Phylogenetic and trait-based assembly of arbuscular mycorrhizal fungal communities

Both competition and environmental filtering are expected to influence the community structure of microbes, but there are few tests of the relative importance of these processes because trait data on these organisms is often difficult to obtain. Using phylogenetic and functional trait information, we tested whether arbuscular mycorrhizal (AM) fungal community composition in an old field was influenced by competitive exclusion and/or environmental filtering. Communities at the site were dominated by species from the most speciose family of AM fungi, the Glomeraceae, though species from two other lineages, the Acaulosporaceae and Gigasporaceae were also found. Despite the dominance of species from a single family, AM fungal species most frequently co-existed when they were distantly related and when they differed in the ability to colonize root space on host plants. The ability of AM fungal species to colonize soil did not influence co-existence. These results suggest that competition between closely related and functionally similar species for space on plant roots influences community assembly. Nevertheless, in a substantial minority of cases communities were phylogenetically clustered, indicating that closely related species could also co-occur, as would be expected if i) the environment restricted community membership to single functional type or ii) competition among functionally similar species was weak. Our results therefore also suggest that competition for niche space between closely related fungi is not the sole influence of mycorrhizal community structure in field situations, but may be of greater relative importance than other ecological mechanisms.     

Rice flooding negatively impacts root branching and arbuscular mycorrhizal colonization,but not fungal viability

Rice is mostly cultivated in wetlands, where arbuscular mycorrhization (AM) is reported to decrease. The mechanisms regulating such events are largely unknown. Rice uninoculated and inoculated with Rhizophagus irregularis were grown in dry and flooded conditions, allowing also for the transfer of plants from one water regime to the other. Roots were sampled at different times, from 7 to 35 d post‐inoculation (dpi). The morphological and molecular parameters (root branching, aerenchyma formation, mycorrhizal colonization, AM marker gene expression) were evaluated. Root branching was more pronounced in dry conditions, and such phenotype was enhanced by the fungus. In wetlands, the colonization level was comparable till 21 dpi, when the mycorrhization then decreased, paralleled by an increase in aerenchyma. Expression of the fungal transporters was comparable under the two conditions. The root apparatus, when shifted from one water regime to the other, rapidly adapted to the new condition, revealing a marked plasticity. The reversibility of the AM rice symbiosis was also mirrored by expression changes of plant marker genes. The results demonstrate that the water regime is the driving force that regulates AM colonization under flooding conditions, by directly influencing root architecture and anatomy, but without impacting the basic AM functionality.     

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).     

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.     

Plant traits determine the phylogenetic structure of arbuscular mycorrhizal fungal communities

Functional diversity in ecosystems has traditionally been studied using aboveground plant traits. Despite the known effect of plant traits on the microbial community composition, their effects on the microbial functional diversity are only starting to be assessed. In this study, the phylogenetic structure of arbuscular mycorrhizal (AM) fungal communities associated with plant species differing in life cycle and growth form, that is, plant life forms, was determined to unravel the effect of plant traits on the functional diversity of this fungal group. The results of the 454 pyrosequencing showed that the AM fungal community composition differed across plant life forms and this effect was dependent on the soil collection date. Plants with ruderal characteristics tended to associate with phylogenetically clustered AM fungal communities. By contrast, plants with resource‐conservative traits associated with phylogenetically overdispersed AM fungal communities. Additionally, the soil collected in different seasons yielded AM fungal communities with different phylogenetic dispersion. In summary, we found that the phylogenetic structure, and hence the functional diversity, of AM fungal communities is dependent on plant traits. This finding adds value to the use of plant traits for the evaluation of belowground ecosystem diversity, functions and processes.     

