兰科植物菌根真菌的研究进展

兰科（Orehidaceae）是仅次于菊科（Composltae）的一个兴盛、复杂的植物类群,广布于全球,多数种是著名的药用植物和珍贵花卉。兰科植物具有三大特点：第一、其花形状奇特,色彩艳丽,芳香宜人,授粉机制独特而复杂;第二、种子细小,仅具未分化的原胚;第三、在生活史中,与真菌共生形成内生苗根。因而,长期以来受到人们的普遍喜爱,带来了较高的商业利益,科学家,园艺工作者也从各个角度对兰科植物作了大量的研究。本文从兰科菌根的共生物之———菌根真菌的角度对有关其分类及与植物之间的专一性的研究状况进行了分析讨论,旨在为从…     

兰科植物黄花白及Bletilla ochracea内生真菌多样性分析

黄花白及Bletilla ochracea是一种地生兰科植物,也是中国传统中药材之一。以黄花白及为实验材料,对其叶和根组织中内生真菌类群组成及多样性进行分析。结果表明,从黄花白及植株的叶片和根组织块(段)中分离到可培养内生真菌140株,根据形态特征将它们鉴定为16个分类单元,其中包括10种子囊菌和6种担子菌。从叶片组织中分离的内生真菌有6种,其中刺盘孢属Colletotrichum真菌为叶片组织中的优势种类;从根组织中分离内生真菌有10种,瘤菌根菌属Epulorhiza和腊壳菌属Sebacina真菌构成了根组织内生真菌的优势类群。根组织可培养内生真菌的多样性(H′=1.968)要高于叶片组织(H′=1.459)。     

菌根真菌多样性与植物多样性的相互作用研究进展

菌根共生双方多样性影响着生态系统的过程与功能。菌根真菌-寄主植物之间的共生组合存在偏好性或特异性,这导致菌根真菌对寄主植物的效益差异和寄主植物对菌根真菌的利益差别：两者在互利共生过程中不仅相互选择,还存在相互促进与制约的关系(如互补与选择效应、竞争),从而在一定程度上决定生态系统的演化与发展。本文概述了植物多样性与菌根真菌多样性的相互影响,探讨了两者互作可能存在的调控因素与机制,对存在的问题和争议进行了总结,并提出了进一步研究的方向。深入阐明植物多样性与菌根真菌多样性之间的互作关系,将丰富生物共生学理论,增强菌根应用潜力及生物多样性的维持。     

菌根真菌是唱响生物共生交响曲的主角——菌根真菌专辑序言

地球上任何生物都不是单独进行生命活动和进化的,即生物之间更多的是依靠密切联合、共生互助、需求互补和共同发展。其中,陆地生态系统中的强大分解者真菌和强大生产者植物关系密切,特别是以菌根真菌(mycorrhizal fungi)为代表的植物共生真菌自始至终与植物共生,这一强-强共生联合在维持生态平衡、保存生态系统可持续生产力与生态系统综合服务功能体系中,其分布之广、作用之多、功能之强和贡献之大,可谓名副其实的生物共生体系中的主导者。近年来,中国在真菌与植物共生研究领域成绩斐然,占据世界重要地位。本期《菌物学报》"菌根真菌专刊"刊登了2篇综述和20篇研究报告。综述分别对丛枝菌根(AM)真菌在煤矿区生态修复应用研究和兰科植物与菌根真菌的营养关系进行了总结和热点论述;研究报告分别对菌物界球囊菌门AM真菌全球公认种的中文学名给予了全面规范正确的描述、对中国部分林区红桦外生菌根真菌多样性、华山松印度块菌菌根中的块菌交配型基因、AM真菌对枳吸收磷和分泌磷酸酶的影响、转Bt基因棉叶片腐熟物抑制AM真菌定殖及菌根对磷的吸收、低pH影响AM真菌丛枝发育和磷的吸收、接种AM真菌与间作对红壤上玉米和大豆种间氮素竞争的影响、AM真菌及其菌种组合对植物根结线虫病害的影响以及接种AM真菌和施加铁可协同降低水稻砷累积等方面进行了研究。本期内容基本体现了中国菌根真菌分类、物种多样性、生理学、生态学、生理生态效应与作用机制研究的最新进展,对当前和今后开展植物共生真菌的研究具有重要的引领作用。     

