西北干旱区AM真菌多样性研究

为了探明我国新疆、甘肃河西走廊等西北干旱区丛枝菌根(AM)真菌的资源与分布, 我们调查了荒漠、草原、针阔叶混交林、草甸和绿洲农田5种植被类型中AM真菌类群分布特征, 以及不同生态因子对AM真菌多样性特征的影响。在西北干旱区的26科60种植物的根区土壤中, 共分离鉴定出AM真菌6属40种, 其中35个鉴定到种, 5个鉴定到属。在调查区域内球囊霉属(Glomus)为优势属, G. claroideum为优势菌种。AM真菌在不同植被类型中的发生和分布规律明显不同: 内养囊霉属(Entrophospora)只在荒漠中有分布, 类球囊霉属(Paraglomus)在绿洲农田和荒漠中有分布, 原囊霉属(Archaeospora)在草原、荒漠和绿洲农田中分布, 盾巨孢囊霉属(Scutellospora)在草甸、针阔叶混交林和草原中有分布。研究观察到在绿洲农田和草甸中AM真菌的物种多样性指数高于其他植被类型, 这反映出在干旱区土壤水分条件可能是AM真菌发生和分布的重要制约因素。     

AM真菌物种多样性:生态功能、影响因素及维持机制

AM真菌物种多样性是土壤生态系统生物多样性的重要组分之一。尽管对AM真菌多样性已有多年研究,但是,已有研究绝大多数仅停留在对AM真菌群落种属解析层面上,对AM真菌物种多样性生态功能及维持机制方面的认识较浅。从生态功能、影响因素及维持机制3个方面系统地综述了近年来AM真菌多样性领域的研究进展。认为AM真菌多样性对植物群落生产力的调控机制及结合理论与实践解析AM真菌多样性维持机制是该领域未来的重点研究方向。     

Uniting species- and community-oriented approaches to understand arbuscular mycorrhizal fungal diversity

Arbuscular mycorrhizal fungi (AMF) occur in the majority of terrestrial, and some aquatic, habitats worldwide. They are important for the functioning of individual plants and of entire ecosystems. Here, we review trends in research on species- (species recognition, phylogenetic relationships, autecology) and community-level AMF diversity patterns and aim to identify ways of improving the complementarity of these approaches. Research into many aspects of AMF diversity has flourished in parallel with the increasing availability of molecular biology techniques. However, despite their shared goal of understanding AMF biodiversity, species- and community-level perspectives, and the morphological and molecular approaches that underpin them, currently have limited capacity for information exchange. We indicate critical research gaps in AMF species and community characterisation and outline important research directions. We propose steps that could link research using different methods and targeting different aspects of diversity, in order to maximise our understanding of AMF.     

Arbuscular mycorrhizal fungal propagules in a salt marsh

The tolerance of indigenous arbuscular mycorrhizal fungi (AMF) to stressful soil conditions and the relative contribution of spores of these fungi to plant colonization were examined in a Portuguese salt marsh. Glomus geosporum is dominant in this salt marsh. Using tetrazolium as a vital stain, a high proportion of field-collected spores were found to be metabolically active at all sampling dates. Spore germination tests showed that salt marsh spores were not affected by increasing levels of salinity, in contrast to two non-marsh spore isolates, and had a significantly higher ability to germinate under increased levels of salinity (20) than in the absence of or at low salinity (10). Germination of salt marsh spores was not affected by soil water levels above field capacity, in contrast to one of the two non-marsh spore isolates. For the evaluation of infectivity, a bioassay was established with undisturbed soil cores (containing all types of AM fungal propagules) and soil cores containing only spores as AM fungal propagules. Different types of propagules were able to initiate and to expand the root colonization of a native plant species, but spores were slower than mycelium and/or root fragments in colonizing host roots. The AM fungal adaptation shown by this study may explain the maintenance of AMF in salt marshes.     

