The influence of arbuscular mycorrhizae on the relationship between plant diversity and productivity

Ecological theory predicts a positive and asymptotic relationship between plant diversity and ecosystem productivity based on the ability of more diverse plant communities to use limiting resources more fully. This is supported by recent empirical evidence. Additionally, in natural ecosystems, plant productivity is often a function of the presence and composition of mycorrhizal associations. Yet, the effect of mycorrhizal fungi on the relationship between plant diversity and productivity has not been investigated. We predict that in the presence of AMF, productivity will saturate at lower levels of species richness because AMF increase the ability of plant species to utilize nutrient resources. In this study we manipulated old-field plant species richness in the presence and absence of two species of AMF. We found that in the absence of AMF, the relationship between plant species richness and productivity is positive and linear. However, in the presence of AMF, the relationship is positive but asymptotic, even though the maximum plant biomass was significantly different between the two AMF treatments. This is consistent with the hypothesis that AMF increase the redundancy of plant species in the productivity of plant communities, and indicates that these symbionts must be considered in future investigations of plant biodiversity and ecosystem function.     

Shifting the balance from qualitative to quantitative analysis of arbuscular mycorrhizal communities in field soils

Population studies of arbuscular mycorrhizal fungi (AMF) have traditionally been achieved by indirect analyses of soil-borne spore populations. These studies are not necessarily reflective of populations of AMF within the roots. Advances in molecular biology have revolutionized the analysis of fungal populations colonizing roots and forming mycorrhizas. Initially these studies were qualitative and reported presence or absence of particular AMF species in soils or in roots for comparison between different environments. More recently, the methodology has developed for direct quantification of AMF within roots. Quantitative PCR provides the means to study spatial distribution and individual quantification of AMF in mixed communities over time. In this review, we discuss the progress and application of indirect, direct and finally quantitative methodologies for studying arbuscular mycorrhizal communities. We conclude that the molecular tools now exist to quantitatively analyse the effect of environment, management or inoculation of soils on AMF communities within roots.     

Seasonal dynamics of arbuscular mycorrhizal fungal communities in roots in a seminatural grassland

Symbiotic arbuscular mycorrhizal fungi (AMF) have been shown to influence both the diversity and productivity of grassland plant communities. These effects have been postulated to depend on the differential effects of individual mycorrhizal taxa on different plant species; however, so far there are few detailed studies of the dynamics of AMF colonization of different plant species. In this study, we characterized the communities of AMF colonizing the roots of two plant species, Prunella vulgaris and Antennaria dioica, in a Swedish seminatural grassland at different times of the year. The AMF small subunit rRNA genes were subjected to PCR, cloning, sequencing, and phylogenetic analysis. Nineteen discrete sequence types belonging to Glomus groups A and B and to the genus Acaulospora were distinguished. No significant seasonal changes in the species compositions of the AMF communities as a whole were observed. However, the two plant species hosted significantly different AMF communities. P. vulgaris hosted a rich AMF community throughout the entire growing season. The presence of AMF in A. dioica decreased dramatically in autumn, while an increased presence of Ascomycetes species was detected.     

中国大陆地区丛枝菌根真菌菌种资源的分离鉴定与形态学特征

【目的】分离收集保藏中国大陆各个地区不同生态环境的丛枝菌根真菌菌种资源,为丛枝菌根的研究提供资源、奠定基础。【方法】以高粱为宿主植物,采用诱导培养、单孢培养和扩繁培养分离土壤样品中的丛枝菌根真菌菌种并鉴定。【结果】从我国大陆的45个地区50余种宿主植物根区土壤中分离到丛枝菌根真菌135株,隶属于23个种;对各个菌株的形态特征进行了描述。【结论】我国蕴藏着丰富的丛枝菌根真菌菌种资源,文中描述的菌种资源是目前从我国大陆地区获得的种类和数量最多、覆盖范围最广的AM真菌菌种资源。     

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。     

Seasonal Dynamics of Arbuscular Mycorrhizal Fungal Communities in Roots in a Seminatural Grassland

