丛枝菌根真菌提高植物抗逆性的效应及其机制研究进展

丛枝菌根（arbuscularmycorrhizal,AM）真菌是土壤中重要的生物成员之一,对植物具有多种有益效应。AM真菌的基本功能之一是增强植物的抗逆性,在全球气候变化的今天尤其重要。本文总结了AM真菌降低温度胁迫、水分胁迫、盐胁迫、重金属胁迫、病虫害、以及杂草对植物造成的危害和提高植物抗逆性的效应;阐述了AM真菌提高植物抗逆性的作用机制;并讨论了当前该领域研究存在的难题及今后的展望。旨在为探讨提高植物抗逆性策略与途径提供参考。     

丛枝菌根在植物修复重金属污染土壤中的作用

丛枝菌根（Arbuscular mycorrhizae,AM）是自然界中分布最广的一类菌根,AM真菌能与陆地上绝大多数的高等植物共生,常见于包括重金属污染土壤在内的各种生境中。在重金属污染条件下,AM真菌可以减轻重金属对植物的毒害,影响植物对重金属的吸收和转运,在重金属污染土壤的植物修复中显示出极大的应用潜力。重点介绍了AM真菌对植物重金属耐性的影响及其在植物提取和植物稳定中的应用等方面的进展,讨论了未来研究所面临的任务和挑战。     

丛枝菌根真菌的生态分布及其影响因子研究进展

丛枝菌根(arbuscular mycorrhiza,AM)共生体系对于植物适应各种逆境胁迫具有重要积极作用。AM真菌还能够通过根外菌丝网络调节植物群落结构和演替,深刻影响生态系统结构和功能的稳定性。AM真菌生态生理功能的发挥主要取决于其生态适应性,明确AM真菌在不同环境中的多样性、生态适应性以及对各种生态因子的响应机制,是AM真菌资源管理、功能发掘与利用的前提。迄今为止,有关各种生态因子对AM真菌多样性的影响已有不少研究,但是AM真菌生态分布及其形成机制仍缺乏系统的研究和理论分析。综述了生物因子和非生物因子对AM真菌生态分布的影响,结合大型生物空间分布理论探讨了AM真菌生态分布规律和建成机制,分析了当前本研究领域所存在的问题和动向,以期推动相关研究进展。     

Interactions between arbuscular mycorrhizal fungi and soil bacteria

The soil environment is interesting and complicated. There are so many interactions taking place in the soil, which determine the properties of soil as a medium for the growth and activities of plants and soil microorganisms. The soil fungi, arbuscular mycorrhiza (AM), are in mutual and beneficial symbiosis with most of the terrestrial plants. AM fungi are continuously interactive with a wide range of soil microorganisms including nonbacterial soil microorganisms, plant growth promoting rhizobacteria, mycorrhiza helper bacteria and deleterious bacteria. Their interactions can have important implications in agriculture. There are some interesting interactions between the AM fungi and soil bacteria including the binding of soil bacteria to the fungal spore, the injection of molecules by bacteria into the fungal spore, the production of volatiles by bacteria and the degradation of fungal cellular wall. Such mechanisms can affect the expression of genes in AM fungi and hence their performance and ecosystem productivity. Hence, consideration of such interactive behavior is of significance. In this review, some of the most important findings regarding the interactions between AM fungi and soil bacteria with some new insights for future research are presented.     

丛枝菌根真菌与根围促生细菌相互作用的效应与机制

丛枝菌根(arbuscular mycorrhiza,AM)真菌是植物活体营养专性共生菌,广泛存在于陆地各生态系统中.研究表明,AM真菌与根围促生细菌(plant growth promoting rhizobacteria,PGPR)之间的相互作用,尤其是它们之间的协同作用不仅影响植物养分吸收利用、病原物发生发展、土壤理化特性与生物修复等,而且对于可持续农、林、牧业生产、稳定生态系统都具有十分重要的意义.因此,近年来给予众多关注和研究.综述了AM真菌与PGPR之间的相互影响及其可能的作用机制,以及AM真菌与PGPR协同改善植物营养和生长、协同抑制病原菌、协同修复土壤方面的作用,旨在总结AM真菌与PGPR相互作用的效应与机制方面的最新研究进展,为今后研究发展提供依据.     

