WSWXTB微生物学通报Microbiology China0253-2654微生物学通报编辑部中国北京wswxtb-48-11-428210.13344/j.microbiol.china.210059专论与综述Reviews丛枝菌根真菌在蔬菜生产中的研究进展Research progress of arbuscular mycorrhizal fungi in vegetable production王蕾WANGLei1张春楠ZHANGChunnan1李洪波LIHongbo2王红WANGHong345王鑫鑫WANGXinxinsywxx@hebau.edu.cn345*河北农业大学资源与环境科学学院 河北 保定 071001College of Resources and Environment Science, Hebei Agricultural University, Baoding, Hebei 071001, China中国农业科学院农业环境与可持续发展研究所 北京 100081Institute of Environment and Sustainable Development in Agriculture, Chinese Academy of Agricultural Sciences, Beijing 100081, China河北省山区农业技术创新中心 河北 保定 071001Agricultural Technology Innovation Center in Mountainous Areas of Hebei Province, Baoding, Hebei 071001, China河北农业大学河北省山区研究所 河北 保定 071001Mountainous Area Research Institute of Hebei Province, Hebei Agricultural University, Baoding, Hebei 071001, China国家北方山区农业工程技术研究中心 河北 保定 071001Agricultural Engineering Technology Research Center of National North Mountainous Area, Baoding, Hebei 071001, China
王鑫鑫, Tel:0312-7526344;E-mail:sywxx@hebau.edu.cn
WANG Xinxin, Tel: 86-312-7526344; E-mail: sywxx@hebau.edu.cn
In recent years, the application of arbuscular mycorrhizal fungi (AMF) in greenhouse vegetable production has shown that AMF inoculation during vegetable seedling breeding can produce mycorrhizal seedlings with good quality and strong resistance. Via various mycorrhizal effects, the application of AMF in vegetables production has proven that AMF can improve the nutrition and water supply of vegetables, enhance the tolerance of vegetables to environmental stress, increase the resistance of vegetable root diseases and nematodes, and improve the yield and product quality of vegetables. These functions of AMF are of importance in agriculture. Based on the beneficial effects of AMF on vegetables, this paper reviews the research progress of AMF in promoting vegetable growth, increasing vegetable yield and quality, alleviating its abiotic stress, controlling pathogens, and applying AMF in combination with pathogenic bacteria (or pesticides), so as to provide reference for the application of AMF in vegetable production in the future.
丛枝菌根真菌蔬菜生产菌根接种联合接种生物防治arbuscular mycorrhizal fungivegetable productionmycorrhizal inoculationcombined inoculationbiological control华北作物改良与调控国家重点实验室自主课题NCCIR2020ZZ-4宁夏回族自治区重点研发专项2019BBF02026华北作物改良与调控国家重点实验室自主课题(NCCIR2020ZZ-4);宁夏回族自治区重点研发专项(2019BBF02026)Independent Program of State Key Laboratory of North China Crop Improvement and RegulationNCCIR2020ZZ-4Key Research and Development Program of Ningxia Hui Autonomous Region2019BBF02026Independent Program of State Key Laboratory of North China Crop Improvement and Regulation (NCCIR2020ZZ-4); Key Research and Development Program of Ningxia Hui Autonomous Region (2019BBF02026)