Symbiotic arbuscular mycorrhizal fungi (AMF) have been shown to influence both the diversity and productivity of grassland plant communities. These effects have been postulated to depend on the differential effects of individual mycorrhizal taxa on different plant species; however, so far there are few detailed studies of the dynamics of AMF colonization of different plant species. In this study, we characterized the communities of AMF colonizing the roots of two plant species, Prunella vulgaris and Antennaria dioica, in a Swedish seminatural grassland at different times of the year. The AMF small subunit rRNA genes were subjected to PCR, cloning, sequencing, and phylogenetic analysis. Nineteen discrete sequence types belonging to Glomus groups A and B and to the genus Acaulospora were distinguished. No significant seasonal changes in the species compositions of the AMF communities as a whole were observed. However, the two plant species hosted significantly different AMF communities. P. vulgaris hosted a rich AMF community throughout the entire growing season. The presence of AMF in A. dioica decreased dramatically in autumn, while an increased presence of Ascomycetes species was detected.     

Host identity and functional traits determine the community composition of the arbuscular mycorrhizal fungi in facultative epiphytic plant species

The epiphytic vascular mycobiota is scarce and facultative in semi-arid Mediterranean ecosystems. However, unlike in soil conditions, little is known about the factors driving mycorrhizal communities in epiphytic environments. Here, we investigated the arbuscular mycorrhizal fungi (AMF) harboured by 31 plant species occurring on the trunks of Phoenix dactylifera. We wanted to ascertain if host identity and plant functional traits shape mycorrhizal communities. Specifically, we tested the plant life-cycle (perennial versus annual), the plant life-form (herbaceous versus woody), the plant origin (exotic versus native) and the plant species. The plant affiliation to species strongly influenced the AMF community composition. Plant life-form and plant life-cycle also shaped indicator taxa. The AMF structure differed between annual and perennial species and higher AMF richness was detected in perennial plants. The epiphytic plants associated with AMF irrespective of whether they were native or not, probably because here no functional differences derive from plant origin.     

Host preferences of arbuscular mycorrhizal fungi colonizing annual herbaceous plant species in semiarid mediterranean prairies

In this study, we have analyzed and compared the diversities of the arbuscular mycorrhizal fungi (AMF) colonizing the roots of five annual herbaceous species (Hieracium vulgare, Stipa capensis, Anagallis arvensis, Carduus tenuiflorus, and Avena barbata) and a perennial herbaceous species (Brachypodium retusum). Our goal was to determine the differences in the communities of the AMF among these six plant species belonging to different families, using B. retusum as a reference. The AMF small-subunit rRNA genes (SSU) were subjected to nested PCR, cloning, sequencing, and phylogenetic analysis. Thirty-six AMF phylotypes, belonging to Glomus group A, Glomus group B, Diversispora, Paraglomus, and Ambispora, were identified. Five sequence groups identified in this study clustered to known glomalean species or isolates: group Glomus G27 to Glomus intraradices, group Glomus G19 to Glomus iranicum, group Glomus G10 to Glomus mosseae, group Glomus G1 to Glomus lamellosum/etunicatum/luteum, and group Ambispora 1 to Ambispora fennica. The six plant species studied hosted different AMF communities. A certain trend of AMF specificity was observed when grouping plant species by taxonomic families, highlighting the importance of protecting and even promoting the native annual vegetation in order to maintain the biodiversity and productivity of these extreme ecosystems.     

High compatibility between arbuscular mycorrhizal fungal communities and seedlings of different land use types in a tropical dry ecosystem