樟根生产法培育的苗与常规苗的丛枝菌根真菌多样性

以湖南省林业科学院龙伏实验基地的3年生樟根生产（root production method,RPM）苗和同龄常规技术培育的苗木（以下简称常规苗）为研究对象,采集两种苗木的根系和根际土,观察根系上菌根形态,并对其苗木样品进行高通量测序研究,探索二者根系菌根特征及根系、根际土丛枝菌根真菌（arbuscular mycorrhizal fungi,AMF）群落结构及土壤微生境差异,旨在为进行接种有效性菌根苗木来改善常规苗木的土壤微生境,进而培育高质量苗木提供理论依据。结果显示：2种樟苗木根系均显示AM菌根特征,未见外生菌根结构。测序得到属于AMF群落的有效序列共25 738条,隶属1门,2纲,3目,5科,5属,按有效序列占比从高到低依次为Acaulospora、Glomus、Diversispora、Rhizophagus和Paraglomus。根生产法（RPM）苗和常规苗的优势属均为Acaulospora和Glomus,只是RPM苗中Glomus占比高（在根、根际土内占比分别为28.04%、57.36%）而常规苗中Acaulospora占比高（在根、根际土内占比分别为99.58%、42.2%）。2种苗木均是根际土内的丰富度指数（Ace和Chao）远高于根系,并且RPM苗根际土的丰富度指数高于常规苗,但二者根系内的丰富度指数无显著差异（P<0.05）。冗余分析结果显示,土壤pH和有效磷是决定AMF群落结构的主要影响因子。不同理化因子对AMF群落的菌属影响程度也不同。     

Arbuscular mycorrhizal fungal diversity and species dominance in a temperate soil with long-term conventional and low-input cropping systems

The aim of this work was to study the effect of long-term contrasting cropping systems on the indigenous arbuscular mycorrhizal fungal (AMF) spore populations in the soil of a field experiment located in western Finland. Conventional and low-input cropping systems were compared, each with two nutrient management regimes. The conventional cropping system with a non-leguminous 6-year crop rotation (barley–barley–rye–oat–potato–oat) was fertilized at either full (rotation A) or half (rotation B) the recommended rate. In the low-input cropping system, plant residues were returned to the plots either as such (rotation C) or composted (rotation D). In the rotation of this system, 1 year with barley was replaced by clover, and oat was cultivated mixed with pea. Thus, the 6-year rotation was barley–red clover–rye–oat + pea–potato–oat + pea. Each rotation was replicated three times, starting the 6-year rotation in three different years, these being designated point 1, point 2, and point 3, respectively. In the low-input system, biotite and rock phosphate were used to compensate for K and P in the harvested yield, while animal manure was applied at the start only. After 13 years, rotation points 1 and 3 were studied. Barley was the standing crop in all plots of rotation point 1, while oat and oat + pea were grown in rotations C and D, respectively. AMF spores were studied by direct extraction and by trapping, sampled on 15 June and 15 August. In addition, a special assay was designed for isolation of fast colonizing, dominating AMF. The cropping system did not significantly affect AMF spore densities, although the low-input cropping system with composted plant residues had the highest density with 44 spores on average and the conventional system with full fertilization 24 spores per 100 cm³ soil in the autumn samples. Species richness was low in the experimental area. Five Glomus spp., one Acaulospora, and one Scutellospora were identified at the species level. In addition to these, three unidentified Glomus spp. were found. Species richness was not affected by cropping system, rotation point, or their interactions. The Shannon–Wiener index of AMF spore distributions was significantly higher in the fully fertilized than in the half-fertilized conventional plots. Glomus claroideum was the most commonly identified single species in the experimental area. It occurred in all the cropping systems and their various rotation points, representing about 30% of the total number of identified spores. In August, G. claroideum accounted for as much as 45–55% of the total numbers of spores identified in the conventional system with halved fertilization. In contrast, Glomus mosseae occurred more commonly in June (26%) than in August (9%). A bioassay using roots as inoculum for isolation and culture of dominating AMF was successfully developed and yielded only G. claroideum. This indicates a high probability of being able to more generally identify, isolate, and culture fast colonizing generalist AMF for use as inoculants in agriculture and horticulture.     