Host plant species effects on arbuscular mycorrhizal fungal communities in tallgrass prairie

Symbiotic associations between plants and arbuscular mycorrhizal (AM) fungi are ubiquitous in many herbaceous plant communities and can have large effects on these communities and ecosystem processes. The extent of species-specificity between these plant and fungal symbionts in nature is poorly known, yet reciprocal effects of the composition of plant and soil microbe communities is an important assumption of recent theoretical models of plant community structure. In grassland ecosystems, host plant species may have an important role in determining development and sporulation of AM fungi and patterns of fungal species composition and diversity. In this study, the effects of five different host plant species [Poa pratensis L., Sporobolus heterolepis (A. Gray) A. Gray, Panicum virgatum L., Baptisia bracteata Muhl. ex Ell., Solidago missouriensis Nutt.] on spore communities of AM fungi in tallgrass prairie were examined. Spore abundances and species composition of fungal communities of soil samples collected from patches within tallgrass prairie were significantly influenced by the host plant species that dominated the patch. The AM fungal spore community associated with B. bracteata showed the highest species diversity and the fungi associated with Pa. virgatum showed the lowest diversity. Results from sorghum trap cultures using soil collected from under different host plant species showed differential sporulations of AM fungal species. In addition, a greenhouse study was conducted in which different host plant species were grown in similar tallgrass prairie soil. After 4 months of growth, AM fungal species composition was significantly different beneath each host species. These results strongly suggest that AM fungi show some degree of host-specificity and are not randomly distributed in tallgrass prairie. The demonstration that host plant species composition influences AM fungal species composition provides support for current feedback models predicting strong regulatory effects of soil communities on plant community structure. Differential responses of AM fungi to host plant species may also play an important role in the regulation of species composition and diversity in AM fungal communities. Received: 29 January 1999 / Accepted: 20 October 1999     

Mechanical soil disturbance as a determinant of arbuscular mycorrhizal fungal communities in semi-natural grassland

While the effect of disturbance on overall abundance and community composition of arbuscular mycorrhizal (AM) fungi has been researched in agricultural fields, less is known about the impact in semi-natural grasslands. We sampled two AM plant species, Festuca brevipila and Plantago lanceolata, from an ongoing grassland restoration experiment that contained replicated plowed and control plots. The AM fungal community in roots was determined using nested PCR and LSU rDNA primers. We identified 38 phylotypes within the Glomeromycota, of which 29 belonged to Glomus A, six to Glomus B, and three to Diversisporaceae. Only three phylotypes were closely related to known morphospecies. Soil disturbance significantly reduced phylotype richness and changed the AM fungal community composition. Most phylotypes, even closely related ones, showed little or no overlap in their distribution and occurred in either the control or disturbed plots. We found no evidence of host preference in this system, except for one phylotype that preferentially seemed to colonize Festuca. Our results show that disturbance imposed a stronger structuring force for AM fungal communities than did host plants in this semi-natural grassland.     

Spatial variability of arbuscular mycorrhizal fungal spores in two natural plant communities

Geostatistical techniques were used to assess the spatial patterns of spores of arbuscular mycorrhizal fungi (AMF) in soils from two contrasting plant communities: a salt marsh containing only arbuscular mycorrhizal and non-mycorrhizal plants in a distinct clumped distribution pattern and a maquis with different types of mycorrhiza where most plants were relatively randomly distributed. Also evaluated was the relationship between the spatial distribution of spores and AM plant distribution and soil properties. A nested sampling scheme was applied in both sites with sample cores taken from nested grids. Spores of AMF and soil characteristics (organic matter and moisture) were quantified in each core, and core sample location was related to plant location. Semivariograms for spore density indicated strong spatial autocorrelation and a patchy distribution within both sites for all AM fungal genera found. However, the patch size differed between the two plant communities and AM fungal genera. In the salt marsh, AM fungal spore distribution was correlated with distance to AM plants and projected stand area of AM plants. In maquis, spatial AM fungal spore distribution was correlated with organic matter. These results suggest that spore distribution of AMF varied between the two plant communities according to plant distribution and soil properties.     

