根系分泌物介导下植物-土壤-微生物互作关系研究进展与展望

doi:10.3724/SP.J.1258.2014.00027

摘要/Abstract

摘要：

根系分泌物是植物与土壤进行物质交换和信息传递的重要载体物质, 是植物响应外界胁迫的重要途径, 是构成植物不同根际微生态特征的关键因素, 也是根际对话的主要调控者。根系分泌物对于生物地球化学循环、根际生态过程调控、植物生长发育等均具有重要功能, 尤其是在调控根际微生态系统结构与功能方面发挥着重要作用, 调节着植物-植物、植物-微生物、微生物-微生物间复杂的互作过程。植物化感作用、作物间套作、生物修复、生物入侵等都是现代农业生态学的研究热点, 它们都涉及十分复杂的根际生物学过程。越来越多的研究表明, 不论是同种植物还是不同种植物之间相互作用的正效应或是负效应, 都是由根系分泌物介导下的植物与特异微生物共同作用的结果。近年来, 随着现代生物技术的不断完善, 有关土壤这一“黑箱”的研究方法与技术取得了长足的进步, 尤其是各种宏组学技术(meta-omics technology), 如环境宏基因组学、宏转录组学、宏蛋白组学、宏代谢组学等的问世, 极大地推进了人们对土壤生物世界的认知, 尤其是对植物地下部生物多样性和功能多样性的深层次剖析, 根际生物学特性的研究成果被广泛运用于指导生产实践。深入系统地研究根系分泌物介导下的植物-土壤-微生物的相互作用方式与机理, 对揭示土壤微生态系统功能、定向调控植物根际生物学过程、促进农业生产可持续发展等具有重要的指导意义。该文综述了根系分泌物的概念、组成及功能, 论述了根系分泌物介导下植物与细菌、真菌、土壤动物群之间的密切关系, 总结了探索根际生物学特性的各种研究技术及其优缺点, 并对该领域未来的研究方向进行了展望。

关键词: 生态效应, 微生态系统, 根际, 根系分泌物, 信号分子

Abstract:

Root exudates have specialized roles in nutrient cycling and signal transduction between a root system and soil, as well as in plant response to environmental stresses. They are the key regulators in rhizosphere communication, and can modify the biological and physical interactions between roots and soil organisms. Root exudates play important roles in biogeochemical cycle, regulation of rhizospheric ecological processes, and plant growth and development, and so on. Root exudates also serve roles in the plant-plant, plant-microbe, and microbe-microbe interactions. Plant allelopathy, intercropping system, bioremediation, and biological invasion are all the focal subjects in the field of contemporary agricultural ecology. They all involve the complex biological processes in rhizosphere. There are increasing evidences that various positive and negative plant-plant interactions within or among plant populations, such as allelopathy, consecutive monoculture problem, and interspecific facilitation in intercropping system, are all the results of the integrative effect of plant-microbe interactions mediated by root exudates. Recently, with the development of biotechnology, the methods and technologies relating to soil ecological research have achieved a remarkable progress. In particular, the breakthroughs of meta-omics technologies, including environmental metagenomics, metatranscriptomics, metaproteomics, and metabonomics, have largely enriched our knowledge of the soil biological world and the biodiversity and function diversity belowground. Research on plant-soil-microbe interactions mediated by root exudates has important implications for elucidating the functions of rhizosphere microecology and for providing practical guidelines. The concept and components of root exudates as well as the functions are reviewed in this paper. An overview on the root-bacteria, root-fungi, and root-fauna interactions is presented in detail. Methods to study root exudates and microbial communities are reviewed and the aspects needed to be further studied are also suggested.

Key words: ecological effect, microecology, rhizosphere, root exudate, signal molecule

吴林坤, 林向民, 林文雄. 根系分泌物介导下植物-土壤-微生物互作关系研究进展与展望. 植物生态学报, 2014, 38(3): 298-310. DOI: 10.3724/SP.J.1258.2014.00027

WU Lin-Kun, LIN Xiang-Min, LIN Wen-Xiong. Advances and perspective in research on plant-soil-microbe interactions mediated by root exudates. Chinese Journal of Plant Ecology, 2014, 38(3): 298-310. DOI: 10.3724/SP.J.1258.2014.00027