共生真菌对兰科植物种间杂交后代种子萌发的效应

石斛属(Dendrobium)植物在种子共生萌发过程中与真菌有着较为专一的共生关系, 为探讨这种共生关系在种间杂交后代上的进化和适应, 深入理解兰科植物和真菌共生关系的形成机制, 该研究利用能有效促进铁皮石斛(Dendrobium officinale)和齿瓣石斛(D. devonianum)种子萌发形成幼苗, 并具有较强专一性的胶膜菌属(Tulasnella)真菌SSCDO-5和瘤菌根菌属(Epulorhiza)真菌FDd1, 开展真菌对铁皮石斛和D. tortile种间杂交种子萌发效应的研究。结果表明, 在真菌与种子共生培养68天时, SSCDO-5菌株和FDd1菌株都能有效地促进杂交种子形成原球茎和幼苗, 两个接菌处理之间无显著差异, 来源于铁皮石斛的SSCDO-5菌株不但没有表现出优势, 反而在杂交石斛幼苗形成率上低于来源于齿瓣石斛的FDd1菌株(SSCDO-5: (22.13 ± 6.62)%; FDd1: (29.53 ± 5.51)%); SSCDO-5菌株和铁皮石斛在幼苗形成和发育阶段的共生专一性并没有在杂交后代上得到遗传或表现, 或者说是杂交打破了这种专一性的共生关系, 使得杂交后代能够和不同的真菌建立新的共生关系。该结果不支持关于共生真菌专一性是石斛属植物杂交后代形成的重要限制因素的假设, 推测石斛属植物在幼苗分化和发育阶段与真菌这种专一性的共生关系是在适应特定生态环境的过程中形成和建立的。     

腐生型兰科植物研究进展

腐生型兰科植物又称完全菌根异养型兰(Fully Mycoheterotrophic Orchids),无绿叶,不含叶绿素,不能进行光合作用制造有机物,完全依靠与其共生的真菌提供营养。近年来,腐生兰独特的生活方式引起了生物学家对其生理生态及进化问题的广泛关注,但目前仍缺乏系统深入的研究。为此,作者通过整理《中国植物志》和近年来报道的相关文献,发现分布在中国的腐生型兰科植物约有23属81种,主要集中在天麻属(Gastrodia)(24种)、鸟巢兰属(Neottia)(8种)、无叶兰属(Aphyllorchis)(6种)、山珊瑚属(Galeola)(5种)等。由于腐生兰特殊的营养需求,相比于其他营养类型的兰科植物,其菌根真菌有较大区别。腐生兰的菌根真菌主要有两大类,即外生菌根真菌(Ectomycorrhizal fungi,ECM)和非丝核菌类的腐生菌(Non-rhizoctonia SAP fungi)。该文还综述了腐生兰与其菌根真菌的专一性、营养来源以及系统进化等问题,提出了目前研究存在的问题,并对今后的研究工作进行了展望,以期为腐生兰的资源保护及再生研究提供参考。     

春兰和蕙兰菌根真菌rDNA ITS序列及共生专一性分析

春兰(Cymbidium goeringii)和蕙兰(Cymbidium faberi)是我国较为广泛分布的地生类型兰属植物,具有悠久的栽培历史和很高的经济价值.菌根真菌与兰科植物专一性关系一直是兰科茵根研究中的热点.该文对35株分离自浙江天目山野生春兰和蕙兰的菌根真菌进行了rDNA ITS序列分析,并在此基础上初步探讨了菌根真菌与春兰、蕙兰之间的专一性关系.结果表明,分离获得的菌根真菌与其共生兰属植物在种的层面具显著专一性,即物种是影响或决定浙江天目山地区春兰、蕙兰与共生菌根真菌专一性的重要因素;研究同时发现,自蕙兰同一条根分离获得的菌根真菌菌株间亦表现丰富的多样性差异.     

