RIBOSOMAL RNA GENE DIVERSITY,EFFECTIVE POPULATION SIZE,AND EVOLUTIONARY LONGEVITY IN ASEXUAL GLOMEROMYCOTA

Arbuscular mycorrhizal fungi (phylum Glomeromycota) are among the oldest and most successful symbionts of land plants. With no evidence of sexual reproduction, their evolutionary success is inconsistent with the prediction that asexual taxa are vulnerable to extinction due to accumulation of deleterious mutations. To explore why Glomeromycota defy this prediction, we studied ribosomal RNA (rRNA) gene evolution in the Claroideoglomus lineage and estimated effective population size, N_e, in C. etunicatum. We found that rRNA genes of these fungi exhibit unusual and complex patterns of molecular evolution. In C. etunicatum, these patterns can be collectively explained by an unexpectedly large N_e combined with imperfect genome‐wide and population‐level rRNA gene repeat homogenization. The mutations accumulated in rRNA gene sequences indicate that natural selection is effective at purging deleterious mutations in the Claroideoglomus lineage, which is also consistent with the large N_e of C. etunicatum. We propose that in the near absence of recombination, asexual reproduction involving massively multinucleate spores typical for Glomeromycota is responsible for the improved efficacy of selection relative to drift. We postulate that large effective population sizes contribute to the evolutionary longevity of Glomeromycota.     

Evidence that soil nutrient stoichiometry controls the competitive abilities of arbuscular mycorrhizal vs. root-borne non-mycorrhizal fungi

A majority of plant species has roots that are colonized by both arbuscular mycorrhizal (AM) and non-mycorrhizal (NM) fungi. The latter group may include plant mutualists, commensals, parasites and pathogens. The co-occurrence of these two broad groups may translate into competition for root volume as well as for plant-derived carbon (C). Here we provide evidence that the relative availability of soil nitrogen (N) and phosphorus (P) (i.e., soil nutrient stoichiometry) controls the competitive balance between these two fungal guilds. A decrease in the soil available N:P ratio resulted in a lower abundance of AM fungi and a corresponding increase in NM fungi. However, when the same fertilization treatments were applied in a soil in which AM fungi were absent, lowering the soil available N:P ratio did not affect NM fungal abundance. Taken collectively, our results suggest that the increase in NM fungal abundance was not a direct response to soil nutrient stoichiometry, but rather a competitive release from AM fungi responding negatively to higher soil P. We briefly discuss the mechanisms that may be responsible for this competitive release.     

Effects of simultaneous infections of endophytic fungi and arbuscular mycorrhizal fungi on the growth of their shared host grass Achnatherum sibiricum under varying N and P supply

Grasses can be infected by endophytic fungi and arbuscular mycorrhizal (AM) fungi simultaneously. Here, we investigated the interactions of a native grass, Achnatherum sibiricum, with both endophytic and AM fungi (Glomus mosseae, GM and Glomus etunicatum, GE) at different nitrogen (N) and phosphorus (P) levels. The results showed that endophyte infection significantly suppressed the colonization rates and spore density of GE, but had no effect on those of GM. Endophyte infection increased shoot biomass regardless of the nutrient conditions. The effects of AM fungi on host growth were dependent on mycorrhizal species. There was no significant interaction between endophytic fungi and GE on host growth; however, a significant interaction between endophytic fungi and GE occurred in total phenolic concentrations and P uptake. As for GM, a significant interaction among endophytic fungi, AM fungi and nutrient availability occurred in shoot growth. Under sufficient N and P conditions, endophyte infection alleviated the detrimental effects of GM colonization on host growth.     

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.     

