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。     

Australia offers unique insight into the ecology of arbuscular mycorrhizal fungi: An opportunity not to be lost

Typified by ancient soils and unique assemblages of flora, Australia provides opportunities to expand our understanding of arbuscular mycorrhizal (AM) fungi. Despite their ubiquity, key aspects of Australian AM fungal ecology remain buried due to our limited knowledge of their biogeography and their potential adaptation to Australia's environmental conditions. This knowledge gap is particularly extraordinary given that the characteristics of the Australian environment are likely to provide unique insights into AM fungal ecology and evolution. Extensive exploration of the diversity and distribution of AM fungi across the continent is overdue. In pursuit of this goal, ecologists should employ the most effective and pragmatic molecular approaches, while making use of well-curated databases. We urge researchers to examine the biogeography of Australian AM fungi meaningfully, leveraging the distinctive attributes of Australian landscapes, such as the demographics of plant mycorrhizal types and the characteristic interplay with fire. Documenting AM fungal communities across Australia will not only provide unique insights into their ecology but is also pivotal to being able to incorporate these organisms into land management for conservation, restoration and sustainable agriculture.     

The screening and selection of inoculant arbuscular-mycorrhizal and ectomycorrhizal fungi

The initiation of a programme of screening and selection of arbuscular-mycorrhizal fungi (AM fungi) and ectomycorrhizal fungi (ECM fungi) for use as inoculant fungi in agriculture, horticulture of forestry will depend on whether inoculation is more appropriate as a management option than manipulation of the indigenous mycorrhizal populations. The greatest immediate potential for the successful use of mycorrhizal inoculants will be in soils and growth media where phosphorus (P) limits plant growth and where the indigenous mycorrhizal fungi are either ineffective at increasing P uptake by the plant or their numbers have been drastically reduced by human influence or natural disturbance. In recent investigations, however, additional benefits to the plant following colonization of roots by effective AM or ECM fungi have been demonstrated. These additional benefits of an effective mycorrhizal association have necessitated a re-evaluation of the optimum screening procedures for isolates of AM and ECM fungi. Both current methodologies and suggestions for alternative approaches to the screening of AM and ECM fungi are discussed in this paper. The problems inherent in choosing suitable experimental conditions to compare different isolates at the screening stage are also addressed. The review also stresses the importance of continued monitoring of introduced mycorrhizal fungi to learn more about their longer-term ecological role, particularly within reforestation or revegetation studies.     

分室培养装置在丛枝菌根真菌研究中的应用及其发展

重点围绕玻璃珠分室培养系统、H形分室培养系统、根排斥室培养系统、供体自养植物的双分室体外培养系统、丛枝菌根(AM)真菌与普通植物根器官的双重培养系统、AM真菌与Ri T-DNA转型根的双重单胞无菌培养系统、AM真菌与Ri T-DNA转型根双重培养的改良分室单胞培养系统等7个不同的分室培养装置, 对AM真菌的培养类型及其应用进行了系统的评述。其中, 采用玻璃珠分室培养装置易于将AM真菌与培养基质分开, 能获得大量纯净的AM真菌繁殖体, 用于研究AM真菌对矿质元素和微量元素的吸收, 具有不可替代的作用。H形分室培养系统和根排斥室(RECs)培养系统均能够获得连续的、可切断的共生菌根网络(CMNs), 可用于研究植物-植物、植物-昆虫之间化感作用产生的信息交流。供体自养植物的双分室培养系统有益于研究AM真菌对宿主植物在单作和混作条件下生长效应的影响。AM真菌与植物根器官的双重培养系统为研究AM真菌的侵染过程及生理、生化特性提供了极大的方便, 同时为纯培养研究提供了重要的理论依据。AM真菌与Ri T-DNA转型根的双重单胞无菌培养体系可以获得AM真菌纯净菌体, 是研究AM真菌遗传、生理、生化等特性的理想方法。以AM真菌与Ri T-DNA转型根的双重单胞无菌培养系统为基础, 可以在菌丝生长室置换培养基、在根室中补充适量碳源, 并多次收获AM真菌繁殖体。转型根改良双重培养系统是提高AM真菌孢子接种剂产量的有效方法。综上所述, AM真菌的分室培养系统已经取得显著进展, 为开展个体、种群、群落等不同层次的菌根生态学研究提供了依据。     

