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.     

Severe plant invasions can increase mycorrhizal fungal abundance and diversity

Invasions by non-native plants can alter ecosystem functions and reduce native plant diversity, but relatively little is known about their effect on belowground microbial communities. We show that invasions by knapweed (Centaurea stoebe) and leafy spurge (Euphorbia esula, hereafter spurge)—but not cheatgrass (Bromus tectorum)—support a higher abundance and diversity of symbiotic arbuscular mycorrhizal fungi (AMF) than multi-species native plant communities. The higher AMF richness associated with knapweed and spurge is unlikely due to a co-invasion by AMF, because a separate sampling showed that individual native forbs hosted a similar AMF abundance and richness as exotic forbs. Native grasses associated with fewer AMF taxa, which could explain the reduced AMF richness in native, grass-dominated communities. The three invasive plant species harbored distinct AMF communities, and analyses of co-occurring native and invasive plants indicate that differences were partly driven by the invasive plants and were not the result of pre-invasion conditions. Our results suggest that invasions by mycotrophic plants that replace poorer hosts can increase AMF abundance and richness. The high AMF richness in monodominant plant invasions also indicates that the proposed positive relationship between above and belowground diversity is not always strong. Finally, the disparate responses among exotic plants and consistent results between grasses and forbs suggest that AMF respond more to plant functional group than plant provenance.     

环境因子对AM真菌多样性的影响

土壤,气候和地理因子等环境因子对丝枝菌根（AM）真菌多样性有重要影响,本文系统地综述了最近10年来AM真菌生态学在该方面的最新研究成果,分析当前研究中所存在的问题和动向。     

入侵豚草与本地植物竞争对丛枝菌根真菌多样性的影响

外来植物入侵常引起入侵地丛枝菌根真菌（arbuscular mycorrhizal fungi,AMF）多样性的变化,但AMF多样性的变化与入侵域本地植物种类相关性目前尚不清楚。本研究采用野外同质园试验研究豚草与不同本地植物竞争时对AMF多样性的影响。本试验共设入侵豚草单种、本地植物狗尾草Setaria viridis单种、藜 Chenopodium album单种、黄香草木犀Melilotus officinalis单种、豚草和狗尾草混种、豚草和藜混种、豚草和黄香草木犀混种以及裸土（无任何植物生长）对照8个处理。采集根区土样后分离、鉴定AMF,分析植株生物量和AMF物种多样性变化间的关系。结果表明,对豚草而言,单种处理中生物量均高于混种处理;豚草分别与3种本地植物竞争时,与黄香草木犀混种中豚草竞争能力最强,而与藜混种中竞争能力最弱。豚草分别与3种本地植物混种时土壤中AMF种丰度和香农-威纳指数分别显著高于豚草单种和本地植物单种,其中豚草与藜混种中AMF的种丰度和香农-威纳指数最高,与狗尾草混种中AMF的种丰度和香农-威纳指数最低,而豚草分别与3种本地植物混种处理中孢子密度显著低于本地植物单种（豚草与黄香草木犀混种除外）。豚草单种、狗尾草单种、藜单种、豚草分别与狗尾草和藜混种4个处理中AMF优势种均为网状球囊霉Glomus reticulatum和缩隔球囊霉Septoglomus constrictum;黄香草木犀单种中优势种是网状球囊霉、缩隔球囊霉和根内根孢囊霉Rhizophagus Intraradices,豚草与黄香草木犀混种中优势种是网状球囊霉和根内根孢囊霉;狗尾草单种、豚草与狗尾草混种和豚草与藜混种3个处理的特有种分别是木薯根孢囊霉Rhizoglomus manihotis、近明球囊霉Claroideoglomus claroideum和美丽盾巨孢囊霉Scutellospora calospora。狗尾草单种生物量与孢子密度呈显著正相关,豚草与狗尾草混种中豚草的生物量与均匀度呈显著负相关,豚草与黄香草木犀混种中豚草的生物量与孢子密度呈显著正相关,与香农威纳指数呈显著负相关。综上可知,豚草与本地植物竞争对AMF多样性的影响具有异质性,这与本地植物种类密切相关。     