Do soil inoculants accelerate dryland restoration? A simultaneous assessment of biocrusts and mycorrhizal fungi

Biological soil crusts (biocrusts) and arbuscular mycorrhizal (AM) fungi are communities of soil organisms often targeted to assist in the achievement of multiple ecological restoration goals. In drylands, benefits conferred from biocrust and AM fungal inoculation, such as improved native plant establishment and soil stabilization, have primarily been studied separately. However, comparisons between these two types of soil inoculants and investigations into potential synergies between them, particularly at the plant community scale, are needed to inform on‐the‐ground management practices in drylands. We conducted two full‐factorial experiments—one in greenhouse mesocosms and one in field plots—to test the effects of AM fungal inoculation, biocrust inoculation, and their interaction on multiple measures of dryland restoration success. Biocrust inoculation promoted soil stabilization and plant drought tolerance, but had mixed effects on native plant diversity (positive in greenhouse, neutral in field) and productivity (negative in greenhouse, neutral in field). In greenhouse mesocosms, biocrust inoculation reduced plant biomass, which was antagonistic to % root length colonized by AM fungi. Inoculation with native or commercial AM fungi did not influence plant establishment, drought tolerance, or soil stabilization in either study, and few synergistic effects of simultaneous inoculation of AM fungi and biocrusts were observed. These results suggest that, depending on the condition of existing soil communities, inoculation with AM fungi may not be necessary to promote dryland restoration goals, while inoculation with salvaged biocrust inoculation may be beneficial in some contexts.     

干旱条件下AM真菌对植物生长和土壤水稳定性团聚体的影响

采用分室培养系统,模拟正常水分和干旱胁迫两种环境条件,探讨不同丛枝菌根真菌(arbuscular mycorrhizal fungi,AMF)对紫花苜蓿(Medicago sativa L.)生长和土壤水稳性团聚体的影响.试验条件下,Glomus intraradices对苜蓿根系的侵染率均显著高于Acaulospora scrobiculata和Diversispora spurcum接种处理.正常水分条件下,供试AM真菌均能显著提高植株生物量及磷浓度.干旱胁迫显著抑制了植株生长和菌根共生体发育,总体上菌根共生体对植株生长没有明显影响,接种D.spurcum甚至趋于降低植株生物量;同时,仅有G.intraradices显著提高了植株磷浓度.AM真菌主要影响到＞2mm的水稳性团聚体数量,以G.intraradices作用效果最为显著.在菌丝室中,G.intraradices显著提高了总球囊霉素含量.研究表明AM真菌对土壤大团聚体形成具有积极作用,而菌根效应因土壤水分条件和不同菌种而异,干旱胁迫下仅有G.intraradices对土壤结构和植物生长表现出显著积极作用.在应用菌根技术治理退化土壤时,需要选用抗逆性强共生效率高的菌株,对于不同AM真菌抗逆性差异的生物学与遗传学基础尚需进一步研究.     

