四川都江堰亚热带针阔混交林中主要晚生型外生菌根真菌类群的空间分布

晚生型外生菌根真菌通常出现在森林演替的后期,是成熟林中的优势外生菌根真菌类群。对四川都江堰一片亚热带针阔混交林中的菌根真菌地上群落进行调查,并应用二元逻辑回归分析对晚生型外生菌根真菌的主要类群,即鹅膏菌科、牛肝菌科和红菇科,与周围(5m×5m样方)树种组成的关系进行研究。还应用次级变量分析方法对主要外生菌根真菌类群的空间格局进行了分析。结果表明,非外生菌根树种及某些外生菌根树种对特定类群菌根真菌子实体的出现有抑制作用,而不同类群外生菌根真菌在克隆生长上的差异并不是子实体空间分布的决定因素。我们认为,当研究自然林中外生菌根子实体的空间分布时,除了宿主植物的分布,也应考虑非宿主植物的分布以及菌根真菌相互作用的影响。     

杜鹃花科植物菌根的研究进展

杜鹃花科Ericaceae植物可与土壤真菌形成杜鹃花类菌根ericoid mycorrhizas（ERM）共生体,且广泛分布于全球不同的陆地生态系统,特别是在贫瘠、酸性等严酷的环境中占优势。杜鹃花科植物菌根类型多样,绝大多数宿主具有ERM,还有少量宿主具有其他类型的菌根结构,且常与暗隔内生菌（dark septate endophyte,DSE）并存;ERM的宿主植物除已知的杜鹃花科外,岩梅科Diapensiaceae植物也具有ERM结构;ERM真菌以子囊菌和担子菌为主,主要来自柔膜菌目Helotiales和蜡壳耳目Sebacinales;与杜鹃花科宿主形成ERM的真菌也常与壳斗科Fagaceae、松科Pinaceae等宿主植物形成外生菌根（ectomycorrhiza,ECM）结构;ERM对宿主植物在营养吸收、忍耐贫瘠环境、抵抗重金属污染等能力方面都有积极的促进作用,对环境变化的响应是多样的,生境和季节的变化对ERMF群落的组成和分布有着显著影响,资源比率变化可能改变ERM宿主与其他菌根或非菌根植物之间的竞争关系。本文回顾了近40多年来国内外有关ERM的研究进展,还对ERM研究的前景进行了展望,以期在理论和实践中对杜鹃花科及其菌根的研究能取得更丰硕的成果。     

外生菌根共生:共生真菌多样性及菌根形成的分子机制

大约2%的维管植物能够与17～18个目约280个属的真菌形成外生菌根关系.外生菌根关系的形成、宿主植物的专一化和宿主转移等事件可能是共生真菌物种分化的重要驱动力.比较基因组学研究发现,外生菌根真菌丢失了大量与植物细胞壁物质降解相关的基因.在外生菌根形成过程中,真菌与宿主植物之间借助多样化的信号分子完成相互识别,并且外生菌根真菌能够借助分泌效应蛋白抑制宿主植物的防御反应,促进菌根形成.本文对外生菌根真菌的多样性和分布规律,以及外生菌根形成的分子机制等方面进行了综述,并据此对后续研究提出展望.     

黄山地区松树林外生菌根菌资源及生态分布

1998～2003年,对黄山地区松树林下外生菌根菌资源及生态分布进行了调查,鉴定出外生菌根菌43种,隶属于10科17属.其中马尾松林下有43种、黄山松林下有12种.马尾松幼林下的外生菌根菌种类单一,彩色豆马勃、硬皮马勃类为优势菌种;马尾松成熟林下外生菌根菌种类丰富,其优势种类为红菇科、鹅膏菌科、牛肝菌科和鸡油菌科的一些种类.并分析了树种植被对外生菌根菌分布的影响,菌根真菌的分布与温度、湿度及土壤条件的关系.     