We conducted this study to explore limitations for the establishment of mycorrhizal associations in disturbed areas of the tropical dry ecosystem in the Chamela region of Jalisco, Mexico. Specifically, we: (1) assessed the diversity and composition of arbuscular mycorrhizal fungal (AMF) communities through spore morphospecies identification in three common land uses (primary forest, secondary forest, and pasture), (2) tested the inoculum potential of the AMF communities and the effect of water stress on the establishment of mycorrhizal associations in seedlings of various plant species, and (3) explored the importance of AMF community composition on early seedling development. Soil and root samples were taken from 15 random points in each of three plots established in two primary forests, two 26-year-old secondary forests, and two 26-year-old pastures. We expected that because of soil degradation and management, pastures would have the lowest and primary forests the highest AMF species richness. We found evidence for changes in AMF species composition due to land use and for higher morphospecies richness in primary forests than in secondary forests and pastures. We expected also that water stress limited plant and mycorrhizal development and that plants and AMF communities from secondary forests and pastures would be less affected by (better adapted to) water stress than those from the primary forest. We found that although all plant species showed biomass reductions under water stress, only some of the plant species had lower mycorrhizal development under water stress, and this was regardless of the AMF community inoculated. The third hypothesis was that plant species common to all land use types would respond similarly to all AMF communities, whereas plant species found mainly in one land use type would grow better when inoculated with the AMF community of that specific land use type. All plant species were however equally responsive to the three AMF communities inoculated, indicating that all plants established functionally compatible AMF in each community, with no preferences. The results suggest that early seedling growth and mycorrhizal development in secondary forests and pastures is not likely limited by diversity, quantity, or quality of mycorrhizal propagules but by the high temperature and water stress conditions prevailing at those sites.     

丛枝菌根真菌（AMF）对外来植物入侵反馈机制的研究进展

丛枝菌根真菌（AMF）在植物群落竞争演替、物种多样性的形成及群落空间分布格局、植物群落对全球变化的响应中均起着重要的调节作用;同样也能影响外来植物与本地植物的互作,影响外来植物入侵过程中植物群落演替进程,甚至决定入侵的成败。因此,AMF与外来植物共生及其对外来植物入侵的反馈已成为国际上外来植物入侵机制研究的一个热点。本文基于外来植物的入侵过程,从AMF对外来植物生长、外来植物与本地植物竞争关系的影响,以及外来植物入侵对AMF的影响及AMF对入侵的反馈3个方面综述了AMF对外来植物入侵的反馈机制。外来植物可以通过多种途径改变土著AMF的群落结构和功能,而土著AMF也能直接或间接地改变甚至逆转外来植物与入侵地植物的互作关系。未来的研究不仅需要考虑AMF与外来植物共生的菌根特性和对竞争关系的影响,还需要通过大尺度条件下的野外试验及室内补充试验深入探究影响AMF在外来植物与本地植物竞争演替中的作用的生物和非生物因子,以全面解释AMF影响外来植物入侵的反馈机制。     

Disclosing arbuscular mycorrhizal fungal biodiversity in soil through a land-use gradient using a pyrosequencing approach

The biodiversity of arbuscular mycorrhizal fungi (AMF) communities present in five Sardinian soils (Italy) subjected to different land-use (tilled vineyard, covered vineyard, pasture, managed meadow and cork-oak formation) was analysed using a pyrosequencing-based approach for the first time. Two regions of the 18S ribosomal RNA gene were considered as molecular target. The pyrosequencing produced a total of 10924 sequences: 6799 from the first and 4125 from the second target region. Among these sequences, 3189 and 1003 were selected to generate operational taxonomic units (OTUs) and to evaluate the AMF community richness and similarity: 117 (37 of which were singletons) and 28 (nine of which were singletons) unique AMF OTUs were detected respectively. Within the Glomeromycota OTUs, those belonging to the Glomerales order were dominant in all the soils. Diversisporales OTUs were always detected, even though less frequently, while Archaeosporales and Paraglomerales OTUs were exclusive of the pasture soil. Eleven OTUs were shared by all the soils, but each of the five AMF communities showed particular features, suggesting a meaningful dissimilarity among the Glomeromycota populations. The environments with low inputs (pasture and covered vineyard) showed a higher AMF biodiversity than those subjected to human input (managed meadow and tilled vineyard). A reduction in AMF was found in the cork-oak formation because other mycorrhizal fungal species, more likely associated to trees and shrubs, were detected. These findings reinforce the view that AMF biodiversity is influenced by both human input and ecological traits, illustrating a gradient of AMF communities which mirror the land-use gradient. The high number of sequences obtained by the pyrosequencing strategy has provided detailed information on the soil AMF assemblages, thus offering a source of light to shine on this crucial soil microbial group.     