Mycorrhizal associations in woody plant species at the Mt. Usu volcano,Japan

We investigated the association between ectomycorrhizal (ECM) and arbuscular mycorrhizal (AM) fungi and pioneer woody plant species in areas devastated by the eruption of Mt. Usu, Japan, in 2000. We observed eight woody plant species at the research site, most of which were associated with ECM and/or AM fungi. In particular, dominant woody plant species Populus maximowiczii, Salix hultenii var. angustifolia and Salix sachalinensis were consistently associated with ECM fungi and erratically associated with AM fungi. We found one to six morphotypes in the roots of each ECM host and, on average, two in the roots of each seedling, indicating low ECM fungal diversity. ECM colonization ranged from 17 to 42% of root tips. Using morphotyping and molecular analyses, 15 ECM fungi were identified. ECM fungi differed greatly between hosts. However, Laccaria amethystea, Hebeloma mesophaeum, Thelephora terrestris and other Thelephoraceae had high relative colonization, constituting the majority of the ECM colonization in the roots of each plant species. These ECM fungi may be important for the establishment of pioneer woody plant species and further revegetation at Mt. Usu volcano.     

Mycorrhizal status of <Emphasis Type="Italic">Eucalyptus</Emphasis> plantations in south China and implications for management

The aim of this study is to assess the mycorrhizal status of Eucalyptus plantations in south China and to determine the need for inoculation. In four provinces in south China, 155 plantations were sampled for sporocarps of ectomycorrhizal (ECM) fungi, spores of arbuscular mycorrhizal (AM) fungi, and mycorrhizas over 2 years. This study revealed a low above-ground diversity of ECM fungi consisting of 15 taxa fruiting beneath Eucalyptus plantations. The most common ECM genera were Scleroderma and Pisolithus, but they were infrequent. A total of 21 AM fungi, mostly Glomus species, were recognized from spores collected from eucalypt plantations. Four Glomus species were frequently present in soils, but spore density and relative abundance of AM fungi were generally low. Eucalypt roots from all plantation sites were poorly colonized by either ECM fungi or AM fungi. A bioassay with E. urophylla as a bait host, using soils collected from 11 eucalypt plantations, confirmed low levels of inoculum of both ECM and AM fungi in field soil. This is the first integrated study on the mycorrhizal status of eucalypt plantations in China. Findings from this research can be used to encourage adoption of mycorrhizal technology by eucalypt nurseries in the region. The potential of using spores of compatible ECM fungi or collections for forest nurseries is discussed.     

Diversity of arbuscular mycorrhizal fungi colonising roots of the grass species<Emphasis Type="Italic"> Agrostis capillaris</Emphasis> and<Emphasis Type="Italic"> Lolium perenne</Emphasis> in a field experiment

Analysis of arbuscular mycorrhizal (AM) fungal diversity through morphological characters of spores and intraradicular hyphae has suggested previously that preferential associations occur between plants and AM fungi. A field experiment was established to investigate whether AM fungal diversity is affected by different host plants in upland grasslands. Indigenous vegetation from plots in an unimproved pasture was replaced with monocultures of either Agrostis capillaris or Lolium perenne. Modification of the diversity of AM fungi in these plots was evaluated by analysis of partial sequences in the large subunit (LSU) ribosomal RNA (rDNA) genes. General primers for AM fungi were designed for the PCR amplification of partial sequences using DNA extracted from root tissues of A. capillaris and L. perenne. PCR products were used to construct LSU rDNA libraries. Sequencing of randomly selected clones indicated that plant roots were colonised by AM fungi belonging to the genera Glomus, Acaulospora and Scutellospora. There was a difference in the diversity of AM fungi colonising roots of A. capillaris and L. perenne that was confirmed by PCR using primers specific for each sequence group. These molecular data suggest the existence of a selection pressure of plants on AM fungal communities.     

丛枝菌根真菌分类最新进展

近10a来,随着分子生物学技术在丛枝菌根(arbuscularmycorrhiza,AM)研究中的应用,AM真菌分类学得到迅速发展。重点介绍AM真菌的最新分类系统,并对其发展简史作一简单回顾。     

农业生态系统中的AM真菌多样性

农业生态系统复杂庞大,是由如麦田生态系统、水稻田生态系统、果园生态系统、草地生态系统、保护地生态系统等组成的一个复合生态系统。重点介绍农业生态系统中丛枝菌根(AM)和AM真菌多样性,探讨农业生态系统中调控AM真菌多样性的途径以及今后研究的动向。     