兰科植物内生细菌物种多样性及其促生机理研究进展

内生细菌影响兰科植物菌根形成和共生关系的稳定性,在兰科植物的生活史中起着重要作用。内生细菌通过分泌植物激素、采用光合作用、生物固氮或促进矿质营养的循环以及产生铁载体、合成其他活性物质等途径来促进兰科植物生长发育。综述了兰科植物内生细菌物种多样性的研究方法及其对兰科植物的促生机理,基于兰科植物与共生微生物的密切关系,认为内生细菌间、内生细菌与兰科植物菌根真菌间的互作是揭示兰科植物与内生细菌互作机理的重要方向。     

净月潭国家森林公园针叶林根系共生真菌初步调查

采用菌根形态学及分子生物学手段对采自净月潭国家森林公园的红松Pinus koraiensis、落叶松Larix gmelinii和云杉Picea asperata根系共生真菌进行了初步调查。结果表明:外生菌根形态依据不同树种表现不同,红松外生菌根主要表现为淡黄色到褐色,不分支、偶有珊瑚状分支,外延菌丝丰富或浓密;云杉菌根颜色基本为淡黄色到黄色,末端膨大或等粗,表现为弯曲状或蜿蜒状,不分支或不规则分支、偶有羽状分支,外延菌丝少;落叶松菌根淡黄色到黄色,偶有红褐色,不分支、羽状分支或不规则分支,外延菌丝浓密,也有表现外延菌丝少量。共鉴定获得共生真菌9科9属,其中红松根系外生菌根真菌2种:美洲乳牛肝菌Suillus americanus和外生菌根真菌未定种1,内生真菌3种:Phialocephala fortinii,Cylindrocarpon pauciseptatum和Phialocephala helvetica;云杉根系外生菌根真菌5种:臭红菇Russula foetens、外生菌根真菌未定种2和外生菌根真菌未定种3,Tomentella sp.1和Tomentella sp.2;落叶松根系外生菌根真菌4种:平凹盘菌Peziza depressa、拟篦边红菇Russula pectinatoides、外生菌根真菌未定种4和外生菌根真菌未定种5。该研究结果丰富了针叶树根系共生真菌的多样性,可为进一步开展菌根形态解剖学以及菌根生态学等研究提供依据。     

兰科植物与菌根真菌的营养关系

兰科植物与真菌具有天然的菌根共生关系。兰科植物种子细小,无胚乳,自然条件下只有与适宜的真菌共生才能萌发;兰科植物作为有花植物中最大的科之一,有光合自养、混合营养及完全真菌异养等营养类型。近年研究表明,不同营养类型的兰科植物其菌根真菌常具有一定的差异性,表现出不同的营养作用关系。本文对不同营养类型兰科植物与菌根真菌的营养作用关系的研究进展进行综述,并对兰科植物菌根营养机理、营养关系的变化等进行讨论,以期为兰科菌根营养学研究及菌根技术应用于兰科植物快繁和保育工作等提供参考。     

Generalism in the interaction of Tulasnellaceae mycobionts with orchids characterizes a biodiversity hotspot in the tropical Andes of Southern Ecuador

Biotic interactions play an important role in the assembly and stability of communities. All orchids depend on mycobionts for early establishment, but whether individual orchid species depend on a specific or broad spectrum of mycobionts is still a matter of debate. Tulasnellaceae (Basidiomycota) is the richest and most widespread mycobiont worldwide. We assessed Tulasnellaceae richness in epiphytic and terrestrial orchids in different habitats, and evaluated the degree of generalism in orchid-Tulasnellaceae interactions and the robustness of this mutualistic system to the extinction of mycobiont partners. We sampled 114 orchid individuals including all common and rare species in 56 plots of 1 m² in 3 habitats: pristine forest, regenerating forest and a landslide site in a tropical montane rainforest in Southern Ecuador. We found 52 orchid and 29 Tulasnellaceae species. The composition of Tulasnellaceae OTUs was moderately to highly similar across habitats and between orchid growth forms. A significantly nested network architecture indicated the existence of a core of generalist Tulasnellaceae OTUs interacting with both rare and common orchids. Terrestrial and epiphytic orchids showed significant differences in robustness to the extinction of their Tulasnellaceae mycobionts. Thus, generalist mycobionts may be relevant for the preservation of hyperdiverse orchid communities in the tropics.     