来源对丛枝菌根真菌功能的影响

丛枝菌根真菌(arbuscular mycorrhizal fungi,AMF)是一种具有重要生态功能的根际微生物。影响AMF功能的因素主要包括AMF、宿主,以及其它生物与非生物环境。近年来,大量研究表明不同来源的AMF功能存在显著差异。结合该领域的最新研究成果,从基因变异和表型可塑性两方面分析了来源影响AMF功能的原因和维持机制,并对其生态学意义进行了探讨。认为应从群落水平上研究不同来源的AMF在生态系统中的功能,重视宿主的作用,保持研究对象的对等性,并区分AMF群落结构和来源对功能的不同影响。对未来的研究重点和方向进行了讨论和展望,试图为更加清晰、全面地认识AMF在生态系统中的功能,以及AMF适应特定土壤环境与宿主的机制提供参考。     

Effects of Alliaria petiolata (garlic mustard; Brassicaceae) on mycorrhizal colonization and community structure in three herbaceous plants in a mixed deciduous forest

Herbaceous plant species are important components of forest ecosystems, and their persistence in forests may be affected by invasive plant species that reduce mycorrhizal colonization of plant roots. I examined the effect of the invasive plant Alliaria petiolata on arbuscular mycorrhizal fungi (AMF) colonizing the roots of three forest plant species. AMF root colonization and community structure was examined from plants that were growing either in the absence or presence of Alliaria under natural forest conditions. AMF root colonization varied among the plant species but was not significantly affected by Alliaria. With molecular methods, ～12 different taxa of AMF could be distinguished among the root samples, and these taxa belonged to the genera Acaulospora and Glomus, with Glomus dominating AMF communities. There were significant differences between the community of AMF colonizing roots of Maianthemum racemosum and Trillium grandiflorum, but only AMF communities of Maianthemum roots were significantly affected by Alliaria. Indicator species analysis found that an Acaulospora species type was a significant indicator of Maianthemum plants grown in the absence of Alliaria. These results suggest invasive plants like Alliaria may selectively suppress AMF fungi, and this suppression can affect AMF communities colonizing the roots of some native plant species.     

Diverse plant mixtures sustain a greater arbuscular mycorrhizal fungi spore viability than monocultures after 12 years

密集的土地管理可能损害土壤生物多样性,从而危害长期的土壤生产力。丛枝菌根真菌(AMF)通过与植物专一性的共生关系,在促进土壤生产力方面发挥着关键作用。然而,目前尚不清楚植物群落的性质,特别是植物物种丰富度和组成如何影响土壤中AMF种群的生存能力。本研究中,我们测试了来自不同植物功能类群的8种植物的单一栽培和不同植物的混合种植,是否可以维持更有活力的AMF繁殖体。为了回答这个问题,我们从12年的单种植物和混种植物中提取AMF孢子,并通过因子设计将单株AMF孢子与单株植物配对,并考察与AMF孢子来源和植物物种特性的交互作用。研究结果表明,不同植物混合种植的AMF孢子比单一栽培的AMF孢子更能成功地定植于多种植物和植株个体中。此外,我们还发现,来自不同混合种植的AMF孢子比来自单一栽培的AMF更能显著提高白车轴草(Trifolium repens L.)的生物量。 与单一栽培相比,起源于不同植物混合种植的AMF孢子的生存能力和与多种植物相互作用的能力更强。我们的研究结果首次表明,多样的植物群落可以维持土壤中AMF的活力,也证明了多样的植物群落有潜力维持AMF繁殖体的活力对土壤健康和生产力至关重要。     

Evidence for the evolution of native plant response to mycorrhizal fungi in post‐agricultural grasslands