丛枝菌根真菌和根瘤菌互作对苜蓿根际土壤细菌群落结构的影响及PICRUSt功能预测分析

【背景】紫花苜蓿是优良的豆科牧草,可以与丛枝菌根(Arbuscular mycorrhizae,AM)真菌和根瘤菌形成共生关系,接种AM真菌和根瘤菌可以促进土壤氮、磷循环以及提高苜蓿产量。【目的】探究接种AM真菌和根瘤菌对苜蓿根际细菌群落结构和功能的影响。【方法】采集6个不同处理组苜蓿根际、非根际土壤样品,基于细菌16S rRNA基因V3?V4区进行高通量测序,分析比较不同处理组苜蓿根际、非根际土壤中细菌群落分布的规律,并采用PICRUSt软件对不同处理组间菌群功能进行预测。【结果】36个土壤样品中共检测到3 849个OTU,分属于50门59纲132目249科595属398种。其中主要的优势菌门为Proteobacteria (52.81%?81.46%)、Bacteroidetes (7.83%?19.68%)及Actinobacteria (2.21%?16.4%)。与不接种相比,接种根内球囊霉和摩西球囊霉分别提高了Gammaproteobacteria和Bacteroidia有益菌的丰度,接种根瘤菌提高了固氮菌(Alphaproteobacteria)的丰度。PICRUSt功能预测表明,细菌菌群共有35个子功能,菌群功能丰富,代谢为最主要的功能,并且接种根瘤菌可增加氨基酸代谢,从而有利于植株N素循环,而接种AM真菌可能对于N循环有一定的抑制作用,相比于单接种AM真菌,双接种AM真菌和根瘤菌处理组碳水化合物代谢更强,从而更有益于植株的氮、磷循环。【结论】接种AM真菌和根瘤菌可分别提高苜蓿根系与氮、磷循环有关的不同有益菌的丰度,从而更有益于植株的氮、磷循环,该结果为提高植株养分吸收、提高苜蓿产量以及菌肥开发利用提供了科学依据。     

Soil solarization reduces arbuscular mycorrhizal fungi as a consequence of weed suppression

Soil solarization, the process of heating soil by covering fields with clear plastic, is a promising method to reduce populations of soilborne pests and weeds without the use of pesticides. However, the destruction of beneficial organisms such as arbuscular mycorrhizal (AM) fungi also may occur, thereby reducing positive effects of solarization. We compared the effects of solarization and chemical fumigants on the survival of indigenous AM fungi in 1995 and 1996. The infectivity of AM fungi was monitored before and after solarization using a greenhouse bioassay with Sorghum bicolor L. for both years. AM colonization of roots was also monitored in the field 8 months after solarization in 1995. Weed densities were measured 8 months after treatment in 1996. Solarization increased the average daily soil temperature 6-10°C and the maximum soil temperature reached by 10-16°C (5-20 cm depth). Solarization did not reduce the infectivity of AM fungi immediately after the solarization period in either year, as determined by the greenhouse bioassay. Infectivity was greatly reduced in solarized plots 8 months after solarization (over winter) in both years as assessed in the field (1995) or with the greenhouse bioassay (1996). Fumigation with metam sodium at 930 l ha^-1 (350 kg active ingredient ha^-1) reduced the infectivity of AM fungi in both years, and fumigation with methyl bromide at 800 kg ha^-1 eliminated infection by AM fungi. Solarization was as effective as methyl bromide and metam sodium at 930 l ha^-1 in controlling winter annual weeds measured 8 months after treatment. Solarization apparently reduced AM fungi in soil indirectly by reducing weed populations that maintained infective propagules over the winter. Fumigation with metam sodium or methyl bromide directly reduced AM fungi in soil.     

电子废弃物拆解区土壤重金属污染对丛枝菌根真菌多样性的影响

【背景】电子废弃物拆解造成的土壤重金属污染引发的环境问题日益突出,丛枝菌根(arbuscular mycorrhizal,AM)真菌能侵染植物根系并增强植物抵御环境胁迫的能力,具有重要的生态功能和应用潜力。【目的】探究电子废弃物拆解区土壤重金属污染对AM真菌群落结构与多样性的影响,甄别可耐受重金属污染的AM真菌类群。【方法】从浙江台州某典型电子废弃物拆解场地及其周边区域共采集土壤样品12份,针对土壤中AM真菌的18S rRNA基因进行高通量测序以及可操作分类单元(operational taxonomic unit,OTU)相对丰度和多样性指数计算。【结果】该区土壤中AM真菌由原囊霉目(Archaeosporales)、球囊霉目(Glomerales)和多孢囊霉目(Diversisporales)组成,其中球囊霉目占据优势地位。土壤AM真菌多样性指数与重金属的浓度、综合污染指数和潜在生态风险指数间均无显著相关性,但疑似泡囊根生囊霉(Rhizophagus vesiculiferus)的OTU相对丰度与上述重金属污染指标之间均呈显著正相关关系。【结论】R. vesiculiferus可能对重金属污染有极强耐受性,可为今后电子废弃物拆解污染土壤治理提供技术基础。     