黄顶菊与3种本地植物竞争对丛枝菌根真菌种类和多样性的影响

【目的】外来植物黄顶菊对生态环境和农业经济造成了严重危害,了解黄顶菊与3种不同本地植物种植生长对丛生菌根(AM)真菌群落结构和多样性造成的影响,可以从土壤微生物角度进一步解释黄顶菊的入侵机制。【方法】通过同质园小区试验模拟黄顶菊入侵的生态进程,以黄顶菊和3种本地植物狗尾草、藜、黄香草木樨为研究对象,采用AM真菌的形态学鉴定方法,研究黄顶菊与3种本地植物不同种植方式对AM真菌群落结构和多样性的影响。【结果】(1)黄顶菊根际土壤聚集的AM真菌种类与其伴生本地植物种类有关:黄顶菊与狗尾草混种处理中优势种为网状球囊霉和根内球囊霉,而黄顶菊分别与藜、草木樨混种处理中优势种均为网状球囊霉、根内球囊霉和缩球囊霉;(2)黄顶菊分别与狗尾草和黄香草木樨混种处理中AM真菌种类既高于本地单种处理,也高于黄顶菊单种处理,说明随着黄顶菊的入侵和地上植物多样性的改变,AM真菌种类也发生改变;(3)与3种本地植物单种相比,黄顶菊各混种处理和黄顶菊单种处理中黄顶菊根际土壤根内球囊霉的重要值均增加,表明黄顶菊入侵有利于根内球囊霉的生长和发育。【结论】黄顶菊入侵改变了根际土壤AM真菌的群落结构和多样性,AM真菌的改变既与本地植物种类有关,也与入侵程度有关。     

Spatial characterization of arbuscular mycorrhizal fungal molecular diversity at the submetre scale in a temperate grassland

Although arbuscular mycorrhizal fungi (AMF) form spatially complex communities in terrestrial ecosystems, the scales at which this diversity manifests itself is poorly understood. This information is critical to the understanding of the role of AMF in plant community composition. We examined small-scale (submetre) variability of AMF community composition (terminal restriction fragment length polymorphism fingerprinting) and abundance (extraradical hyphal lengths) in two 1 m(2) plots situated in a native grassland ecosystem of western Montana. Extraradical AMF hyphal lengths varied greatly between samples (14-89 m g soil(-1)) and exhibited spatial structure at scales <30 cm. The composition of AMF communities was also found to exhibit significant spatial autocorrelation, with correlogram analyses suggesting patchiness at scales <50 cm. Supportive of overall AMF community composition analyses, individual AMF ribotypes corresponding to specific phylogenetic groups exhibited distinct spatial autocorrelation. Our results demonstrate that AMF diversity and abundance can be spatially structured at scales of <1 m. Such small-scale heterogeneity in the soil suggests that establishing seedlings may be exposed to very different, location dependent AMF communities. Our results also have direct implications for representative sampling of AMF communities in the field.     

Phylogenetic trait conservatism and the evolution of functional trade-offs in arbuscular mycorrhizal fungi

The diversity of functional and life-history traits of organisms depends on adaptation as well as the legacy of shared ancestry. Although the evolution of traits in macro-organisms is well studied, relatively little is known about character evolution in micro-organisms. Here, we surveyed an ancient and ecologically important group of microbial plant symbionts, the arbuscular mycorrhizal (AM) fungi, and tested hypotheses about the evolution of functional and life-history traits. Variation in the extent of root and soil colonization by AM fungi is constrained to a few nodes basal to the most diverse groups within the phylum, with relatively little variation associated with recent divergences. We found no evidence for a trade-off in biomass allocated to root versus soil colonization in three published glasshouse experiments; rather these traits were positively correlated. Partial support was observed for correlated evolution between fungal colonization strategies and functional benefits of the symbiosis to host plants. The evolution of increased soil colonization was positively correlated with total plant biomass and shoot phosphorus content. Although the effect of AM fungi on infection by root pathogens was phylogenetically conserved, there was no evidence for correlated evolution between the extent of AM fungal root colonization and pathogen infection. Variability in colonization strategies evolved early in the diversification of AM fungi, and we propose that these strategies were influenced by functional interactions with host plants, resulting in an evolutionary stasis resembling trait conservatism.     

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.     