三峡库区马尾松林土壤真菌群落特征及影响因素

土壤真菌是地下生物多样性的重要组成部分,在土壤碳和养分循环中发挥重要功能。本研究基于高通量测序技术对三峡库区22个马尾松林表层土壤(0～10 cm)真菌群落进行研究。结果表明: 子囊菌门与担子菌门为优势真菌类群,常见类群包括散囊菌目、红菇目和银耳目。真菌功能群以腐生真菌与外生菌根真菌为主,欧石南类菌根真菌也较常见。冗余分析表明,无论是对整个土壤真菌群落还是不同真菌功能群,环境变量对群落结构的影响均强于空间变量,说明在较小的空间尺度上生境过滤对土壤真菌群落结构的影响超过扩散限制。地上生物量、电导率、有效磷、容重、碳氮比、硝态氮、粉粒占比是影响真菌群落结构的主要环境因子,但影响不同真菌功能群的最关键环境因子存在差异。     

外生菌根对干旱胁迫的响应

从外生菌根真菌、外生菌根共生体以及外生菌根的间接作用等方面阐述外生菌根如何抵制干旱胁迫,并对未来我国外生菌根的研究提出了建议。干旱可以抑制外生菌根真菌的生长并降低其群落中真菌的多样性,干旱胁迫下外生菌根真菌子实体可以利用深度30cm以下的土壤水,子实体的表面积和体积比可作为筛选抗旱真菌的一个重要因子;在遭受干旱胁迫时,外生菌根共生体可以发生形态变化来应对干旱,同时增加了植株水分的吸收并改善了植物的光合作用、活性氧以及激素等相关代谢;外生菌根对植物生长的促进作用、增加土壤碳汇以及对其他根际微生物生长的促进作用等对宿主植物应对干旱胁迫有利。未来我国外生菌根研究应加强对干旱区优良菌-树组合的筛选工作,同时加大对乡土外生菌根真菌资源的调查力度,未来研究应重点向分子生物学领域推进。     

长茎羊耳蒜菌根真菌类群的研究

研究了广西雅长自然保护区和云南西双版纳自然保护区共3个产地的兰科植物羊耳蒜属长茎羊耳蒜Liparis viridiflora的菌根真菌类群区系组成.根内菌根真菌的核糖体基因内转录间隔区序列(rDNA-ITS)采用PCR技术扩增,克隆,测序并构建系统发育树.结果表明,长茎羊耳蒜根内所检测到的真菌大部分为胶膜菌科Tulasnellaceae真菌;根据序列相似性和系统发育分析,所有真菌可归为12个可操作分类单元(OTU),其中胶膜菌科有7个OTUs,达到总数的90.6%,为优势类群.菌根真菌多样性及区系组成在3个不同产地样本之间存在一定的差异;菌根真菌可能和兰科植物的生境适应性存在一定的相关性.     

外生菌根菌与森林树木的相互关系

生态系统的每个过程都伴随着各种微生物的活动,其中最重要的功能群之一是菌根真菌(菌根菌)。一般认为,菌根菌是自然界多数植物生存最基本的组成部分,陆地上约90％以上的高等植物都具有菌根菌。这些菌类的菌丝体与植物根系结合形成菌根,使植物生长成为可能,使不同种类植物的根系联在一起。根据菌根菌入侵植物根系的方式及菌根的形态特征,菌根可分为外生菌根、内生菌根和内外生菌根3组共7种类型。外生菌根主要出现在松科、桦木科、壳斗科等树种的森林生态系统中,在根系表面形成菌丝鞘,部分菌丝进入根系皮层细胞间隙形成哈氏网表面。菌根菌剂在森林经营中得到广泛地应用。外生菌根菌对森林树木的作用可归纳为：1)促进造林或育苗成活与生长;2)提高森林生态系统中植物的多样性、稳定性和生产力;3)对森林生态系统的综合效应,主要表现在增加植物一土壤联结,改善土壤结构,促进土壤微生物,增强植物器官的功能;4)抗拮植物根部病害病原菌等。树木与菌根菌相互关系研究主要包括：1)菌根共生的机理;2)菌根菌在退化森林生态系统恢复与改造中的作用;3)菌根菌的分布格局与森林生态系统服务功能的关系;4)菌根菌对森林生态系统的综合效应,如菌根菌与森林植物群落结构、物种多样性以及森林系统稳定性和生产力的研究。     