Arbuscular mycorrhizal fungal community structure and diversity in response to long-term fertilization: a field case from China

The influences of different fertilizer treatments on spore community structure and diversity of arbuscular mycorrhizal (AM) fungi (AMF) were investigated in a long-term fertilization experiment with seven treatments: organic manure (OM), half organic manure N plus half fertilizer N (1/2 OMN), fertilizer NPK, fertilizer NP, fertilizer NK, fertilizer PK, and the control (without fertilization). Fertilization generally increased the nutrient contained in the fertilizer and treatments with NPK and 1/2 OMN produced the highest crop yields. Thirty-five species of AMF within 6 genera, including 8 previously undescribed species, were recovered. Similarly in all seven treatments, the most abundant genus was Glomus, and followed by Acaulospora. All the fertilization treatments changed AM species composition, and NK treatment had the slightest influence. Fertilization with fertilizers NP, PK and NPK markedly increased AM fungal spore density, while 1/2 OMN, OM and NK treatments showed no significant influences. All the fertilizer treatments, especially OM, significantly decreased species richness and species diversity (Shannon-Weiner index). There were no significant correlations between AM fungal parameters (spore density, species richness and species diversity) and soil properties. The findings indicate that long-term fertilization all can change AM fungal community structure and decrease species diversity, while balanced fertilization with NPK or 1/2 OMN is the most suitable fertilization regime if taking both crop yields and AM species diversity into account.     

Arbuscular mycorrhizal fungal diversity in phosphorus-deficient Alfisols of a dry North-western agro-ecosystem of Tamil Nadu,India

Semi-arid tropical soils, characterized by low soil organic carbon (SOC) with limited available macronutrients and micronutrients for crop plants, are predicated to have a yield sustainability problem in the future due to intensive cropping and limited nutrient management adoptions. Arbuscular mycorrhizal fungi (AMF), the functional link between plant and soil, play a pivotal role in nutrient cycling, organic matter stabilization and soil structure and fertility improvement. Hence, so far unexplored or underutilized, native AMF could be a potential resource for fertility management of these semi-arid tropical soils. Hence, in the present investigation, we assessed the abundance and diversity of AMF in phosphorus-deficient agricultural soils of semi-arid tropics of southern India. Our results show that the spore density and infective propagules of AMF were relatively low in these soils. The morpho-typing of extracted AMF spores revealed that these soils were dominated by glomeraceae (six species of Glomus) while species of Gigaspora, Scutellospora and Acaulospora were found in low abundance. The diversity indices assessed for the AMF species were also globally low. The non-metric multi-dimensional scaling and hierarchical cluster analysis of species richness showed variation in the community composition of AMF in the soils. The principal component analysis of the assessed soil variables suggest that the available phosphorus (P), SOC and dehydrogenase and alkaline phosphatase activities had negative impact on spore density and infective propagules of AMF with no effect on species diversity. The regression analyses reveal that the available P is the significant soil variable that drives the AMF abundance and infectivity. This study opens the possibilities of effective utilization of native mycorrhizae for agriculture in semi-arid tropical soils.     

Arbuscular mycorrhizal and dark septate endophyte fungal associations in South Indian grasses

We examined arbuscular mycorrhizal (AM) and dark septate endophyte (DSE) fungal association in 50 south Indian grasses from four different sites. AM fungal diversity was also compared among the different sites. Forty-four of the 50 grasses examined had AM association and dual association with DSE fungi occurred in 25 grasses. We report for the first time AM and DSE fungal status in 23 and 27 grasses respectively. Arum-type AM morphology was the dominant occurring in 21 grasses with typical Paris-type colonization occurring in 6 grasses. AM morphology is reported for the first time in 35 grasses. Over the different sites, spore density in the soil ranged from 5–22 per 100 g air-dried soil. Spores of 11 AM fungal taxa were isolated from the soil samples of grasses of which nine belonged to Glomus, one to Acaulospora and one to Scutellospora. No significant relationship existed between AM fungal colonization and spore numbers. Species richness was high in site II and Glomus aggregatum, Glomus viscosum and Glomus mosseae were most frequent species at different sites. Overall species diversity indices (Simpson index, Shannon-Weaver index, species equitability index) differed significantly between sites.     