Phylogenetic diversity of culturable fungi associated with the Hawaiian Sponges Suberites zeteki and Gelliodes fibrosa

Sponges are well documented to harbor large amounts of microbes. Both culture-dependent and molecular approaches have revealed remarkable bacterial diversity in marine sponges. Fungi are commonly isolated from marine sponges, yet no reports on phylogenetic diversity of sponge-inhabiting fungi exist. In this report, we investigated the phylogenetic diversity of culturable fungi from the Hawaiian alien marine sponges Suberites zeteki and Gelliodes fibrosa. A total of 44 independent isolates were recovered from these two sponge species, representing 7 orders and 22 genera of Ascomycota. The majority (58%) of fungal isolates from S. zeteki resided in the Pleosporales group, while the predominant isolates (52%) from G. fibrosa were members of the Hypocreales group. Though differing in fungal species composition and structure, culturable communities of these two sponges displayed similar phylogenetic diversity. At the genus level, only two genera Penicillium and Trichoderma in the Eurotiales and Hypocreales orders, respectively, were present in both sponge species. The other genera of the fungal isolates were associated with either S. zeteki or G. fibrosa. Some of these fungal genera had been isolated from sponges collected in other marine habitats, but more than half of these genera were identified for the first time in these two marine sponges. Overall, the diversity of culturable fungal communities from these two sponge species is much higher than that observed in studies of marine sponges from other areas. This is the first report of phylogenetic diversity of marine sponge-associated fungi and adds one more dimension to our current understanding of the phylogenetic diversity of sponge-symbiotic microbes.     

Mycorrhizal specificity, preference, and plasticity of six slipper orchids from South Western China

Mycorrhizal fungi of six endangered species, Paphiopedilum micranthum, Paphiopedilum armeniacum, Paphiopedilum dianthum, Cypripedium flavum, Cypripedium guttatum, and Cypripedium tibeticum, from two closely related genera in the Orchidaceae from Southwestern China, were characterized using the nuclear internal transcribed spacer (ITS) and part of the large subunit gene of mitochondrial rDNA (mtLSU) sequences. The most frequently detected fungi belonged to the Tulasnellaceae. These fungi were represented by 25 ITS sequence types and clustered into seven major clades in the phylogenetic analysis of 5.8S sequences. Species of Paphiopedilum and Cypripedium shared no fungal ITS sequence types in common, but their fungal taxa sometimes occurred in the same major clade of the 5.8S phylogenetic tree. Although it had several associated fungal ITS sequence types in a studied plot, each orchid species had in general only a single dominant type. The fungal sequence type spectra of different species of Paphiopedilum from similar habitats sometimes overlapped; however, the dominant sequence types differed among the species and so did the sequence-type spectra within Cypripedium. Orchids of P. micranthum and P. armeniacum transplanted from the field and grown in two greenhouses had a greater number of mycorrhizal associations than those sampled directly from the field. Root specimens from P. micranthum taken from the greenhouses were preferably associated with mycobionts of the Tulasnella calospora complex, while those from the field had mycorrhizal associations of other tulasnelloid taxa. Such plasticity in mycorrhizal associations makes ex situ conservation or even propagation by means of mycorrhization of axenically grown seedlings possible.     

High diversity of root-associated fungi isolated from three epiphytic orchids in southern Ecuador

Knowledge of fungal root-associates is essential for effective conservation of tropical epiphytic orchids. We investigated the diversity of root-associated fungi of Cyrtochilum myanthum, Scaphyglottis punctulata and Stelis superbiens from a tropical mountain rainforest in southern Ecuador, using a culture dependent approach. We identified 115 fungal isolates, corresponding to 49 fungal OTUs, based on sequences of the nrDNA ITS and partial 28S region. Members of Ascomycota were unambiguously dominant (37 OTUs), including Trichoderma sp. as the most frequent taxon. Members of Basidiomycota (Agaricales and Polyporales) and Mucoromycota (Umbelopsidales and Mortierellales) were also identified. Four potential mycorrhizal OTUs of Tulasnellaceae and Ceratobasidiaceae were isolated from C. myanthum and S. superbiens. Fungal community composition was examined using Sørensen and Jaccard indices of similarity. Alfa diversity was significantly different between C. myanthum and S. superbiens. No difference in beta diversity of the fungal communities between the 3 orchid species and the collecting sites was detected. The study revealed a high diversity of fungi associated with orchid roots. Our results contribute to a better understanding of specific relationships between epiphytic orchids and their root-associated fungi.     