Changes in arbuscular mycorrhizal fungal communities along a river delta island in northeastern Brazil

Arbuscular mycorrhizal fungi (AMF) play a key role in the maintenance of the balance of terrestrial ecosystems, but little is known about the biogeography of these fungi, especially on tropical islands. This study aims to compare AMF community structure along a transect crossing a fluvial-marine island and relate these communities with soil and vegetation parameters to shed light on the forces driving AMF community structure on a local scale. We tested the hypothesis that the composition of AMF communities changes across the island, even within short distances among sites, in response to differences in edaphic characteristics and vegetation physiognomies. We sampled roots and soils in five different natural and degraded habitats: preserved mangrove forest (MF), degraded mangrove forest (MD), natural Restinga forest (RF), and two regeneration Restinga forests (RR1 and RR2) on Ilha da Restinga, northeastern Brazil. We determined the mycorrhizal colonization rate and AMF community structure based on morphological spore identification. The island soils were sandy with pH varying from acid to neutral; higher levels of organic matter were registered in RF and lower in MF; other chemical and physical soil attributes differed along the habitat types on the island. In total, 22 AMF species were identified, without any difference in species richness. However, the diversity and composition of AMF communities, spore abundance per families, and mycorrhizal colonization were statistically different among the habitats. The composition of AMF communities was strongly related to soil characteristics, especially the sum of exchangeable bases. Our results indicate that the different habitat types have diverse AMF communities even within short distances among habitats. In conclusion, islands with high spatial heterogeneity in soil parameters and diverse vegetation are potential refuges for the diversity conservation of AM fungi.     

Spatial soil heterogeneity has a greater effect on symbiotic arbuscular mycorrhizal fungal communities and plant growth than genetic modification with Bacillus thuringiensis toxin genes

Maize, genetically modified with the insect toxin genes of Bacillus thuringiensis (Bt), is widely cultivated, yet its impacts on soil organisms are poorly understood. Arbuscular mycorrhizal fungi (AMF) form symbiotic associations with plant roots and may be uniquely sensitive to genetic changes within a plant host. In this field study, the effects of nine different lines of Bt maize and their corresponding non‐Bt parental isolines were evaluated on AMF colonization and community diversity in plant roots. Plants were harvested 60 days after sowing, and data were collected on plant growth and per cent AMF colonization of roots. AMF community composition in roots was assessed using 454 pyrosequencing of the 28S rRNA genes, and spatial variation in mycorrhizal communities within replicated experimental field plots was examined. Growth responses, per cent AMF colonization of roots and AMF community diversity in roots did not differ between Bt and non‐Bt maize, but root and shoot biomass and per cent colonization by arbuscules varied by maize cultivar. Plot identity had the most significant effect on plant growth, AMF colonization and AMF community composition in roots, indicating spatial heterogeneity in the field. Mycorrhizal fungal communities in maize roots were autocorrelated within approximately 1 m, but at greater distances, AMF community composition of roots differed between plants. Our findings indicate that spatial variation and heterogeneity in the field has a greater effect on the structure of AMF communities than host plant cultivar or modification by Bt toxin genes.     

Diversity of root-associated arbuscular mycorrhizal fungal communities in a rubber tree plantation chronosequence in Northeast Thailand

Rubber tree (Hevea brasiliensis) is of major economic importance in Southeast Asia and for small land holders in Thailand in particular. Due to the high value of latex, plantations are expanding into unsuitable areas, such as the northeast province of Thailand where soil fertility is very low and therefore appropriate management practices are of primary importance. Arbuscular mycorrhizal fungi (AMF) contribute to plant growth through a range of mechanisms and could play a key role in a more sustainable management of the rubber plantations. We described the diversity of AMF associated with rubber tree roots in Northeast Thailand in relation to tree age and soil parameters along a chronosequence of rubber tree plantations. Cassava fields were included for comparison. Rubber tree and cassava roots harbored high diversity of AMF (111 Virtual Taxa, VT), including 20 novel VT. AMF VT richness per sample was consistently high (per site mean 16 to 21 VT per sample) along the chronosequence and was not related to soil properties. The composition of AMF communities differed between cassava and rubber tree plantations and was influenced by soil texture and nutrient content (sand, K, P, Ca). AMF community composition gradually shifted with the age of the trees. Our results suggest that the high diversity of AMF in this region is potentially significant for maintaining high functionality of AMF communities.     