Arbuscular mycorrhizal fungal communities in plant roots are not random assemblages

We investigated whether arbuscular mycorrhizal fungal (AMF) communities in plant roots are random subsets of the local taxon pool or whether they reflect the action of certain community assembly rules. We studied AMF small subunit rRNA gene sequence groups in the roots of plant individuals belonging to 11 temperate forest understorey species. Empirical data were compared with null models assuming random association. Distinct fungal species pools were present in young and old successional forest. In both forest types, the richness of plant-AMF associations was lower than expected by chance, indicating a degree of partner selectivity. AMF communities were generally not characteristic of individual plant species, but those associated with ecological groups of plant species - habitat generalists and forest specialists - were nonrandom subsets of the available pool of fungal taxa and differed significantly from each other. Moreover, these AMF communities were the least distinctive in spring, but developed later in the season. Comparison with a global database showed that generalist plants tend to associate with generalist AMF. Thus, the habitat range of the host and a possible interaction with season played a role in the assembly of AMF communities in individual plant root systems.     

泰山丛枝菌根真菌群落结构特征

2007年对泰山植被根围内丛枝菌根(arbuscular mycorrhiza,AM)真菌群落组成、数量、分布及其与植物多样性的关系进行了研究。从泰山傲徕峰、黑龙潭库区等样地共分离出4属16种AM真菌:球囊霉属Glomus 9种、无梗囊霉属Acaulospora 4种、巨孢囊霉属Gigaspora 2种和盾巨孢囊霉属Scutellospora1种。其中,球囊霉属Glomus及聚球囊霉Glomus fasciculatum的孢子密度、相对多度、分布频度和重要值均最高,分别为泰山植被区根围内AM真菌优势属和优势种。各样地之间Sorenson相似系数在0.60和0.85之间。植被数量与孢子密度(r=0.80,p0.01)、植物种的丰富度与AM真菌种的丰富度(r=0.77,p0.01)以及与孢子密度(r=0.59,p0.01)均呈极显著正相关关系。研究结果表明植物多样性对于提高AM真菌多样性发挥极为重要的作用。     

Arbuscular mycorrhizal fungus identity and diversity influence subtropical tree competition

Arbuscular mycorrhizal (AM) fungi can influence plant nutrient uptake and, therefore, may alter interspecific plant competition. However, the role of AM fungi in subtropical tree competition is poorly understood. In this study, we investigated the effects of AM fungus identity (four species) and diversity (a mixture of the same four species) on the competitive relationships between seedlings of a pioneer tree Rhus chinensis and a late-pioneer tree Celtis sinensis, and between R. chinensis and a mid-successional tree Cinnamomum camphora. In seedlings, AM fungi significantly promoted a competitive advantage of R. chinensis over both Ce. sinensis and Ci. camphora. Furthermore, the extent to which AM fungi affected interspecific plant competition outcomes was dependent on AM fungus identity, and the effect of AM fungus diversity on interspecific competition outcomes may derive from the most beneficial AM fungal species.     

中国丛枝菌根真菌分子多样性

【背景】随着分子生物学技术的发展,通过分子生物学方法鉴定丛枝菌根真菌(Arbuscular mycorrhizal fungi,AMF)多样性的研究越来越多。目前,AMF在全球的分子多样性及其分布已引起关注,但在我国关于AMF分子多样性的研究进展状况尚未见报道。【目的】探究我国AMF分子多样性及其分布,研究不同气候区域以及不同生态系统中AMF分子多样性并揭示其分布规律,为我国AMF在分子水平上的研究提供基础数据。【方法】利用Davison于2015年发表于"Science"上的一篇有关全球AMF分子多样性的数据库,选用中国AMF多样性的数据,并补充了近年来已发表的该数据库中未涉及的数据,建立中国AMF分子多样性的新数据库。【结果】共得到145个分子生物学鉴定出的虚拟种(Virtualtaxa,VT),隶属于8科12属。对于不同生态系统而言,草地生态系统中AMF的分子多样性最高,占到总VT数的60.7%,人为生态系统和森林生态系统也分别达到55.2%和43.4%。同样,AMF的VT数也随气候区域的不同而有所变化,温带气候区域所占比例最高(64.1%),亚热带气候区域次之(60.7%),极北气候区域最少(20.7%)。而对于根系和土壤样品中AMF的分子多样性而言,根系样品中发现的AMF分子多样性(80.0%)高于土壤中AMF的分子多样性(48.3%)。进一步对不同生态系统、气候区域和样品来源分别在属的水平上进行研究,同样也存在一定的差异。【结论】我国具有丰富的AMF分子多样性,且不同气候区域、生态系统以及样品来源都影响着AMF的分子多样性及其分布。     