Indigenous arbuscular mycorrhizal fungal assemblages protect grassland host plants from pathogens

Plant roots can establish associations with neutral, beneficial and pathogenic groups of soil organisms. Although it has been recognized from the study of individual isolates that these associations are individually important for plant growth, little is known about interactions of whole assemblages of beneficial and pathogenic microorganisms associating with plants.We investigated the influence of an interaction between local arbuscular mycorrhizal (AM) fungal and pathogenic/saprobic microbial assemblages on the growth of two different plant species from semi-arid grasslands in NE Germany (Mallnow near Berlin). In a greenhouse experiment each plant species was grown for six months in either sterile soil or in sterile soil with one of three different treatments: 1) an AM fungal spore fraction isolated from field soil from Mallnow; 2) a soil pathogen/saprobe fraction consisting of a microbial community prepared with field soil from Mallnow and; 3) the combined AM fungal and pathogen/saprobe fractions. While both plant species grew significantly larger in the presence of AM fungi, they responded negatively to the pathogen/saprobe treatment. For both plant species, we found evidence of pathogen protection effects provided by the AM fungal assemblages. These results indicate that interactions between assemblages of beneficial and pathogenic microorganisms can influence the growth of host plants, but that the magnitude of these effects is plant species-specific.     

The effects of arbuscular mycorrhizal (AM) fungal and garlic mustard introductions on native AM fungal diversity

Introduced, non-native organisms are of global concern, because biological invasions can negatively affect local communities. Arbuscular mycorrhizal (AM) fungal communities have not been well studied in this context. AM fungi are abundant in most soils, forming symbiotic root-associations with many plant species. Commercial AM fungal inocula are increasingly spread worldwide, because of potentially beneficial effects on plant growth. In contrast, some invasive plant species, such as the non-mycorrhizal Alliaria petiolata, can negatively influence AM fungi. In a greenhouse study we examined changes in the structure of a local Canadian AM fungal community in response to inoculation by foreign AM fungi and the manipulated presence/absence of A. petiolata. We expected A. petiolata to have a stronger effect on the local AM fungal community than the addition of foreign AM fungal isolates. Molecular analyses indicated that inoculated foreign AM fungi successfully established and decreased molecular diversity of the local AM fungal community in host roots. A. petiolata did not affect molecular diversity, but reduced AM fungal growth in the greenhouse study and in a in vitro assay. Our findings suggest that both introduced plants and exotic AM fungi can have negative impacts on local AM fungi.     

桑及菌根桑在库区消落带的生态重建功能及应用潜力

三峡工程在提供丰富水电资源造福人类的同时,也引发了一些生态环境问题,对库区环境、生物、气候以及人们的生活都产生了不同程度的影响。桑树休眠和生长发育节律与库区水体涨落规律的反向偶联,桑的菌根结构在改善根际土质量,促进桑树水分和养分吸收,增强桑树对盐碱、干旱、重金属的耐受性,提高幼苗的存活率,促进桑苗生长,促进植物群落正向演替等生态过程中发挥着重要的作用。分析了三峡库区消落带周期性水淹的生态特征和岸生植物生态恢复障碍,综述了桑及菌根桑在消落带的生态特性和应用潜力,最后提出了菌根桑生物技术的应用所面临的新课题和挑战。     

Plant performance in stressful environments: interpreting new and established knowledge of the roles of arbuscular mycorrhizas

Arbuscular mycorrhizal (AM) symbioses are formed by approximately 80% of vascular plant species in all major terrestrial biomes. In consequence an understanding of their functions is critical in any study of sustainable agricultural or natural ecosystems. Here we discuss the implications of recent results and ideas on AM symbioses that are likely to be of particular significance for plants dealing with abiotic stresses such as nutrient deficiency and especially water stress. In order to ensure balanced coverage, we also include brief consideration of the ways in which AM fungi may influence soil structure, carbon deposition in soil and interactions with the soil microbial and animal populations, as well as plant-plant competition. These interlinked outcomes of AM symbioses go well beyond effects in increasing nutrient uptake that are commonly discussed and all require to be taken into consideration in future work designed to understand the complex and multifaceted responses of plants to abiotic and biotic stresses in agricultural and natural environments.     