Characterization of arbuscular mycorrhizal fungal communities with respect to zonal vegetation in a coastal dune ecosystem

Coastal dune vegetation distributes zonally along the environmental gradients of, e.g., soil disturbance. In the preset study, arbuscular mycorrhizal fungal communities in a coastal dune ecosystem were characterized with respect to tolerance to soil disturbance. Two grass species, Elymus mollis and Miscanthus sinensis, are distributed zonally in the seaward and landward slopes, respectively, in the primary dunes in Ishikari, Japan. The seaward slope is severely disturbed by wind, while the landward slope is stabilized by the thick root system of M. sinensis. The roots and rhizosphere soils of the two grasses were collected from the slopes. The soils were sieved to destruct the fungal hyphal networks, and soil trap culture was conducted to assess tolerance of the communities to disturbance, with parallel analysis of the field communities using a molecular ecological tool. In the landward communities, large shifts in the composition and increases in diversity were observed in the trap culture compared with the field, but in the seaward communities, the impact of trap culture was minimal. The landward field community was significantly nested within the landward trap culture community, implying that most members in the field community did not disappear in the trap culture. No nestedness was observed in the seaward communities. These observations suggest that disturbance-tolerant fungi have been preferentially selected in the seaward slope due to severe disturbance in the habitat. Whereas a limited number of fungi, which are not necessarily disturbance-sensitive, dominate in the stable landward slope, but high-potential diversity has been maintained in the habitat.     

The composition of arbuscular mycorrhizal fungal communities in the roots of a ruderal forb is not related to the forest fragmentation process

Land‐use changes and forest fragmentation have strong impact on biodiversity. However, little is known about the influence of new landscape configurations on arbuscular mycorrhizal fungal (AMF) community composition. We used 454 pyrosequencing to assess AMF diversity in plant roots from a fragmented forest. We detected 59 virtual taxa (VT; phylogenetically defined operational taxonomic units) of AMF – including 10 new VT – in the roots of Euphorbia acerensis. AMF communities were mainly composed of members of family Glomeraceae and were similar throughout the fragmented landscape, despite variation in forest fragment size (i.e. small, medium and large) and isolation (i.e. varying pairwise distances). AMF communities in forest fragments were phylogenetically clustered compared with the global, but not regional and local AMF taxon pools. This indicates that non‐random community assembly processes possibly related to dispersal limitation at a large scale, rather than habitat filtering or biotic interactions, may be important in structuring the AMF communities. In this system, forest fragmentation did not appear to influence AMF community composition in the roots of the ruderal plant. Whether this is true for AMF communities in soil and the roots of other ecological groups of host plants or in other habitats deserves further study.     

Heterogeneity in arbuscular mycorrhizal fungal communities may contribute to inconsistent plant-soil feedback in a Neotropical forest

Background and aims

Plant-soil feedback may vary across host species and environmental gradients. The relative importance of these biotic versus abiotic drivers of feedback will determine the stability of plant and microbial communities across environments. If plant hosts are the main driver of soil microbial communities, plant-soil feedback may be stable across changing environments. However, if microbial communities vary with environmental gradients, feedback may also vary, limiting its capacity to predict plant distributions.

Methods

We characterized arbuscular mycorrhizal (AM) fungi across tree plantations and a primary Neotropical rainforest. We then performed a plant-soil feedback pot experiment of AM fungi from these plantations on three plant species and related feedback and AM fungal communities in the field.

Results

In the field, temporal and spatial variation in AM fungal composition was similar in magnitude to variation across plant host species. Composition of AM fungi in the pot experiment significantly differed from the field plots. Furthermore, differential feedback was explained by shifts in AM fungal composition only for one plant host species (Hyeronima alchorneoides) in the pot experiment.

Conclusions

Natural AM fungal communities were temporally and spatially heterogeneous and AM fungal communities in the greenhouse did not reflect natural soils. These factors led to heterogeneous and unpredictable feedback responses, which suggests that applying greenhouse derived plant-soil feedback trends to predict plant coexistence in natural systems may be misleading.