Studying fungal pathogens of humans and fungal infections: fungal diversity and diversity of approaches

Seminal work by Louis Pasteur revealed the contribution of fungi – yeasts and microsporidia to agroindustry and disease in animals, respectively. More than 150 years later, the impact of fungi on human health and beyond is an ever-increasing issue, although often underestimated. Recent studies estimate that fungal infections, especially those caused by Candida, Cryptococcus and Aspergillus species, kill more than one million people annually. Indeed, these neglected infections are in general very difficult to cure and the associated mortality remains very high even when antifungal treatments exist. The development of new antifungals and diagnostic tools that are both necessary to fight fungal diseases efficiently, requires greater insights in the biology of the fungal pathogens of humans in the context of the infection, on their epidemiology, and on their role in the human mycobiota. We also need a better understanding of the host immune responses to fungal pathogens as well as the genetic basis for the increased sensitivity of some individuals to fungal infections. Here, we highlight some recent progress made in these different areas of research, in particular based on work conducted in our own laboratories. These progresses should lay the ground for better management of fungal infections, as they provide opportunities for better diagnostic, vaccination, the development of classical antifungals but also strategies for targeting virulence factors or the host.     

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。     

Arbuscular mycorrhizal fungal abundance in elevation belts of the hyperarid Atacama Desert

Arbuscular mycorrhizal (AM) fungi affect ecosystem processes improving plant tolerance in (hyper)arid/saline environments. However, there are no previous studies on the presence of AM fungi in the Atacama Desert (Northern Chile), the driest desert in the world. Here, we studied the root and rhizosphere in 111 samples of representative plants from three elevation gradients: (i) hyperarid desert (700–2000 m a.s.l.), (ii) pre-Puna (2000–3100 m a.s.l.) and (iii) Puna (3100–4500 m a.s.l.) elevation belts. Soil pH, electrical conductivity (EC), organic matter, cations and the AM colonization and fungal structures were determined. All plants showed colonization and fungal structures. Root colonization ranged from 3.5 to 87%, hyphae showed densities from 0.13 to 204 m g⁻¹, and spore densities between 20 and 45,500 per 100 g of soil. The highest fungal structure abundances were found in Prosopis tamarugo, Baccharis scandens, Werneria pinnatifida, Deyeuxia curvula and Festuca deserticola rhizospheres. In general, EC and cations showed strong relationship with fungal structure abundance. Here, we reported for the first time the widespread presence of AM symbiosis in all the elevation belts of the Atacama Desert, constituting a first step to understand the ecological role that AM fungi play under extreme aridity and salinity conditions.     