Shifts in symbiotic associations in plants capable of forming multiple root symbioses across a long‐term soil chronosequence

Changes in soil nutrient availability during long‐term ecosystem development influence the relative abundances of plant species with different nutrient‐acquisition strategies. These changes in strategies are observed at the community level, but whether they also occur within individual species remains unknown. Plant species forming multiple root symbioses with arbuscular mycorrhizal (AM) fungi, ectomycorrhizal (ECM) fungi, and nitrogen‐(N) fixing microorganisms provide valuable model systems to examine edaphic controls on symbioses related to nutrient acquisition, while simultaneously controlling for plant host identity. We grew two co‐occurring species, Acacia rostellifera (N₂‐fixing and dual AM and ECM symbioses) and Melaleuca systena (AM and ECM dual symbioses), in three soils of contrasting ages (c. 0.1, 1, and 120 ka) collected along a long‐term dune chronosequence in southwestern Australia. The soils differ in the type and strength of nutrient limitation, with primary productivity being limited by N (0.1 ka), co‐limited by N and phosphorus (P) (1 ka), and by P (120 ka). We hypothesized that (i) within‐species root colonization shifts from AM to ECM with increasing soil age, and that (ii) nodulation declines with increasing soil age, reflecting the shift from N to P limitation along the chronosequence. In both species, we observed a shift from AM to ECM root colonization with increasing soil age. In addition, nodulation in A. rostellifera declined with increasing soil age, consistent with a shift from N to P limitation. Shifts from AM to ECM root colonization reflect strengthening P limitation and an increasing proportion of total soil P in organic forms in older soils. This might occur because ECM fungi can access organic P via extracellular phosphatases, while AM fungi do not use organic P. Our results show that plants can shift their resource allocation to different root symbionts depending on nutrient availability during ecosystem development.     

丛枝菌根共生建成的信号识别机制

丛枝菌根(Arbuscular mycorrhiza,AM)共生是自然界中普遍存在的一种互惠共生现象,对促进土壤生态系统物质循环及维持生态系统稳定具有重要的意义。AM共生体的建立需要AM真菌和宿主植物间一系列复杂的信号识别、交换和传导。本文总结近年来相关文献,从AM共生体形成前期及AM共生体形成期两个阶段,分别综述了信号物质的生物合成过程、调控过程及其作用机制,希望有助于进一步认识AM共生体建成过程,同时通过分析当前研究工作的不足及未来研究动向,期望推动相关研究工作。     

Functional compatibility in cucumber mycorrhizas in terms of plant growth performance and foliar nutrient composition

Functional compatibility in cucumber mycorrhizas in terms of plant and fungal growth, and foliar nutrient composition from all possible combinations of six cucumber varieties and three species of arbuscular mycorrhizal (AM) fungi was evaluated. Measurements of foliar nutrient composition included N, P, K, Mg, Ca, Na, Fe, Zn, Mn and Cu. Growth of AM fungi was measured in terms of root colonisation, as examined with microscopy and the AM fungus biomarker fatty acid 16:1ω5 from both phospholipids and neutral lipids. Different responses of plant growth and foliar nutrient profiles were observed for the different AM symbioses examined. The AM fungus Claroideoglomus claroideum caused growth depression in association with four out of six cucumber varieties; Rhizophagus irregularis caused growth promotion in one of six cucumber varieties; whereas Funneliformis mosseae had no effect on the growth performance of any of the cucumber varieties examined. All three AM fungi markedly altered host plant shoot nutrient composition, with the strongest contrast observed between cucumber–R. irregularis symbioses and non‐mycorrhizal cucumber plants, independent of cucumber variety. On the other hand, AM fungal growth in roots differed between the three AM fungi, but was unaffected by host genotype. Strong build‐up of storage lipids was observed for R. irregularis, which was more moderate in the two other AM fungi. In conclusion, strong differential responses of cucumber varieties to inoculation with different AM fungi in terms of growth and shoot nutrient composition revealed high functional diversity in AM symbioses in cucumber plants.     