盐胁迫下接种AMF可以通过促进植株体内的渗透调节和提高抗氧化酶活性有效降低细胞膜脂过氧化水平,进而缓解盐胁迫对蔬菜造成的伤害以及盐胁迫条件对蔬菜幼苗生长的抑制[45]。接种AMF (F. mosseae)的番茄植株在盐胁迫下的生长、光合速率、抗氧化酶活性、根和茎叶质量均有明显提高[46]。刘耀臣等[47]发现AMF对盐胁迫下芹菜(Apium graveliens)的生长和生理指标会产生显著影响,接种AMF的处理与未接种相比,其株高、茎粗、地上部鲜重和地下部鲜重的受抑制程度显著降低,芹菜植株内可溶性糖和脯氨酸含量有所升高,超氧化物歧化酶、过氧化物酶和过氧化氢酶活性都得到了显著提升。在对盐胁迫下黄瓜(Cucumis sativus)生长、果实产量和品质的研究中发现,与未接种植株相比,接种AMF的黄瓜果实产量更高、营养品质更好,体内的氮、磷、钾、铜和锌含量分别比对照提高了7%、12%、28%、14%和10%,说明AMF在盐渍化土壤依然可以有效促进黄瓜植株生长和对矿物质营养的吸收,改善黄瓜的营养品质,提高其果实产量[48]。
干旱胁迫下,AMF通过改变蔬菜的生理特性和相关基因表达,从而增强蔬菜作物的耐旱性能。在Jezdinský等[52]的一项试验中,与未接种对照相比,接种AMF可以提高干旱胁迫下辣椒植株的光合速率和水分利用效率。在对培育番茄的抗旱性研究中发现,轻度水分胁迫和重度水分胁迫条件下,接种AMF (D. epigaea和F. mosseae)可提高培育番茄公顷产量2%−28%,同时可以防治培育番茄脐腐病的发生,增加培育番茄叶片叶绿素含量、超氧化物歧化酶和过氧化氢酶活性[53]。接种AMF还可以提高干旱胁迫下番茄[54]、西瓜[50]等的水分利用效率,这是因为接种AMF提高了宿主作物的蒸腾效率,改变了宿主的营养吸收[55]。
土壤中重金属浓度升高,无论是必需的还是非必需的,都会对植物产生毒害作用,导致植物生长及营养获取受阻、生物量减少,最终导致植物死亡[73]。AMF与宿主植物根共生可以增加植物根系吸收面积,这是因为真菌菌丝在根际以外探索了更大的非根际土壤,从而提高了植物对水和矿质养分的吸收,在胁迫条件下产生了更大的生物量[74]。在重金属镉(20 mg/kg)的污染下,同时接种3种AMF (F. mosseae、Claroideoglomusetunicatum和R. intraradices)显著提高了番茄的根、茎、叶和果实的干物质重,总干重也显著增加[75]。
AMF与其他生物制剂联合接种对黄瓜幼苗的影响已有报道。Saldajeno等[88]研究表明,在黄瓜幼苗接种镰刀菌(Fusarium spp.)的前6−12天内接种AMF,可以提高黄瓜幼苗对腐烂病的防御率。此后,Saldajeno等[88-89]又对AMF与其他真菌复合施用抑制黄瓜病毒进行了研究,结果表明,在黄瓜种植过程中,腐殖质生长的真菌与AMF (F. mosseae)复合施用,促进了黄瓜生长,抑制了由黄瓜花叶病毒(Cucumber mosaic virus,CMV)、炭疽病菌(Colletotrichum spp.)和立枯丝核菌(Rhizoctonia solani)导致的黄瓜叶病毒、炭疽病和幼苗的腐烂引起的病害。胡玉金等[90]的研究发现,在黄瓜根内接种AMF和DSE (暗隔内生真菌)组合处理的各指标显著高于单接种AMF或DSE,其中组合处理的株高、茎粗、地上及地下部干重和产量均高于其他处理,而且降低线虫繁殖数量及黄瓜发病率的效果最好。通过以上试验研究发现,联合接种不仅可以提高蔬菜作物的抗病能力,而且蔬菜作物的生长及营养的吸收都要高于单接种,说明菌株联合接种对作物的有益作用具有协同效应。
Tanwar等[91]研究了2种AMF (F. mosseae和Acaulospora laevis)与木霉菌(Trichoderma Pers.)联合施用对尖孢镰刀菌(Fusarium oxysporum)引起的番茄枯萎病的生物防治潜力,结果发现,在盆栽条件下,土壤接种F. mosseae,同时在番茄移栽之前根部接种木霉菌,番茄幼苗具有较好的存活能力和抗枯萎病能力。AMF (Glomus fasciculatum)、木霉菌和荧光假单胞菌(Pseudomonas fluorescens)这3种生物防治剂结合使用能够促进番茄生长,对感染尖孢镰刀菌的番茄植株疾病传播的控制效果最高可达94%[92]。Reddy等[93]通过番茄抗病性参数发现,双接种AMF (G. fasciculatum)和腐霉菌(Pythium aphanidermatum)显著抑制了病原菌在番茄根组织中的生长发育。单独接种AMF和双接AMF与荧光假单胞菌联合接种豆类植株,在田间条件下都可以减少根腐病发病率,但AMF与荧光假单胞菌联合接种对于控制疾病和增加产量更有效[94]。由此表明,通过使用多种生物防治剂诱导多种防御机制,比单独使用生物接种剂更能提高抗病性水平。
印楝饼剂与AMF联用能提高蔬菜作物对线虫的防治效果。Rao等[95]研究了印楝饼剂与AMF (F. mosseae)组合对番茄线虫的防控作用,结果表明,菌根番茄幼苗在印楝饼剂改良土中移栽时受线虫侵染最少,在印楝饼剂改良土中进行菌根幼苗移栽处理后,植株生长参数增加,根结指数下降,最终线虫种群减少。Reimann等[96]发现,在番茄线虫防治中,AMF (R. intraradices)与根瘤菌联合接种产生了叠加效应,这些微生物的单一接种使瘿的数量减少了24%−39%,而双接种则减少了60%。这些研究表明,植物促进有益微生物与AMF的特定组合可以减少线虫侵染,共同作为促进蔬菜健康生长的有效工具。
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