亚热带草地中植物优势种与从属种对AM真菌的差异性生长反应

AM真菌能够影响植物生态系统的群落结构.以亚热带草地生态系统为研究对象,调查了两块草地中优势种和从属种的菌根,并在盆栽试验中比较了优势种和从属种对AM真菌的土著菌种和外源菌种Glomus mosseae的生长反应、养分吸收.结果表明,两块草地各自的优势种藿香蓟和两耳草对土著菌种的菌根依赖性分别是41.5%和77.4%,远远高于从属种莎草和毛蓼（16.0%和7.9%）;但是它们对Glomus mosseae的菌根依赖性有所变化,分别是79.6%、44.2%、74.1%和24.9%.这表明,土著菌种是优势种和从属种的形成机制之一,而外源菌种可能改变基于土著菌种而形成的植物群落结构.植株磷营养的分析结果表明,AM真菌对优势种和从属种生长的促进与对磷吸收的促进高度相关,表明AM真菌促进养分吸收是其影响植物群落结构的机制之一.     

丛枝菌根真菌物种多样性研究进展

丛枝菌根(arbuscular mycorrhiza, AM)真菌是生态系统中生物多样性的重要组分之一,具有十分丰富的物种多样性、遗传多样性和功能多样性.该真菌分类地位不断提高已上升至门,下设1个纲、4个目、13个科,19个属,现已报道214种.丛枝菌根对保持生态平衡、稳定和提高生态系统可持续生产力具有重要作用.本文分析了世界范围内丛枝菌根真菌物种多样性研究现状、不同生态系统中影响丛枝菌根真菌物种多样性的关键因子及其调控途径;认为分子生物学技术是今后丛枝菌根真菌物种多样性研究的主要方法.     

青藏高原高寒草地AM真菌分布及其对近自然恢复的生态作用

超载过牧以及全球气候变化等导致大部分青藏高原高寒草地呈现持续退化态势。青藏高原高寒草地退化致使地上植物群落逐渐发生更替,地下土壤微生物群落多样性和丰富度发生改变。本文旨在探析青藏高原高寒草地丛枝菌根（arbuscular mycorrhizal,AM）真菌的分布特征、对近自然恢复的生理生态效应及其作用机制。青藏高原高寒草地中已报道4目14属61种AM真菌,约占已知AM真菌物种的20%。高寒草地禾本科植物根围AM真菌物种丰度最高,而莎草科植物根围AM真菌孢子密度最高。3种高寒草地植被类型中,高寒草原AM真菌丰度最高（33种）,山地灌丛草原次之（32种）,高寒草甸最低（22种）。高寒草原以光壁无梗囊霉Acaulospora laevis和闪亮和平囊霉Pacispora scintillans为优势种,山地灌丛草原以摩西斗管囊霉Funneliformis mosseae为优势种,高寒草甸以光壁无梗囊霉A. laevis、近明球囊霉Claroideoglomus claroideum和闪亮和平囊霉P. scintillans为优势种。高寒草地土著AM真菌与植物构建的菌根网络可以通过调节营养元素吸收、分配,促进植物建植和生长;但是毒杂草入侵可以改变土著AM真菌物种多样性和菌根网络,限制本地植被的实际生态位扩张。退化高寒草地中,AM真菌群落具有高的环境适应性和恢复力,其不仅调控地上植物群落建植和多样性,同时AM真菌建植也增加了代谢产物-球囊霉素相关土壤蛋白产生,进而协同改善地下土壤微生态系统,为退化高寒草地早期植被恢复塑造土壤生境。因此,AM真菌在退化高寒草地近自然恢复中具有较大的应用潜力。     

