM formationis less understood. In this study, tomato wild-type plants and its mutant spr2with JAdeficiency were used to comparatively investigate the role of endogenous JA in the AMformation between plant roots and AMF, with an emphasis on the function of JA levels andsystemic role of JA in above-ground parts in AM formation. Data were collected includingthe activities of superoxide dismutase (SOD), peroxidase (POD) and catalase (CAT), andtheir isoenzymes shown by polyacrylamide gel electrophoresis (PAGE), malondialdehyde(MDA) contents, soluble sugar contents, and proline levels.
The results showed that:
1. The colonisation of AM in the spr2mutant was lower than that in wild-type plants,indicating that endogrnous JA in plants had a function in AM formation.
2. JA in the above-ground parts of plants had a systemic function in AM formation.
3. At the early stage of AF formation, the activities of SOD and CAT were reduced inwild-type plants relative to their uninoculated controls, suggesting that the inhibition of theseantioxidative enzymes was a prerequisite to the infection of AMF to plants.
4. The MDA levels were higher in wild-type plants with high AM colonisation than in spr2with low AM colonisation, showing that the AM colonisation induce lipid peroxidation.
5. The contents of proline and soluble sugar in wild-type plants were higher than thosein the spr2mutant plants after AM colonization, as well as in the non-inoculated control,together with the contents of proline and soluble sugar in the grafting plants with high AMcolonization being higher than those in the grafting plants with low AM colonization,indicating that the AM colonization was favourable for these soluble compounds productionsor accumulations in plants.
我们以番茄野生型植株及其茉莉酸缺失的突变体spr2为材料，采用嫁接技术及人工接种菌根真菌，在实验室控制的条件下，初步研究了番茄内源JA在植物根与菌根真菌共生过程中的作用机理，重点关注JA水平的作用以及地上部分JA对菌根形成的系统性作用。检测的指标包括抗氧化酶超氧化物岐化酶（SOD）、过氧化物酶（POD）和过氧化氢酶（CAT）及其同工酶活性、可溶性糖含量以及脯氨酸水平等。结果显示：
1.内源JA生物合成缺失突变体spr2的AM定殖水平低于野生型的AM定殖水平，表明JA对AM的形成具有一定促进作用。
2.地上部分的JA对AM形成具有一定的系统性促进作用。
3.菌根真菌侵染初期，AM定殖水平高的野生型植株SOD和CAT活性低于未接种的对照，表明菌根形成初期真菌抑制抗氧化酶活性是其侵染的条件之一。
4. AM定殖水平高的野生型植株MDA含量高于定殖水平低的突变体，表明AM定殖引发了植株氧化胁迫。
5.野生型植株AM定殖后（接种60天后）的可溶性糖和脯氨酸含量高于JA缺失突变体，也高于未接种的对照，此外，AM形成效率高的嫁接植株可溶性糖和脯氨酸含量高于AM形成效率低的嫁接，表明AM的形成有利于这些可溶性物质的合成与积累。
Arbuscular mycorrhiza (AM) is the most widespread terrestrial symbiosis and is formedby at least80%of land plant species with fungi that belong to Glomeromycota. The AMhyphal network supply plants with nutrients (predominantly phosphate) and water, in turnobtain carbohydrates from plants. Therefore, AM symbiosis contributes significantly toglobal phosphate and carbon cycling and influences primary productivity in terrestrialecosystems. In addition, mycorrhizal can enhance markedly its host plant resistance topathogens and abiotic stresses. However, so far, the understanding of the mechanism of AMformation still remains limited due to the difficulty to cultivate the pure AM fungi, coupledwith the low frequency infection for the host and the lack of a synchronicity. Fortunately,recent studies demonstrate that several new plant hormones are involved directly in the AMformation, such as strigolactones and jasmonic acid (JA). In the case of JA, however, eventhough there are several lines of evidence including our previous study have shown thatendogenous JA or exogenous application of JA to plants play a crucial role in the symbiosisbetween hosts and AM fungi, the molecular mechanism by which JA regulates AM formationis less understood. In this study, tomato wild-type plants and its mutant spr2with JAdeficiency were used to comparatively investigate the role of endogenous JA in the AMformation between plant roots and AMF, with an emphasis on the function of JA levels andsystemic role of JA in above-ground parts in AM formation. Data were collected includingthe activities of superoxide dismutase (SOD), peroxidase (POD) and catalase (CAT), andtheir isoenzymes shown by polyacrylamide gel electrophoresis (PAGE), malondialdehyde(MDA) contents, soluble sugar contents, and proline levels.
The results showed that:
1. The colonisation of AM in the spr2mutant was lower than that in wild-type plants,indicating that endogrnous JA in plants had a function in AM formation.
2. JA in the above-ground parts of plants had a systemic function in AM formation.
3. At the early stage of AF formation, the activities of SOD and CAT were reduced inwild-type plants relative to their uninoculated controls, suggesting that the inhibition of theseantioxidative enzymes was a prerequisite to the infection of AMF to plants.
4. The MDA levels were higher in wild-type plants with high AM colonisation than in spr2with low AM colonisation, showing that the AM colonisation induce lipid peroxidation.
5. The contents of proline and soluble sugar in wild-type plants were higher than thosein the spr2mutant plants after AM colonization, as well as in the non-inoculated control,together with the contents of proline and soluble sugar in the grafting plants with high AMcolonization being higher than those in the grafting plants with low AM colonization,indicating that the AM colonization was favourable for these soluble compounds productionsor accumulations in plants.
引文
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