喀斯特植被演替过程土壤丛枝菌根真菌(AMF)多样性

喀斯特生态系统维持了丰富的微生物多样性,丛枝菌根真菌(Arbuscular mycorrhizal fungi,AMF)结构和组成会随喀斯特植被演替而改变。以贵州贵阳花溪、毕节织金和关岭花江典型喀斯特区域按时空替代法采集了乔木林、灌木林和草本群落样地土壤,采用Illumina HiSeq分子测序技术,通过OTU聚类分析、物种注释及数据库比对,探索了喀斯特不同演替阶段土壤AMF物种多样性。结果表明:(1)喀斯特生境土壤获得球囊菌门Glomeromycota OTU为275个,分属于4目8科13属19种,属水平上AMF丰度表明根内根孢囊霉属Rhizophagus为优势属,花江拥有最高AMF丰富度,缩隔球囊霉Septoglomus constrictum、根内根孢囊霉Rhizophagus intraradices、Claroideoglomus sp. MIB8381和稀有内养囊霉Entrophospora infrequens均分布于各样地的不同植被演替阶段,为常见种。(2)AM真菌多样性Shannon指数与Simpson指数在不同演替阶段表现为花溪:乔木≈灌木草本(P0.05)、花江:灌木≈草本乔木(P0.05)、织金:乔木灌木草本,但差异不显著,Chao1和Abundance-based coverag estimator(ACE)指数表现为花江灌木≈草地乔木(P0.05)。(3)Spearman相关性分析表明土壤全磷与AMF ACE指数显著负相关,且与Chao1指数极显著负相关;速效磷与Shannon和Simpson指数显著负相关。(4)典范对应分析(Canonical Correlation Analysis,CCA)表明土壤全氮、速效氮、有机质、全磷和速效钾与AMF群落分布有显著相关性。结果表明喀斯特植被演替过程中土壤丛枝菌根真菌多样性随着演替进行或升高或降低,无一致变化规律,并与土壤理化性质关系密切,其中以磷的影响最大。     

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。     

喀斯特地区丛枝菌根真菌遗传多样性

为探明喀斯特地区丛枝菌根真菌( AMF)的遗传多样性特征,利用巢式PCR和DGGE相结合的分子生物学方法对茂兰喀斯特多个植被类型下的AMF遗传多样性进行了研究.结果表明,喀斯特地区AMF遗传多样性指数和物种丰富度分别平均为3.50和41,远高于非喀斯特对照样地的2.68和17,分析表明,喀斯特地区较高的AMF多样性与此地区丰富的植物多样性以及特殊的生态环境有关,是与喀斯特生态系统长期相互选择的结果.不同植被类型下的AMF多样性差异显著,相似性指数最高为0.34,喀斯特地区AMF的群落结构随着植被类型的改变发生显著变化;基因测序显示,喀斯特地区AMF的优势菌属是生态适应性很强的球囊霉属,在喀斯特石漠化生态恢复中具有较强的利用潜力.     

丛枝菌根真菌物种多样性研究进展

丛枝菌根(arbuscular mycorrhiza, AM)真菌是生态系统中生物多样性的重要组分之一,具有十分丰富的物种多样性、遗传多样性和功能多样性.该真菌分类地位不断提高已上升至门,下设1个纲、4个目、13个科,19个属,现已报道214种.丛枝菌根对保持生态平衡、稳定和提高生态系统可持续生产力具有重要作用.本文分析了世界范围内丛枝菌根真菌物种多样性研究现状、不同生态系统中影响丛枝菌根真菌物种多样性的关键因子及其调控途径;认为分子生物学技术是今后丛枝菌根真菌物种多样性研究的主要方法.     

Effects of long-term NP-fertilization on abundance and diversity of arbuscular mycorrhizal fungi under a maize cropping system

Diversity of arbuscular mycorrhizal fungi (AMF) in 27-year long-term NP-fertilization plots under a maize cropping system in Thailand was studied through spore morphological characterization. The plots received 0–0, 60–60, 120–120 and 180–180 kg N-P₂O₅ ha^–1 year^–1 as ammonium sulfate and triple superphosphate. The plots were sampled monthly for one year, the AMF spores were counted and morphotyped, and taxa were identified after morphotyping and monospecific pot culture. Spore number g^–1 soil, relative spore abundance and Shannon-Wiener indexes were calculated. Sixteen putative taxa were recorded from the field of which nine sporulated on maize roots in pot culture. The long-term fertilization caused decreases in AMF total spore numbers and variation in species diversity depended on sampling time. Effects of fertilization on spore number and also relative spore abundance varied with species and sampling time. Among the nine species sporulating under maize, only Acaulospora sp.1 showed no change (P > 0.003 after Bonferroni correction) in spore number with fertilization in the field; and was therefore classified as an AMF species insensitive to fertilization. Spores of Entrophospora schenckii, Glomus mosseae, Glomus sp.1, Glomus geosporum-like and Scutellospora fulgida, though they decreased in absolute numbers in response to fertilization, showed no change (P > 0.003 after Bonferroni correction) in relative abundance; these species were classified as AMF species slightly sensitive to fertilization. Three unidentified species of Glomus, though they decreased in absolute numbers in response to fertilization, showed decreases (P < 0.003 after Bonferroni correction) in relative abundance; these species were classified as AMF species highly sensitive to fertilization.     