尖峰岭热带山地雨林根部真菌-植物互作网络结构特征

不同功能群的根部真菌可能会与植物差异性地互作, 并进一步影响地下真菌与植物群落构建。本研究采用Illumina Miseq测序方法检测了海南尖峰岭热带山地雨林中常见植物的根部真菌; 采用网络分析法比较了丛枝菌根(AM)真菌、外生菌根(ECM)真菌, 以及所有根部真菌与植物互作的二分网络(bipartite networks)结构特性。从槭树科、番荔枝科、夹竹桃科、冬青科、棕榈科、壳斗科、樟科和木犀科等8科植物的根系中, 检测到297,831条真菌ITS1序列, 这些序列被划为1,279个真菌分类单元(OTUs), 其中子囊菌门748个、担子菌门354个、球囊菌亚门80个, 以及未知真菌97个。核心根部真菌群落(420个OTUs)中, 至少有三类不同生态功能的真菌常见, 即丛枝菌根真菌(40个OTUs, 占总序列数23.4%)、外生菌根真菌(48个OTUs, 13.9%)和腐生型真菌(83个OTUs, 19.8%)。尖峰岭山地雨林根部真菌-植物互作网络结构特性的指标普遍显著高于/低于假定物种随机互作的零模型期待值。在群落水平, 不同功能型的根部真菌-植物互作网络表现出不同或相反的结构特性, 如丛枝菌根互作网络表现为比零模型预测值高的嵌套性和连接性, 以及比零模型低的专一性, 而外生菌根互作网络呈现出比零模型预测值低的嵌套性和连接性, 以及比零模型高的专一性。在功能群水平, 植物的生态位重叠度在AM互作网络高, 而ECM互作网络低; 真菌的生态位宽度在ECM互作网络窄, 而在AM互作网络较宽。共现(co-occurrence)网络分析进一步揭示, ECM群落的物种对资源的高度种间竞争(植物、真菌高C-score), 以及AM群落的物种无明显种间竞争(低C-score), 可能分别是形成反嵌套ECM互作网络及高嵌套AM互作网络结构的原因。上述结果说明, 尖峰岭山地雨林中至少有两种及以上的种间互作机制调节群落构建: 驱动AM互作网络冗余(nestedness)及ECM互作网络的高生态位分化(专一性)。本研究在同一个森林内探讨了不同功能型的真菌-植物互作特性, 对深入理解热带森林的物种共存机制和生态恢复具有重要意义。     

丛枝菌根真菌对大气CO2浓度升高和增温响应研究进展

全球变化深刻影响着陆地生态系统生物多样性及生态功能。丛枝菌根(AM)真菌可与绝大多数陆生植物根系形成互惠共生体,在协助宿主养分吸收、促进植物生长、维持植物多样性等方面发挥着重要作用。本文主要分析了大气CO₂浓度升高(eCO₂)和增温对森林和草地生态系统AM真菌群落组成及其功能的影响。eCO₂主要通过影响宿主植物、土壤碳(C)输入等方式间接影响AM真菌,可增加AM真菌的多度和活性,影响AM真菌的多样性与群落组成。增温可直接或间接地(通过宿主植物和土壤途径)影响AM真菌,显著改变森林土壤AM真菌的群落组成,但对草地土壤AM真菌群落组成的影响尚无定论。我们提出了当前研究中存在的主要问题及未来应重点关注的内容。本文旨在明晰AM真菌对eCO₂和增温的响应和适应,增进对AM真菌介导的土壤生态功能的认识,为利用AM真菌缓解全球变化、增强土壤功能的韧性和全球变化的生态系统适应性提供依据。     

Dispersal of arbuscular mycorrhizal fungi and plants during succession

Arbuscular mycorrhizal (AM) fungi are important root symbionts that enhance plant nutrient uptake and tolerance to pathogens and drought. While the role of plant dispersal in shaping successional vegetation is well studied, there is very little information about the dispersal abilities of AM fungi. We conducted a trap-box experiment in a recently abandoned quarry at 10 different distances from the quarry edge (i.e. the potential propagule source) over eleven months to assess the short term, within-year, arrival of plant and AM fungal assemblages and hence their dispersal abilities. Using DNA based techniques we identified AM fungal taxa and analyzed their phylogenetic diversity. Plant diversity was determined by transporting trap soil to a greenhouse and identifying emerging seedlings. We recorded 30 AM fungal taxa. These contained a high proportion of ruderal AM fungi (30% of taxa, 79% of sequences) but the richness and abundance of AM fungi were not related to the distance from the presumed propagule source. The number of sequences of AM fungi decreased over time. Twenty seven plant species (30% of them ruderal) were recorded from the soil seed traps. Plant diversity decreased with distance from the propagule source and increased over time. Our data show that AM fungi with ruderal traits can be fast colonizers of early successional habitats.     