华北特有腐生型兰科植物北京无喙兰根际土壤微生物多样性研究

北京无喙兰(Holopogon pekinensis X. Y. Mu & Bing Liu)为华北地区特有珍稀腐生型兰科植物,分布在海拔约1100 m的杂木林内,生境与本区域内其他腐生型兰科植物(常在1600 m以上桦木林中生长)显著不同。本研究针对北京玉渡山和百花山两个北京无喙兰种群的根际土壤样品开展基于高通量测序技术的根际土壤微生物多样性分析,解析北京无喙兰根际土壤微生物群落组成及多样性。测序分析结果显示,共得到4973个细菌OTU(Operational taxonomic unit),发现北京无喙兰根际土壤的优势细菌类群为变形菌门、放线菌门、拟杆菌门等7个门,优势属有MND1、硝化螺菌属(Nitrospira)和Haliangium等。1914个真菌OTU的分析结果表明,根际土壤优势真菌类群为子囊菌门、担子菌门、结合菌门等;优势属有Archaeorhizomyces、蜡壳耳属(Sebacina)和被孢霉属(Mortierella)等;优势真菌多为外生菌根真菌,可能是北京无喙兰潜在的菌根真菌。多样性指数分析显示,北京无喙兰玉渡山种群根际土壤中的真菌和细菌群落的丰富度和均匀度均高于百花山种群,各种群土壤微生物多样性与北京无喙兰所在种群的乔木种类多样性具有一定的相关性。     

大青山外生菌根真菌资源与生态研究

历经5a,采用踏查与样线调查相结合的方法,对内蒙古大青山主要林区的外生菌根真菌资源与生态进行了系统研究。通过对子实体鉴定及对菌根组织的分离与回接研究,并借鉴前人对大青山大型真菌调查资料,初步认为在大青山分布的外生菌根真菌共163种,占现有国内报道种的26.6%,它们分别隶属于18科、41属。在大青山此次调查发现内蒙新记录种54种,占大青山记录种的33%,还有5个存疑种;通过对分布区内各生态因子研究表明,在大青山外生菌根真菌喜欢生长在有机质丰富、林分郁闭度0.7~0.9、海拔1300~1900 m、阴坡的中部、坡度为20~40°的环境中,而在陡坡、山顶或山麓分布较少;通过对生境土壤理化因子分析结果表明,菌根真菌的存在能够明显降低土壤pH值、并对环境中全N含量有显著的提高,对全P、全K无显著影响,但对速效态N、P、K释放均有显著的促进作用,尤其速效K的释放与对照存在着极显著的差异;从对共生关系研究表明,白蘑科(Tricholomataceae)、红菇科(Russulaceae)、丝膜菌科(Cortinariaceae)、鹅膏科(Amanitaceae)的大部分真菌,以白桦(Betula platyphyllaSuks.)、山杨(Populus davidianaDode.)、云杉(Picea asperataMast.)、蒙古栎[Quercus mongolica(Fisch.)Turcz.]、虎榛子(OstryopsisdavidianaDecne.)为主要宿主,而牛肝菌科(Boletaceae)的大部分真菌主要以油松(Pinus tabulaeformisCarr.)、华北落叶松(Larixprincipisr-upprechtiiMayr.)、云杉、白桦、虎榛子等为主要宿主植物。     

Spatial structure and diversity of woody plants and ectomycorrhizal fungus sporocarps in a natural subtropical forest

Spatial patterns of ectomycorrhizal fungi, ectomycorrhizal plants, and non-ectomycorrhizal plants were investigated in a natural subtropical forest using second-order analysis. The results of spatial pattern analysis showed that the degree of clumping of woody plants and ectomycorrhizal sporocarps were correlated. There was a significantly positive correlation of relative aggregation indices between ectomycorrhizal fungi and both non-ectomycorrhizal trees and ectomycorrhizal saplings. Correlations between percentage of ectomycorrhizal trees and sporocarp occurrence of ectomycorrhizal fungi and between diversities of woody plants and ectomycorrhizal fungi were distance-dependent or scale-related. A significantly high percentage of ectomycorrhizal trees was found only at relatively short distance from ectomycorrhizal fungal sporocarps, and significantly positive correlation of the diversity between woody plants and ectomycorrhizal fungi was found only at relative long distance, which implied that ectomycorrhizal sporocarps prefer ectomycorrhizal-tree-dominant micro-sites at near distances and at relatively large scales, diverse ectomycorrhizal sporocarps could be found in woodlands with high diversity of woody plants. Important factors affecting the spatial distribution, occurrence, and diversity of ectomycorrhizal fungi include spatial pattern of ectomycorrhizal plants and non-ectomycorrhizal plants, percentage of ectomycorrhizal plants, and plant diversity in a natural forest.     