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链接本文: https://www.plant-ecology.com/CN/10.3724/SP.J.1258.2014.00027

https://www.plant-ecology.com/CN/Y2014/V38/I3/298

图/表 3

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组成 Component	物质 Substance
糖类 Sugar	葡萄糖、果糖、核糖、蔗糖、木糖、鼠李糖、阿拉伯糖、寡糖、聚多糖 Glucose, fructose, ribose, sucrose, xylose, rhamnose, arabinose, oligosaccharide, polysaccharides
氨基酸类 Amino acid	精氨酸、赖氨酸、组氨酸、亮氨酸、天冬氨酸、谷氨酸、脯氨酸、苯丙氨酸 Arginine, lysine, histidine, leucine, aspartic acid, glutamic acid, proline, phenylalanine
有机酸 Organic acid	柠檬酸、草酸、苹果酸、酒石酸、乳酸、丙二酸、丙酮酸、丁酸 Citrate, oxalate, malate, tartaric acid, lactate, malonic acid, pyruvic acid, butyrate
酚酸类 Phenolic	对羟苯甲酸、香豆酸、丁香酸、香草酸、阿魏酸、肉桂酸 p-hydroxybenzoic acid, cumaric acid, syringic acid, vanillic acid, ferulic acid, cinnamic acid
脂肪酸 Fatty acid	油酸、亚麻酸、硬脂酸、软脂酸、棕榈酸 Oleic acid, linolenic acid, stearic acid, palmitic acid, palmitic acid
甾醇类 Sterol	油菜素甾醇、胆甾醇、谷甾醇、豆甾醇 Brassinosteroid, cholesterol, sitosterol, stigmasterol
蛋白质 Protein	过氧化物酶、半乳糖苷酶、磷酸水解酶、吲哚乙酸氧化酶、蛋白酶、多肽 Peroxidase, galactosidase, phosphohydrolase, IAA oxidase, protease, polypeptide
生长因子 Growth factor	生物素、泛酸、胆碱、肌醇、硫胺素、尼克酸、维生素B₆ Biotin, pantothenic acid, choline, inositol, thiamine, nicotinic acid, vitamin B₆
其他 Others	CO₂、乙烯、质子、核苷、黄酮类化合物、植物生长素、植物抗毒素 CO₂, ethylene, proton, nucleoside, flavonoid, auximone, phytoalexin

组成 Component	物质 Substance
糖类 Sugar	葡萄糖、果糖、核糖、蔗糖、木糖、鼠李糖、阿拉伯糖、寡糖、聚多糖 Glucose, fructose, ribose, sucrose, xylose, rhamnose, arabinose, oligosaccharide, polysaccharides
氨基酸类 Amino acid	精氨酸、赖氨酸、组氨酸、亮氨酸、天冬氨酸、谷氨酸、脯氨酸、苯丙氨酸 Arginine, lysine, histidine, leucine, aspartic acid, glutamic acid, proline, phenylalanine
有机酸 Organic acid	柠檬酸、草酸、苹果酸、酒石酸、乳酸、丙二酸、丙酮酸、丁酸 Citrate, oxalate, malate, tartaric acid, lactate, malonic acid, pyruvic acid, butyrate
酚酸类 Phenolic	对羟苯甲酸、香豆酸、丁香酸、香草酸、阿魏酸、肉桂酸 p-hydroxybenzoic acid, cumaric acid, syringic acid, vanillic acid, ferulic acid, cinnamic acid
脂肪酸 Fatty acid	油酸、亚麻酸、硬脂酸、软脂酸、棕榈酸 Oleic acid, linolenic acid, stearic acid, palmitic acid, palmitic acid
甾醇类 Sterol	油菜素甾醇、胆甾醇、谷甾醇、豆甾醇 Brassinosteroid, cholesterol, sitosterol, stigmasterol
蛋白质 Protein	过氧化物酶、半乳糖苷酶、磷酸水解酶、吲哚乙酸氧化酶、蛋白酶、多肽 Peroxidase, galactosidase, phosphohydrolase, IAA oxidase, protease, polypeptide
生长因子 Growth factor	生物素、泛酸、胆碱、肌醇、硫胺素、尼克酸、维生素B₆ Biotin, pantothenic acid, choline, inositol, thiamine, nicotinic acid, vitamin B₆
其他 Others	CO₂、乙烯、质子、核苷、黄酮类化合物、植物生长素、植物抗毒素 CO₂, ethylene, proton, nucleoside, flavonoid, auximone, phytoalexin