AM真菌在草原生态系统中的功能

AM真菌是土壤生态系统中重要的微生物类群,能与陆地生态系统中80%以上的高等植物建立共生体系。目前,AM真菌在维持草原生态系统稳定性中的功能已经成为生态学研究的热点问题之一。基于此,从植物个体、种群、群落和生态系统等不同层次探究AM真菌在维持植物群落多样性和草原生态系统稳定性中的功能。分析发现在个体水平上,AM真菌对宿主植物具有促生效应、抑制效应或中性效应。在种群水平上,分析AM真菌对不同宿主植物吸收土壤矿质营养的分配和调控策略,围绕构成草原植被的两大组成成分:牧草和有毒植物,论述AM真菌对植物种群增长和衰败的调控机制,并从草原植物群落的物种多样性和稳定性角度,探讨AM真菌与植物群落之间的相关性。在生态系统水平上,围绕AM真菌对草原生态系统的演替和退化草原的修复等展开论述,以期为利用AM真菌开展草原生态系统保护和恢复治理提供理论依据,并对草原菌根生态学领域未来的研究进行展望。     

Effects of the soil microbiome on the demography of two annual prairie plants

1. Both mutualistic and pathogenic soil microbes are known to play important roles in shaping the fitness of plants, likely affecting plants at different life cycle stages.
2. In order to investigate the differential effects of native soil mutualists and pathogens on plant fitness, we compared survival and reproduction of two annual tallgrass prairie plant species (Chamaecrista fasciculata and Coreopsis tinctoria) in a field study using 3 soil inocula treatments containing different compositions of microbes. The soil inocula types included fresh native whole soil taken from a remnant prairie containing both native mutualists and pathogens, soil enhanced with arbuscular mycorrhizal (AM) fungi derived from remnant prairies, and uninoculated controls.
3. For both species, plants inoculated with native prairie AM fungi performed much better than those in uninoculated soil for all parts of the life cycle. Plants in the native whole prairie soil were either generally similar to plants in the uninoculated soil or had slightly higher survival or reproduction.
4. Overall, these results suggest that native prairie AM fungi can have important positive effects on the fitness of early successional plants. As inclusion of prairie AM fungi and pathogens decreased plant fitness relative to prairie AM fungi alone, we expect that native pathogens also can have large effects on fitness of these annuals. Our findings support the use of AM fungi to enhance plant establishment in prairie restorations.

     

Mutually beneficial legume symbioses with soil microbes and their potential for plant production

Legumes develop different mutually beneficial symbioses with soil microbes, such as arbuscular mycorrhizal (AM) fungi, nodule bacteria and plant growth promoting bacteria. Symbioses supply the plants with nutrients (predominantly with nitrogen and phosphorus), protect them from pathogens and abiotic stresses and improve soil microbial biodiversity and fertility. The synergistic activity of beneficial soil microbes (BSM) on the plants has great importance for the use of multi-component symbiotic systems in low-input sustainable environmentally-friendly agrotechnologies. However, the complex nature of the AM symbiosis when in a multi-component symbiosis (plant-fungus-bacteria) creates complications for the fungus to produce AM fungal propagules and poses questions (a) about the effectiveness of the fungus per se in interactions with the plants, without associates, and (b) about the necessity of using sterile/axenic conditions for the production of the AM fungi based inoculants because of any mixing and competition by microbes from the inoculants with the local soil microbial consortia. The legume genes controlling interactions with BSM (including genes responsible for effectiveness of such interactions) should be considered as a united genetic system. The plant genome is more stable than that of microbes and therefore crop plants should select beneficial microbes and control the effectiveness of the whole plant-microbe system in the field for the benefit of the crop and therefore of human beings. There is clearly a need to breed legume crops with improved performance under sustainable conditions involving interactions with BSM and optimising the use of agrochemicals.     