Plant–microbe interactions play an important role in structuring plant communities. Arbuscular mycorrhizal fungi (AMF) are particularly important. Nonetheless, increasing anthropogenic disturbance will lead to novel plant–AMF interactions, altering longstanding co‐evolutionary trajectories between plants and their associated AMF. Although emerging work shows that plant–AMF response can evolve over relatively short time scales due to anthropogenic change, little work has evaluated how plant AMF response specificity may evolve due to novel plant–mycorrhizal interactions. Here, we examine changes in plant–AMF interactions in novel grassland systems by comparing the mycorrhizal response of plant populations from unplowed native prairies with populations from post‐agricultural grasslands to inoculation with both native prairie AMF and non‐native novel AMF. Across four plant species, we find support for evolution of differential responses to mycorrhizal inocula types, that is, mycorrhizal response specificity, consistent with expectations of local adaptation, with plants from native populations responding most to native AMF and plants from post‐agricultural populations responding most to non‐native AMF. We also find evidence of evolution of mycorrhizal response in two of the four plant species, as overall responsiveness to AMF changed from native to post‐agricultural populations. Finally, across all four plant species, roots from native prairie populations had lower levels of mycorrhizal colonization than those of post‐agricultural populations. Our results report on one of the first multispecies assessment of local adaptation to AMF. The consistency of the responses in our experiment among four species provides evidence that anthropogenic disturbance may have unintended impacts on native plant species'' association with AMF, causing evolutionary change in the benefit native plant species gain from native symbioses.     

Exploring the symbiont diversity of ancient western redcedars: arbuscular mycorrhizal fungi of long‐lived hosts

Arbuscular mycorrhizal fungi (AMF) are globally distributed, monophyletic root symbionts with ancient origins. Their contribution to carbon cycling and nutrient dynamics is ecologically important, given their obligate association with over 70% of vascular plant species. Current understanding of AMF species richness and community structure is based primarily on studies of grasses, herbs and agricultural crops, typically in disturbed environments. Few studies have considered AMF interactions with long‐lived woody perennial species in undisturbed ecosystems. Here we examined AMF communities associated with roots and soils of young, mature and old western redcedar (Thuja plicata) at two sites in the old‐growth temperate rainforests of British Columbia. Due to the unique biology of AMF, community richness and structure were assessed using a conservative, clade‐based approach. We found 91 AMF OTUs across all samples, with significantly greater AMF richness in the southern site, but no differences in richness along the host chronosequence at either site. All host age classes harboured AMF communities that were overdispersed (more different to each other than expected by chance), with young tree communities most resembling old tree communities. A comparison with similar clade richness data obtained from the literature indicates that western redcedar AMF communities are as rich as those of grasses, tropical trees and palms. Our examination of undisturbed temperate old‐growth rainforests suggests that priority effects, rather than succession, are an important aspect of AMF community assembly in this ecosystem.     

Plant–mycorrhizal fungus co‐occurrence network lacks substantial structure

The interactions between plants and arbuscular mycorrhizal fungi (AMF) maintain a crucial link between macroscopic organisms and the soil microbial world. These interactions are of extreme importance for the diversity of plant communities and ecosystem functioning. Despite this importance, only recently has the structure of plant–AMF interaction networks been studied. These recent studies, which used genetic data, suggest that these networks are highly structured, very similar to plant–animal mutualistic networks. However, the assembly process of plant–AMF communities is still largely unknown, and an important feature of plant–AMF interactions has not been incorporated: they occur at an extremely localized scale. Studying plant–AMF networks in a spatial context seems therefore a crucial step. This paper studies a plant–AMF spatial co‐occurrence network using novel methodology based on information theory and a unique set of spatially explicit species‐level data. We apply three null models of which only one accounts for spatial effects. We find that the data show substantial departures from null expectations for the two non‐spatial null models. However, for the null model considering spatial effects, there are few significant co‐occurrences compared with the other two null models. Thus, plant–AMF spatial co‐occurrences seem to be mostly explained by stochasticity, with a small role for other factors related to plant–AMF specialization. Furthermore, we find that the network is not significantly nested or modular. We conclude that this plant–AMF spatial co‐occurrence network lacks substantial structure and, therefore, plants and AMF species do not track each other over space. Thus, random encounters seem more important in the first step of the assembly of plant–AMF communities. Synthesis The symbiotic interaction between plants and arbuscular mycorrhizal fungi (AMF) is crucial for ecosystem functioning. However, the factors affecting the assembly of plant‐AMF communities are poorly understood. An important factor of the assembly of plant‐AMF communities has been overlooked: plant‐AMF interactions occur at a localized spatial scale. Our study investigated the importance of space in the structure of plant‐AMF communities. We studied a plant‐AMF spatial co‐occurrence network using a unique set of spatially explicit data and applied three null models. We found that plant‐AMF spatial co‐occurrences seem to be mostly explained by stochasticity. In particular, our study shows that this plant‐AMF spatial co‐occurrence network lacks substantial structure and, therefore, plants and AMF species do not track each other over space. Thus, random encounters seem to drive the assembly of plant‐AMF communities.     