Patterns and regulation of mycorrhizal plant and fungal diversity

The diversity of mycorrhizal fungi does not follow patterns of plant diversity, and the type of mycorrhiza may regulate plant species diversity. For instance, coniferous forests of northern latitudes may have more than 1000 species of ectomycorrhizal (EM) fungi where only a few ectomycorrhizal plant species dominate, but there are fewer than 25 species of arbuscular mycorrhizal (AM) fungi in tropical deciduous forest in Mexico with 1000 plant species. AM and EM fungi are distributed according to biome, with AM fungi predominant in arid and semiarid biomes, and EM fungi predominant in mesic biomes. In addition, AM fungi tend to be more abundant in soils of low organic matter, perhaps explaining their predominance in moist tropical forest, and EM fungi generally occur in soils with higher surface organic matter.EM fungi are relatively selective of host plant species, while AM tend to be generalists. Similar morphotypes of AM fungi collected from different sites confer different physiological benefits to the same plant species. While the EM fungi have taxonomic diversity, the AM fungi must have physiological diversity for individual species to be so widespread, as supported by existing studies. The environmental adaptations of mycorrhizal fungi are often thought to be determined by their host plant, but we suggest that the physiology and genetics of the fungi themselves, along with their responses to the plant and the environment, regulates their diversity. We observed that one AM plant species,Artemisia tridentata, was associated with different fungal species across its range, indicating that the fungi can respond to the environment directly and must not do so indirectly via the host. Different species of fungi were also active during different times of the growing season on the same host, again suggesting a direct response to the environment.These patterns suggest that even within a single functional group of microorganisms, mycorrhizal fungi, considerable diversity exists. A number of researchers have expressed the concept of functional redundancy within functional groups of microorganisms, implying that the loss of a few species would not be detectable in ecosystem functioning. However, there may be high functional diversity of AM fungi within and across habitats, and high species diversity as well for EM fungi. If one species of mycorrhizal fungus becomes extinct in a habitat, field experimental data on AM fungi suggest there may be significant shifts in how plants acquire resources and grown in that habitat.     

Arbuscular mycorrhizal and septate endophyte fungal associations in lycophytes and ferns of south India

We studied extent and type of arbuscular mycorrhizal (AM) and septate endophytic (SE) fungal associations in five lycophytes and 50 ferns collected from Eastern and Western Ghats regions. Of the 54 species and one variety (belonging to 31 genera) examined; 54 taxa had AM association and AM fungal structures were absent in Marsilea quadrifolia. This is the first report of AM and SE fungal status for 26 species each. Of the 55 taxa examined, AM morphology has been evaluated for the first time in 51 species. The hydrophytic fern Salvinia molesta was mycorrhizal and non-mycorrhizal at different sites. All the epiphytic and saxicolous species examined were mycorrhizal. The percentage of AM colonization ranged from 22.23 (Christella parasitica) to 82.20 (Adiantum lunulatum) in ferns and 53.46 (Selaginella bryopteris) to 84.34 (Selaginella sp.) in lycophytes. Epiphytic life-forms had the maximum average AM colonization levels, whereas aquatic life-forms had the minimum colonization levels. The percentage root length colonized by septate fungi ranged between 0.59 in Ophioglossum reticulatum and 16.36 in Pteris pellucida. The root length with AM and SE fungal structures as well as their total colonization significantly varied among the taxa examined. Most of the lycophytes and ferns had intermediate-type of AM morphology with a few exhibiting Paris-type. AM fungal spore numbers ranged from 1.0 (Angiopteris evecta, Pteridium aquilinum) to (Nephrolepis exaltata) 9.3 spores per 25 g soil and varied significantly among taxa. AM fungal spore morphotypes belonging to Claroideoglomus, Funneliformis, Glomus and Rhizophagus were recorded.     

Arbuscular mycorrhizal potential and mycorrhizal fungi diversity associated with Agave potatorum Zucc. in Oaxaca, Mexico

The status of the arbuscular mycorrhizal association in wild Agave potatorum Zucc. was studied at three semiarid sites over a 1-year period of rhizospheric soil sampling. Root colonization present at all sites and at all times of year, ranged from 20 to 83 %. The extraradical mycelia length was estimated to be from 2.64 to 5.22 m g^?1 of dry soil. Spore number ranged from 20 to 192 in 100 g of soil. The number of viable mycorrhizal propagules ranged from 500 to 2,640 in 100 g of soil. Twenty species of arbuscular mycorrhizal fungi (AMF) were identified, the family Glomeracea comprising the greatest number of species (45 %). Two other fungi families Acaulosporaceae and Gigasporaceae made up 35 and 15 %, respectively. High alpha diversity and low beta diversity of AMF were found in this study. Arbuscular mycorrhizal fungi species richness in A. potatorum is high and the associated fungi appear to be an important component in semiarid ecosystems in this region of Mexico. Arbuscular mycorrhizal fungi species with small spores might be better adapted to the local environment.