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。     

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。     

Nitrogen and phosphorus additions impact arbuscular mycorrhizal abundance and molecular diversity in a tropical montane forest

Increased nitrogen (N) depositions expected in the future endanger the diversity and stability of ecosystems primarily limited by N, but also often co‐limited by other nutrients like phosphorus (P). In this context a nutrient manipulation experiment (NUMEX) was set up in a tropical montane rainforest in southern Ecuador, an area identified as biodiversity hotspot. We examined impacts of elevated N and P availability on arbuscular mycorrhizal fungi (AMF), a group of obligate biotrophic plant symbionts with an important role in soil nutrient cycles. We tested the hypothesis that increased nutrient availability will reduce AMF abundance, reduce species richness and shift the AMF community toward lineages previously shown to be favored by fertilized conditions. NUMEX was designed as a full factorial randomized block design. Soil cores were taken after 2 years of nutrient additions in plots located at 2000 m above sea level. Roots were extracted and intraradical AMF abundance determined microscopically; the AMF community was analyzed by 454‐pyrosequencing targeting the large subunit rDNA. We identified 74 operational taxonomic units (OTUs) with a large proportion of Diversisporales. N additions provoked a significant decrease in intraradical abundance, whereas AMF richness was reduced significantly by N and P additions, with the strongest effect in the combined treatment (39% fewer OTUs), mainly influencing rare species. We identified a differential effect on phylogenetic groups, with Diversisporales richness mainly reduced by N additions in contrast to Glomerales highly significantly affected solely by P. Regarding AMF community structure, we observed a compositional shift when analyzing presence/absence data following P additions. In conclusion, N and P additions in this ecosystem affect AMF abundance, but especially AMF species richness; these changes might influence plant community composition and productivity and by that various ecosystem processes.     

Environmental modulation of plant mycorrhizal traits in the global flora

Mycorrhizal symbioses are known to strongly influence plant performance, structure plant communities and shape ecosystem dynamics. Plant mycorrhizal traits, such as those characterising mycorrhizal type (arbuscular (AM), ecto-, ericoid or orchid mycorrhiza) and status (obligately (OM), facultatively (FM) or non-mycorrhizal) offer valuable insight into plant belowground functionality. Here, we compile available plant mycorrhizal trait information and global occurrence data ($\sim$100 million records) for 11,770 vascular plant species. Using a plant phylogenetic mega-tree and high-resolution climatic and edaphic data layers, we assess phylogenetic and environmental correlates of plant mycorrhizal traits. We find that plant mycorrhizal type is more phylogenetically conserved than plant mycorrhizal status, while environmental variables (both climatic and edaphic; notably soil texture) explain more variation in mycorrhizal status, especially FM. The previously underestimated role of environmental conditions has far-reaching implications for our understanding of ecosystem functioning under changing climatic and soil conditions.     

不同喀斯特小生境中土壤丛枝菌根真菌的遗传多样性

为探明西南喀斯特地区小生境土壤中丛枝菌根真菌(AMF)的遗传多样性特征, 利用巢式PCR和变性梯度凝胶电泳相结合的分子生物学方法, 对茂兰3种植被类型下的小生境(石缝、石沟、土面) AMF遗传多样性进行了研究。结果发现: 各类小生境都含有丰富的AMF遗传多样性, 灌木林土面的多样性指数和物种丰富度最高, 为4.06和68; 次生林石缝的最低, 为3.16和29, 所研究的9个小生境多样性指数和物种丰富度的平均值分别高达3.67和48, 高于同类研究在其他地区的结果, 这可能主要与喀斯特生态系统复杂的结构和较高的植物多样性有关; 聚类分析显示各类小生境间的AMF群落结构差异显著, 相似性指数最高仅为0.45, 说明小生境所带来的空间异质性对AMF的遗传多样性产生了显著影响; 基因测序显示球囊霉属(Glomus)极有可能是喀斯特地区AMF的优势菌属, 在以后筛选喀斯特地区的高效生态恢复菌种时可重点考虑球囊霉属的一些菌种。     

丛枝菌根真菌最新分类系统与物种多样性研究概况

丛枝菌根真菌(arbuscular mycorrhizal fungi,AMF)是自然界分布最广泛的一类植物共生真菌,能够与大部分高等植物的根系形成共生关系.由于它们在农林、环境等领域的巨大应用潜力,国内外关于AMF物种多样性的研究一直受到较高的关注.然而,AMF专性共生的特征以及研究方法不够理想等因素长期阻碍了AMF物种多样性的研究进展.近年来,研究方法的改进与新技术的应用为AMF物种多样性的研究提供了极好的机遇.简述了AMF的最新分类系统及全球物种数量、AMF物种多样性影响因素以及AMF物种多样性研究方法三个方面的研究进展,并分析了今后在AMF物种多样性相关领域值得关注的研究方向.