The role of local environment and geographical distance in determining community composition of arbuscular mycorrhizal fungi at the landscape scale

Arbuscular fungi have a major role in directing the functioning of terrestrial ecosystems yet little is known about their biogeographical distribution. The Baas-Becking hypothesis (‘everything is everywhere, but, the environment selects'') was tested by investigating the distribution of arbuscular mycorrhizal fungi (AMF) at the landscape scale and the influence of environmental factors and geographical distance in determining community composition. AMF communities in Trifolium repens and Lolium perenne roots were assessed in 40 geographically dispersed sites in Ireland representing different land uses and soil types. Field sampling and laboratory bioassays were used, with AMF communities characterised using 18S rRNA terminal-restriction fragment length polymorphism. Landscape-scale distribution of AMF was driven by the local environment. AMF community composition was influenced by abiotic variables (pH, rainfall and soil type), but not land use or geographical distance. Trifolium repens and L. perenne supported contrasting communities of AMF, and the communities colonising each plant species were consistent across pasture habitats and over distance. Furthermore, L. perenne AMF communities grouped by soil type within pasture habitats. This is the largest and most comprehensive study that has investigated the landscape-scale distribution of AMF. Our findings support the Baas-Becking hypothesis at the landscape scale and demonstrate the strong influence the local environment has on determining AMF community composition.     

Nonlegumes, legumes, and root nodules harbor different arbuscular mycorrhizal fungal communities

Legumes are an important plant functional group since they can form a tripartite symbiosis with nitrogen-fixing Rhizobium bacteria and phosphorus-acquiring arbuscular mycorrhizal fungi (AMF). However, not much is known about AMF community composition in legumes and their root nodules. In this study, we analyzed the AMF community composition in the roots of three nonlegumes and in the roots and root nodules of three legumes growing in a natural dune grassland. We amplified a portion of the small-subunit ribosomal DNA and analyzed it by using restriction fragment length polymorphism and direct sequencing. We found differences in AMF communities between legumes and nonlegumes and between legume roots and root nodules. Different plant species also contained different AMF communities, with different AMF diversity. One AMF sequence type was much more abundant in legumes than in nonlegumes (39 and 13%, respectively). Root nodules contained characteristic AMF communities that were different from those in legume roots, even though the communities were similar in nodules from different legume species. One AMF sequence type was found almost exclusively in root nodules. Legumes and root nodules have relatively high nitrogen concentrations and high phosphorus demands. Accordingly, the presence of legume- and nodule-related AMF can be explained by the specific nutritional requirements of legumes or by host-specific interactions among legumes, root nodules, and AMF. In summary, we found that AMF communities vary between plant functional groups (legumes and nonlegumes), between plant species, and between parts of a root system (roots and root nodules).     

陆地农业生态系统丛枝菌根真菌物种多样性研究进展

丛枝菌根真菌(AMF)是一种古老的、在自然界中普遍存在的土壤微生物,能与大部分陆生植物形成互惠互利的菌根共生体.在这种共生关系中,AMF从植物获取自身生长所需碳源的同时,帮助宿主吸收氮、磷等营养物质.AMF在农业生态系统中具有重要作用,能够促进植物生长、改善作物品质、提高植物抗逆性、稳定土壤结构、维护生态平衡和维持农业可持续发展.本文总结了近几年来陆地农业生态系统AMF的研究进展,着重从我国陆地农业生态系统AMF物种多样性、AMF生物多样性时空分布特征及影响AMF多样性的因素等几个方面,综述了陆地农业生态系统AMF的物种多样性,并对以后的研究进行了展望.