The functioning of ectomycorrhizal fungi in the field: linkages in space and time

Individual trees, either of the same or different species, can be linked spatially and temporally by the hyphae of ectomycorrhizal (ECM) fungi that allow carbon and nutrients to pass among them and promote forest establishment following disturbance. Spatial and temporal linkages between plants influence the function of ECM fungi in the field. Studies indicate that ECM linkages can reduce plant competition for resources, promote forest recovery, and influence the pattern of plant succession. The degree of influence depends on many factors, including the composition and arrangement of the vegetative community and soil and climatic conditions. Management practices that create intense disturbance and loss of organic matter or promote the introduction of non-ectomycorrhizal host species can decrease the ability of plants to form linkages with ECM fungi. Management practices that retain living trees and shrubs and input of organic matter provide the energy source and substrate necessary for ECM linkages. More research is needed to determine the degree to which ECM fungal linkages occur in the field and their role in ecosystem function and long-term health.     

Population genetic structure of an ectomycorrhizal fungus Amanita manginiana in a subtropical forest over two years

The population genetic structure of the late-stage fungus Amanita manginiana in a natural forest in Dujiangyan, southwest China was examined over two years using inter-simple sequence repeat (ISSR) markers. Seven ISSR primers were used and 170 bands were obtained in this population: 134/160 and 135/153 bands were polymorphic for sporocarps of 2001 and 2002, respectively. Each sporocarp represented a single genet in 2001 and 2002, and no identical genets were found between the two years. The results of genetic similarity comparison, using unweighted pair group method with arithmetic means, and analysis of molecular variance, indicated that although genetic variances were mainly within individuals of the same year the genetic variance between years was statistically significant (P<0.001). Relationships between genetic similarity and spatial distance of pairwise sporocarps were also found to be different in the two years. The differences in genetic structure and genetic similarity between individuals of the two years implied that the sporocarps were not likely to be derived from continuous generations, i.e., the sporocarps collected in 2002 were not developed from sexual spores dispersed by sporocarps of 2001. We suggest that the life-cycle traits of ectomycorrhizal (ECM) fungi should be considered in genetic studies on ECM fungal populations.     

Ectomycorrhizal fungi are shared between seedlings and adults in a monodominant Gilbertiodendron dewevrei rain forest in Cameroon

Ectomycorrhizal networks may facilitate the establishment and survival of seedlings regenerating under the canopies of tropical forests and are often invoked as a potential contributor to monodominance. We identified ectomycorrhizal fungi in a monodominant Gilbertiodendron dewevrei (Fabaceae) rain forest in Cameroon, using sporocarps and ectomycorrhizae of three age categories (seedlings, intermediate trees, and large trees) and tentatively revealed nutrient transfer through ectomycorrhizal networks by measuring spontaneous isotopic (¹³C and ¹⁵N) abundances in seedlings. Sporocarp surveys revealed fewer ectomycorrhizal fungal taxa (59 species from 1030 sporocarps) than molecular barcoding of ectomycorrhizal roots (75 operational taxonomic units from 828 ectomycorrhizae). Our observations suggest that ectomycorrhizal fungal diversity is similar to that in other mixed tropical forests and provide the first report of the Tuber‐Helvella lineage in a tropical forest. Despite some differences, all age categories of G. dewevrei had overlapping ectomycorrhizal fungal communities, with families belonging to Thelephoraceae, Russulaceae, Sebacinaceae, Boletaceae, and Clavulinaceae. Of the 49 operational taxonomic units shared by the three age categories (65.3% of the ectomycorrhizal fungal community), 19 were the most abundant on root tips of all categories (38.7% of the shared taxa), supporting the likelihood of ectomycorrhizal networks. However, we obtained no evidence for nutrient transfer from trees to seedlings. We discuss the composition of the ectomycorrhizal fungal community among the G. dewevrei age categories and the possible role of common ectomycorrhizal networks in this rain forest.     

Mycorrhizal colonization status of plant species established in an exposed area following the 2000 eruption of Mt. Usu, Hokkaido, Japan

We investigated the ectomycorrhizal (ECM) and arbuscular mycorrhizal (AM) colonization status of plant seedlings that established in areas devastated by the eruption in March 2000 of Mt. Usu, Hokkaido, Japan. In 2005, we estimated the mycorrhizal colonization ratios and frequencies in seedlings of twelve herbaceous and seven woody plant species. Although arbuscular mycorrhizas were found to colonize Equisetum arvense and Polygonum sachalinense, they were presented at very low frequencies and colonization ratios. Other herbaceous plants exhibited higher frequencies of AM associations and either AM and/or ECM fungal associations were observed in all of the woody plant seedlings investigated. The dominant woody plant species (Populus maximowiczii, Salix sachalinensis and Salix hultenii var. angustifolia) associated mainly with ECM fungi and exhibited variable associations with AM fungi. Mycorrhizal associations were common and significant events for plant establishment, even in the early stages of the revegetation process.     