Testing Potential Effects of Maize Expressing the Bacillus thuringiensis Cry1Ab Endotoxin (Bt Maize) on Mycorrhizal Fungal Communities via DNA- and RNA-Based Pyrosequencing and Molecular Fingerprinting

The cultivation of genetically modified (GM) crops has increased significantly over the last decades. However, concerns have been raised that some GM traits may negatively affect beneficial soil biota, such as arbuscular mycorrhizal fungi (AMF), potentially leading to alterations in soil functioning. Here, we test two maize varieties expressing the Bacillus thuringiensis Cry1Ab endotoxin (Bt maize) for their effects on soil AM fungal communities. We target both fungal DNA and RNA, which is new for AM fungi, and we use two strategies as an inclusive and robust way of detecting community differences: (i) 454 pyrosequencing using general fungal rRNA gene-directed primers and (ii) terminal restriction fragment length polymorphism (T-RFLP) profiling using AM fungus-specific markers. Potential GM-induced effects were compared to the normal natural variation of AM fungal communities across 15 different agricultural fields. AM fungi were found to be abundant in the experiment, accounting for 8% and 21% of total recovered DNA- and RNA-derived fungal sequences, respectively, after 104 days of plant growth. RNA- and DNA-based sequence analyses yielded most of the same AM fungal lineages. Our research yielded three major conclusions. First, no consistent differences were detected between AM fungal communities associated with GM plants and non-GM plants. Second, temporal variation in AMF community composition (between two measured time points) was bigger than GM trait-induced variation. Third, natural variation of AMF communities across 15 agricultural fields in The Netherlands, as well as within-field temporal variation, was much higher than GM-induced variation. In conclusion, we found no indication that Bt maize cultivation poses a risk for AMF.     

Symbiotic soil fungi enhance ecosystem resilience to climate change

Substantial amounts of nutrients are lost from soils through leaching. These losses can be environmentally damaging, causing groundwater eutrophication and also comprise an economic burden in terms of lost agricultural production. More intense precipitation events caused by climate change will likely aggravate this problem. So far it is unresolved to which extent soil biota can make ecosystems more resilient to climate change and reduce nutrient leaching losses when rainfall intensity increases. In this study, we focused on arbuscular mycorrhizal (AM) fungi, common soil fungi that form symbiotic associations with most land plants and which increase plant nutrient uptake. We hypothesized that AM fungi mitigate nutrient losses following intensive precipitation events (higher amount of precipitation and rain events frequency). To test this, we manipulated the presence of AM fungi in model grassland communities subjected to two rainfall scenarios: moderate and high rainfall intensity. The total amount of nutrients lost through leaching increased substantially with higher rainfall intensity. The presence of AM fungi reduced phosphorus losses by 50% under both rainfall scenarios and nitrogen losses by 40% under high rainfall intensity. Thus, the presence of AM fungi enhanced the nutrient interception ability of soils, and AM fungi reduced the nutrient leaching risk when rainfall intensity increases. These findings are especially relevant in areas with high rainfall intensity (e.g., such as the tropics) and for ecosystems that will experience increased rainfall due to climate change. Overall, this work demonstrates that soil biota such as AM fungi can enhance ecosystem resilience and reduce the negative impact of increased precipitation on nutrient losses.     

Field microplot performance of the peach-almond hybrid GF-677 after inoculation with arbuscular mycorrhizal fungi in a replant soil infested with root-knot nematodes

The effects of arbuscular mycorrhizae (AM) on the development and nutrition of the peach almond hybrid GF-677 rootstock in a replant soil heavily infested with Meloidogyne javanica were evaluated in field microplot conditions for two growing seasons. There was a significant beneficial effect of mycorrhizal inoculation on plant growth and nutrition in previously pasteurized replant soil. In natural replant soil, early inoculation with a mixed AM inoculum of Glomus intraradices, Glomus mosseae and Glomus etunicatum did not affect growth parameters. Whilst inoculation with these AM fungi led to suppression of root-knot nematode reproduction, natural mycorrhizal colonization of the replant soil with native AM fungi did not. Accepted: 6 December 2000