不同盐碱度土壤中AM菌根真菌群落结构组成及影响因子

为研究黄河三角洲盐渍土壤中植物根围AM菌根真菌多样性及影响多样性的因素,从东营孤东和孤岛油区采集碱蓬和柽柳植物的根围土壤,鉴定了4种土壤试样中丛枝菌根(Arbuscular mycorrhiza,AM)的群落组成。结果表明:球囊霉属(Glomus)是盐碱地中的优势种,同时还有许多未知真菌;考察不同盐碱度情况下菌根真菌群落结构差异,结果表明:碱蓬根围土壤中AM真菌的多样性高于柽柳,孤东根围土壤AM真菌多样性比孤岛高。相关分析表明,铵态氮含量与AM真菌多样性呈现显著负相关。     

Ericoid mycorrhizal fungi: some new perspectives on old acquaintances

Many ericaceous species colonize as pioneer plants substrates ranging from arid sandy soils to moist mor humus, in association with their mycorrhizal fungi. Thanks to the symbiosis with ericoid mycorrhizal fungi, ericaceous plants are also able to grow in highly polluted environments, where metal ions can reach toxic levels in the soil substrate. For a long time this mycorrhizal type has been regarded as an example of a highly specific interaction between plants and fungi. More recent studies have been challenging this view because some ericoid mycorrhizal endophytes seem also able to colonise plants from very distant taxa. A molecular approach has allowed the investigation of genetic diversity and molecular ecology of ericoid mycorrhizal fungi, and has revealed that ericaceous plants can be very promiscuous, with multiple occupancy of their thin roots. The molecular analysis of sterile morphotypes involved in this symbiosis has also led to deeper understanding of the species diversity of ericoid fungi. Genetic polymorphism of ericoid fungi is wider than previously thought, and often increased by the presence of Group I introns in the nuclear small subunit rDNA.     

亚热带典型树种根毛特征及其与共生真菌的关系

根毛和共生真菌增加了吸收面积,提高了植物获取磷等土壤资源的能力。由于野外原位观测根表微观结构较为困难,吸收细根、根毛、共生真菌如何相互作用并适应土壤资源供应,缺乏相应的数据和理论。该研究以受磷限制的亚热带森林为对象,选取了21种典型树种,定量了根毛存在情况、属性变异,分析了根毛形态特征与共生真菌侵染率、吸收细根功能属性之间的关系,探讨了根表结构对低磷土壤的响应和适应格局。结果表明:1)在亚热带森林根毛不是普遍存在的, 21个树种中仅发现7个树种存有根毛, 4个为丛枝菌根(AM)树种, 3个为外生菌根(ECM)树种。其中,马尾松(Pinus massoniana)根毛出现率最高,为86%;2)菌根类型是理解根-根毛-共生真菌关系的关键,AM树种根毛密度与共生真菌侵染率正相关,但ECM树种根毛直径与共生真菌侵染率负相关; 3) AM树种根毛长度和根毛直径、ECM树种根毛出现率与土壤有效磷含量呈负相关关系。该研究揭示了不同菌根类型树种根毛-共生真菌-根属性的格局及相互作用,为精细理解养分获取策略奠定了基础。     

Evolutionary histories and mycorrhizal associations of mycoheterotrophic plants dependent on saprotrophic fungi

Mycoheterotrophic plants (MHPs) are leafless, achlorophyllous, and completely dependent on mycorrhizal fungi for their carbon supply. Mycorrhizal symbiosis is a mutualistic association with fungi that is undertaken by the majority of land plants, but mycoheterotrophy represents a breakdown of this mutualism in that plants parasitize fungi. Most MHPs are associated with fungi that are mycorrhizal with autotrophic plants, such as arbuscular mycorrhizal (AM) or ectomycorrhizal (ECM) fungi. Although these MHPs gain carbon via the common mycorrhizal network that links the surrounding autotrophic plants, some mycoheterotrophic lineages are associated with saprotrophic (SAP) fungi, which are free-living and decompose leaf litter and wood materials. Such MHPs are dependent on the forest carbon cycle, which involves the decomposition of wood debris and leaf litter, and have a unique biology and evolutionary history. MHPs associated with SAP fungi (SAP-MHPs) have to date been found only in the Orchidaceae and likely evolved independently at least nine times within that family. Phylogenetically divergent SAP Basidiomycota, mostly Agaricales but also Hymenochaetales, Polyporales, and others, are involved in mycoheterotrophy. The fungal specificity of SAP-MHPs varies from a highly specific association with a single fungal species to a broad range of interactions with multiple fungal orders. Establishment of symbiotic culture systems is indispensable for understanding the mechanisms underlying plant–fungus interactions and the conservation of MHPs. Symbiotic culture systems have been established for many SAP-MHP species as a pure culture of free-living SAP fungi is easier than that of biotrophic AM or ECM fungi. Culturable SAP-MHPs are useful research materials and will contribute to the advancement of plant science.