     

减磷配施有机肥对丛枝菌根真菌群落的复杂度和稳定性的短期效应

【目的】探究减磷配施有机肥条件下土壤中丛枝菌根（arbuscular mycorrhiza,AM）真菌群落特性、网络复杂性及群落的稳定性之间的关系,揭示有机肥替代背景下,土壤理化性质与AM真菌的群落结构对网络特征和群落稳定性的短期效应。【方法】在2012年开始的无机磷肥长期定位试验的基础上,于2018年实施减磷配施有机肥裂区试验,共设6个处理：施无机磷0、75、150 kg/hm²;无机磷肥施用量减少30%,即0、52.5、105 kg/hm²,并配施有机肥（猪粪）3 187 kg/hm²,每个处理重复3次。通过高通量测序和生物信息学分析,探究减磷配施有机肥对土壤中AM真菌群落的网络特征及稳定性的短期效应。【结果】相比于无机磷施用,减磷配施有机肥整体上降低了AM真菌群落的α多样性,各处理中的AM真菌优势类群均为球囊霉属（Glomus）和类球囊霉属（Paraglomus）。网络的平均度、平均加权度在无机磷肥及减磷配施有机肥处理中均在适量施磷下达到最大值,且无机磷肥处理大于减磷配施有机肥处理;网络负相关连接线数在无机磷肥处理中随施磷量增加而增加,而在减磷配施有机肥处理中随施磷量增加而减少。减磷配施有机肥通过抑制AM真菌群落间正相互作用来提高负正凝聚力比值,从而促进群落稳定性。相比于AM真菌的指示物种（indicator species）和关键类群（keystone taxa）,优势类群（dominant taxa）与AM真菌群落的稳定性密切相关。【结论】在酸性紫色土中,短期减磷配施有机肥通过改变土壤pH、速效磷和有机质,调控AM真菌群落的α多样性和优势类群,进而影响AM真菌群落的网络复杂度和群落稳定性。     

长期肥料施用对华北大田玉米土壤丛枝菌根真菌群落结构的影响

华北平原是我国北方最重要的粮食主产区之一,过去30年间其粮食产量稳定提高的同时,肥料等生产资料的投入也大幅增加。丛枝菌根(Arbuscular mycorrhiza,AM)真菌作为一类重要的土壤功能性微生物,研究施肥措施对其群落、功能的影响,在挖掘农田有益微生物的潜在作用方面具有重要意义。本研究采用分子生物学方法测定了华北平原典型集约化农田体系中不同养分用量和投入模式对土壤AM真菌群落的影响。试验样地于2007年开始对冬小麦-夏玉米轮作体系进行施肥处理:有机肥(牛粪9 000kg/hm~2)、高量化肥(296.4kg N/hm~2、177.3kg P/hm~2、287.8kg K/hm~2,较当地常规化肥施用量上调25%)、低量化肥(74.1kg N/hm~2、44.3kg P/hm~2、71.9kg K/hm~2,较当地常规化肥施用量下调75%)和不施肥对照,每个处理3次重复,于2011年10月玉米收获季进行土壤取样。试验获得的31个OTUs主要属于Glomerales目(球囊霉目)(Glomus Group A/B)及Diversisporales目(多孢囊霉目),其中部分AM真菌类群如Glo12、Aca3、Scutellospora aurigloba(Scut.aurigloba全球盾巨孢囊霉)以及Div2仅在有机肥处理中被检测到。与其他处理相比,有机肥处理的116bp片段(Glo9/12/13/14;Fun1/2;Rhi1)相对丰度显著升高,而140bp片段(Uncultured Glomus)(球囊霉属)则主要受到化肥处理的促进。对于Diversisporales目AM真菌所对应的片段(141、142、169bp),施肥明显对其有促进作用。4年的肥料施用对土壤AM真菌物种多样性无显著影响,施肥处理0–20、20–40、40–60cm深度土壤AM真菌群落的多样性水平分别为1.14–1.78、1.61–1.84、1.23–2.07,丰富度分别为3.7–7.5、8.0–8.3、5.0–7.3。表明深层土壤是AM真菌重要的库,多样性水平并不低,对于维持土壤微生物多样性水平具有重要作用。本研究发现,p H值是影响表层土壤AM真菌群落分布的主要原因,土壤全氮、全碳、速效磷等肥力因子也是影响表层土壤AM真菌群落分布的重要因子,而深层土壤AM真菌的群落组成与固有的盐分特征(土壤电导率EC)有着较好的相关性。