功能 Function (Neumann, 2007)	组成 Compound
营养功能 Nutritional function
为微生物生长提供碳、氮源 Available sources of carbon and nitrogen for microbes	低分子量糖类、氨基酸、羧酸类 Low molecular weight sugar, amino acid, carboxylic acid
植物营养吸收 Plant nutrient acquisition
通过金属螯合提高对P、Fe、Zn、Mn的吸收 Mobilization of P, Fe, Zn and Mn, by metal chelation	柠檬酸、草酸、苹果酸、酒石酸、植物铁载体 Citrate, oxalate, malate, malonate, tartaric acid, phytosiderophore
通过还原作用提高对Fe、Mn的吸收 Mobilization of Fe and Mn, by contact reduction	酚类、苹果酸、柠檬酸 Phenolic, malate, citrate
回收利用有机磷酯类中的P元素 Mobilization and retrieval of P from organic P esters	根细胞分泌磷酸水解酶 Root-secretory phosphohydrolase
吸引固氮菌 Chemoattraction of N₂-fixing microbes	类黄酮、苹果酸、植物黏液 Flavonoid, malate, mucilage
诱导共生固氮菌表达Nod基因和抗性相关基因以抵御植物抗毒素 Nod-gene inducers and resistance-inducers against phytoalexins in symbiotic N₂ fixation	黄酮类、黄烷酮(衍生物)、异黄酮 Flavone, flavanone, isoflavone
诱导形成菌根的信号物质 Signal functions in establishment of mycorrhizal associations	类黄酮、糖、氨基酸 Flavonoid, sugar, amino acid
为菌根真菌提高碳源 Available carbon source for mycorrhizal fungi	转化酶 Invertase
保护功能 Protective function
通过络合缓解铝毒 Detoxification of Al₃⁺ by complexation	柠檬酸、草酸、苹果酸、酚类、植物黏液、分泌蛋白 Citrate, oxalate, malate, phenolic, mucilage, secretory protein
响应重碳酸盐毒胁迫 In response to bicarbonate toxicity	苹果酸、柠檬酸 Malate, citrate
保护根部分裂组织, 提高根部与土壤的接触, 提高土壤保水能力 Protection of the root meristem, improved root-soil contact, increased water-holding capacity	植物黏液 Mucilage
植物抗毒素, 抵御病原菌、寄生植物、竞争者 Phytoalexin, defense against of pathogens, parasites and competitors	苯醌、氢化奎宁、皂角苷、几丁质酶、根边缘细胞 Quinone, hydroquinine, saponin, chitinase, root border cell

功能 Function (Neumann, 2007)	组成 Compound
营养功能 Nutritional function
为微生物生长提供碳、氮源 Available sources of carbon and nitrogen for microbes	低分子量糖类、氨基酸、羧酸类 Low molecular weight sugar, amino acid, carboxylic acid
植物营养吸收 Plant nutrient acquisition
通过金属螯合提高对P、Fe、Zn、Mn的吸收 Mobilization of P, Fe, Zn and Mn, by metal chelation	柠檬酸、草酸、苹果酸、酒石酸、植物铁载体 Citrate, oxalate, malate, malonate, tartaric acid, phytosiderophore
通过还原作用提高对Fe、Mn的吸收 Mobilization of Fe and Mn, by contact reduction	酚类、苹果酸、柠檬酸 Phenolic, malate, citrate
回收利用有机磷酯类中的P元素 Mobilization and retrieval of P from organic P esters	根细胞分泌磷酸水解酶 Root-secretory phosphohydrolase
吸引固氮菌 Chemoattraction of N₂-fixing microbes	类黄酮、苹果酸、植物黏液 Flavonoid, malate, mucilage
诱导共生固氮菌表达Nod基因和抗性相关基因以抵御植物抗毒素 Nod-gene inducers and resistance-inducers against phytoalexins in symbiotic N₂ fixation	黄酮类、黄烷酮(衍生物)、异黄酮 Flavone, flavanone, isoflavone
诱导形成菌根的信号物质 Signal functions in establishment of mycorrhizal associations	类黄酮、糖、氨基酸 Flavonoid, sugar, amino acid
为菌根真菌提高碳源 Available carbon source for mycorrhizal fungi	转化酶 Invertase
保护功能 Protective function
通过络合缓解铝毒 Detoxification of Al₃⁺ by complexation	柠檬酸、草酸、苹果酸、酚类、植物黏液、分泌蛋白 Citrate, oxalate, malate, phenolic, mucilage, secretory protein
响应重碳酸盐毒胁迫 In response to bicarbonate toxicity	苹果酸、柠檬酸 Malate, citrate
保护根部分裂组织, 提高根部与土壤的接触, 提高土壤保水能力 Protection of the root meristem, improved root-soil contact, increased water-holding capacity	植物黏液 Mucilage
植物抗毒素, 抵御病原菌、寄生植物、竞争者 Phytoalexin, defense against of pathogens, parasites and competitors	苯醌、氢化奎宁、皂角苷、几丁质酶、根边缘细胞 Quinone, hydroquinine, saponin, chitinase, root border cell