丛枝菌根真菌参与下植物-土壤系统的养分交流及调控

近几年随着有机农业的发展,丛枝菌根的作用受到特别关注。丛枝菌根是由植物根系与丛枝菌根真菌(AMF)形成的一种共生体。在植物-AMF-土壤系统中,AMF为植物提供N、P等营养的同时从根系得到所需的C。概述了植物-AMF-土壤系统中C、N、P等营养物质的交流以及AMF与土壤微生物的互作关系。丛枝菌根的形成可显著提高植物对P的吸收,且在高P条件下多余的P可储存于AMF中。AMF对土壤N循环的影响相当复杂,可能参与调控N循环的多个过程,如硝化作用、反硝化作用和氨氧化作用等。在有机质丰富的土壤中AMF菌丝可快速扩增并吸收其中的N,主要供菌丝自身所需,只有一小部分传递给植物。AMF对土壤C库的影响尚存争议,可能存在时间尺度的差异。短期内可活化土壤C,而在长期尺度上可能有利于土壤C的储存。AMF能够通过改变土壤微生物群落结构而影响植物-土壤体系的物质交流。AMF与解磷菌、根瘤菌和放线菌的协同增效作用可促进土壤有机质的降解或增强其固氮能力;AMF对氨氧化菌的抑制作用可降低氨的氧化减少N2O的释放。AMF与外生共生真菌EMF共存时,表现出协同增效作用,但EMF的优先定殖会限制AMF的侵染。AMF不同类群之间则主要表现为竞争和拮抗关系。AMF与土壤微生物之间的互作关系受土壤无机环境的影响,在养分亏缺条件下微生物之间往往表现为竞争关系。因植物、AMF与土壤微生物之间存在复杂的互作关系,为此AMF并不总是表现出其对植物营养的促进作用。目前关于AMF的作用机理仍以假说为主,需要进一步的实验验证。在植物-AMF-土壤系统中N与C的交流和P与C的交流并未表现出一致性,对N、P循环相互关系的进一步探讨有助于深入理解植物-土壤体系中的养分循环。植物、AMF和土壤微生物的养分来源及其对养分的相对需求强度和吸收效率尚未可知,因此无法深入理解AMF在植物-土壤体系中养分交流和转化的作用。在方法上,传统的土壤学方法在养分动态研究中存在局限性,现代分子生物学手段和化学计量学的结合值得尝试。     

浙江镜湖湿地3种菌根植物根部真菌群落多样性及对金属元素富集能力的研究

城市湿地（urban wetland）植物多样性水平高,并具有积极的净化和美化环境的作用。本研究以浙江绍兴镜湖城市湿地公园为研究样地,通过ICP-AES检测了人工园区和次生林土壤中几种金属元素（Al、Cr、Cu、K和Zn）的含量,比较不同类型的菌根植物——黑麦草（Lolium perenne L.）（AM）、六月霜（Monochasma sauatieri Franch）（AM）和乌饭树（Vaccinium bracteatum Thunb）（ERM）对金属元素的富集作用。通过高通量测序分析各宿主植物根部真菌群落的组成和结构,比较其对宿主植物金属富集能力的影响。结果表明：（1）人工园区土壤中5种金属元素的含量均显著高于次生林下的土壤;（2）乌饭树对Al的富集能力较强,黑麦草和六月霜富集Cr能力都较强;（3）3种植物根部真菌主要来自Ascomycota,而乌饭树根部Basidiomycota也是优势真菌,AM真菌在黑麦草和六月霜根部真菌群落结构中较少,而公认的ERM真菌Helotiales和Sebacinales在乌饭树根部真菌群落中的比例较高,且与所测金属元素无显著相关性。绍兴镜湖城市湿地公园次生林土壤环境保持良好。     

Soil fungal community changes in response to long-term fire cessation and N fertilization in tallgrass prairie

In grasslands, fire management and fertilization are established drivers of plant community change, but associated soil fungal responses are less well defined. We predicted that soil fungal communities would change seasonally, that decades of fire cessation and nitrogen (N) fertilization would alter fungal distributions, and that plant and fungal community change would be correlated. Surface soils were sampled monthly for 1 y from a 30-y fire by fertilization experiment to evaluate fungal community dynamics and assess correlation with plant community heterogeneity. ITS gene community composition was seasonally stable, excepting increased arbuscular mycorrhizal fungal summer abundance in the burned, fertilized treatment. Long-term treatments affected soil fungal and plant communities, with correlated heterogeneity patterns. Despite woody encroachment in the fire cessation treatment, soil fungal communities did not resemble those of forests. This study provides evidence supporting the strength of feedbacks between fungal and plant community change in response to long-term grassland fire and N management changes.