Extreme specificity in epiparasitic Monotropoideae (Ericaceae): widespread phylogenetic and geographical structure

The Monotropoideae (Ericaceae) are nonphotosynthetic plants that obtain fixed carbon from their fungal mycorrhizal associates. To infer the evolutionary history of this symbiosis we identified both the plant and fungal lineages involved using a molecular phylogenetic approach to screen 331 plants, representing 10 of the 12 described species. For five species no prior molecular data were available; for three species we confirmed prior studies which used limited samples; for five species all previous reports are in conflict with our results, which are supported by sequence analysis of multiple samples and are consistent with the phylogenetic patterns of host plants. The phylogenetic patterns observed indicate that: (i) each of the 13 plant phylogenetic lineages identified is specialized to a different genus or species group within five families of ectomycorrhizal Basidiomycetes; (ii) mycorrhizal specificity is correlated with phylogeny; (iii) in sympatry, there is no overlap in mature plant fungal symbionts even if the fungi and the plants are closely related; and (iv) there are geographical patterns to specificity.     

Cryptic diversity,high host specificity and reproductive synchronization in army ant‐associated Vatesus beetles

Army ants and their arthropod symbionts represent one of the most species‐rich animal associations on Earth, and constitute a fascinating example of diverse host–symbiont interaction networks. However, despite decades of research, our knowledge of army ant symbionts remains fragmentary due to taxonomic ambiguity and the inability to study army ants in the laboratory. Here, we present an integrative approach that allows us to reliably determine species boundaries, assess biodiversity, match different developmental stages and sexes, and to study the life cycles of army ant symbionts. This approach is based on a combination of community sampling, DNA barcoding, morphology and physiology. As a test case, we applied this approach to the staphylinid beetle genus Vatesus and its different Eciton army ant host species at La Selva Biological Station, Costa Rica. DNA barcoding led to the discovery of cryptic biodiversity and, in combination with extensive community sampling, revealed strict host partitioning with no overlap in host range. Using DNA barcoding, we were also able to match the larval stages of all focal Vatesus species. In combination with studies of female reproductive physiology, this allowed us to reconstruct almost the complete life cycles of the different beetle species. We show that Vatesus beetles are highly adapted to the symbiosis with army ants, in that their reproduction and larval development are synchronized with the stereotypical reproductive and behavioural cycles of their host colonies. Our approach can now be used to study army ant‐symbiont communities more broadly, and to obtain novel insights into co‐evolutionary and ecological dynamics in species‐rich host–symbiont systems.     

Ecological aspects of mycorrhizal symbiosis: with special emphasis on the functional diversity of interactions involving the extraradical mycelium

Different symbiotic mycorrhizal associations between plantsand fungi occur, almost ubiquitously, in a wide range of terrestrialecosystems. Historically, these have mainly been consideredwithin the rather narrow perspective of their effects on theuptake of dissolved mineral nutrients by individual plants.More recent research has placed emphasis on a wider, multifunctionalperspective, including the effects of mycorrhizal symbiosison plant and microbial communities, and on ecosystem processes.This includes mobilization of N and P from organic polymers,release of nutrients from mineral particles or rock surfacesvia weathering, effects on carbon cycling, interactions withmyco-heterotrophic plants, mediation of plant responses to stressfactors such as drought, soil acidification, toxic metals, andplant pathogens, as well as a range of possible interactionswith groups of other soil micro-organisms. Mycorrhizal fungiconnect their plant hosts to the heterogeneously distributednutrients required for their growth, enabling the flow of energy-richcompounds required for nutrient mobilization whilst simultaneouslyproviding conduits for the translocation of mobilized productsback to their hosts. In addition to increasing the nutrientabsorptive surface area of their host plant root systems, theextraradical mycelium of mycorrhizal fungi provides a directpathway for translocation of photosynthetically derived carbonto microsites in the soil and a large surface area for interactionwith other micro-organisms. The detailed functioning and regulationof these mycorrhizosphere processes is still poorly understoodbut recent progress is reviewed and potential benefits of improvedunderstanding of mycorrhizosphere interactions are discussed. Key words: Arbuscular mycorrhiza, biotic interactions, carbon flow, ectomycorrhiza, ericoid mycorrhiza, mycelium, nutrient uptake, symbiosis, weathering Received 22 January 2008; Revised 7 February 2008 Accepted 7 February 2008