Fungal soil communities in a young transgenic poplar plantation form a rich reservoir for fungal root communities

Fungal communities play a key role in ecosystem functioning. However, only little is known about their composition in plant roots and the soil of biomass plantations. The goal of this study was to analyze fungal biodiversity in their belowground habitats and to gain information on the strategies by which ectomycorrhizal (ECM) fungi form colonies. In a 2-year-old plantation, fungal communities in the soil and roots of three different poplar genotypes (Populus × canescens, wildtype and two transgenic lines with suppressed cinnamyl alcohol dehydrogenase activity) were analyzed by 454 pyrosequencing targeting the rDNA internal transcribed spacer 1 (ITS) region. The results were compared with the dynamics of the root-associated ECM community studied by morphotyping/Sanger sequencing in two subsequent years. Fungal species and family richness in the soil were surprisingly high in this simple plantation ecosystem, with 5944 operational taxonomic units (OTUs) and 186 described fungal families. These findings indicate the importance that fungal species are already available for colonization of plant roots (2399 OTUs and 115 families). The transgenic modification of poplar plants had no influence on fungal root or soil communities. Fungal families and OTUs were more evenly distributed in the soil than in roots, probably as a result of soil plowing before the establishment of the plantation. Saprophytic, pathogenic, and endophytic fungi were the dominating groups in soil, whereas ECMs were dominant in roots (87%). Arbuscular mycorrhizal diversity was higher in soil than in roots. Species richness of the root-associated ECM community, which was low compared with ECM fungi detected by 454 analyses, increased after 1 year. This increase was mainly caused by ECM fungal species already traced in the preceding year in roots. This result supports the priority concept that ECMs present on roots have a competitive advantage over soil-localized ECM fungi.     

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。     

Diversity and Spatial Structure of Belowground Plant–Fungal Symbiosis in a Mixed Subtropical Forest of Ectomycorrhizal and Arbuscular Mycorrhizal Plants

Plant–mycorrhizal fungal interactions are ubiquitous in forest ecosystems. While ectomycorrhizal plants and their fungi generally dominate temperate forests, arbuscular mycorrhizal symbiosis is common in the tropics. In subtropical regions, however, ectomycorrhizal and arbuscular mycorrhizal plants co-occur at comparable abundances in single forests, presumably generating complex community structures of root-associated fungi. To reveal root-associated fungal community structure in a mixed forest of ectomycorrhizal and arbuscular mycorrhizal plants, we conducted a massively-parallel pyrosequencing analysis, targeting fungi in the roots of 36 plant species that co-occur in a subtropical forest. In total, 580 fungal operational taxonomic units were detected, of which 132 and 58 were probably ectomycorrhizal and arbuscular mycorrhizal, respectively. As expected, the composition of fungal symbionts differed between fagaceous (ectomycorrhizal) and non-fagaceous (possibly arbuscular mycorrhizal) plants. However, non-fagaceous plants were associated with not only arbuscular mycorrhizal fungi but also several clades of ectomycorrhizal (e.g., Russula) and root-endophytic ascomycete fungi. Many of the ectomycorrhizal and root-endophytic fungi were detected from both fagaceous and non-fagaceous plants in the community. Interestingly, ectomycorrhizal and arbuscular mycorrhizal fungi were concurrently detected from tiny root fragments of non-fagaceous plants. The plant–fungal associations in the forest were spatially structured, and non-fagaceous plant roots hosted ectomycorrhizal fungi more often in the proximity of ectomycorrhizal plant roots. Overall, this study suggests that belowground plant–fungal symbiosis in subtropical forests is complex in that it includes “non-typical” plant–fungal combinations (e.g., ectomycorrhizal fungi on possibly arbuscular mycorrhizal plants) that do not fall within the conventional classification of mycorrhizal symbioses, and in that associations with multiple functional (or phylogenetic) groups of fungi are ubiquitous among plants. Moreover, ectomycorrhizal fungal symbionts of fagaceous plants may “invade” the roots of neighboring non-fagaceous plants, potentially influencing the interactions between non-fagaceous plants and their arbuscular-mycorrhizal fungal symbionts at a fine spatial scale.     

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.