Interactions between an arbuscular mycorrhizal fungus and a soil microbial community mediating litter decomposition

We investigated arbuscular mycorrhizal fungi (AMF) alteration of microbial mediation of litter decomposition. AMF (Glomus hoi) were either allowed access to or excluded from Plantago lanceolata L. root litter embedded in soil; litter was labeled with either (13) C only or (13) C and (15) N. Plant N uptake was significantly increased if AMF accessed the litter, and (15) N analysis of the plant material indicated that 2-3% of plant N originated from litter. Succession of the soil community mediating decomposition was assessed by phospholipid fatty acids (PLFA) combined with (13) C-PLFA. During the first 21?days of decomposition, saprotrophic fungi and Gram-negative bacteria were the dominant consumers of litter C. As decomposition progressed however, (13) C content of the fungal biomarkers declined substantially, and Gram-negative and Gram-positive bacteria became the primary reservoirs of labeled litter C. The putative PLFA marker for AMF (16:1ω5c) originated primarily from non-AMF sources. In AMF-invaded root litter, Gram-negative, Gram-positive, and 16:1ω5c markers became less (13) C-enriched relative to their counterparts in non-AMF-invaded microcosms during active decomposition. These patterns of (13) C: (12) C enrichment may result from AMF supply of (12) C from the plant to the decomposing soil microbial community; such C inputs could alter the microbial mediation of litter decomposition.     

丛枝菌根真菌在土壤氮素循环中的作用

作为植物需求量最大的营养元素,氮素是陆地生态系统初级生产力的主要限制因子。丛枝菌根真菌能与地球上80%以上的陆生植物形成菌根共生体,帮助宿主植物吸收土壤中的P、N等矿质养分。目前,丛枝菌根真菌与氮素循环相关研究侧重于真菌对氮素的吸收形态以及共生体中氮的传输代谢机制,却忽略了丛枝菌根真菌在固氮过程、矿化与吸收过程、硝化过程、反硝化过程以及氮素淋洗过程等土壤氮素循环过程中所起到的潜在作用,并且越来越多的证据也表明丛枝菌根真菌是影响土壤氮素循环过程的重要因子。总结了丛枝菌根真菌可利用的氮素形态及真菌的氮代谢转运相关基因的研究现状;重点分析了丛枝菌根真菌在调控土壤氮素循环过程中的潜在作用以及在生态系统中的重要生态学意义,同时提出了丛枝菌根真菌在土壤氮素循环过程中一些需要深入研究的问题。     

Influence of arbuscular mycorrhizal mycelial exudates on soil bacterial growth and community structure

Plant root systems colonized by arbuscular mycorrhizal (AM) fungi have previously been shown to influence soil bacterial populations; however, the direct influence of the AM extraradical mycelium itself on bacterial growth and community composition is not well understood. In this study, we investigated the effects of exudates produced by AM extraradical mycelia on the growth and development of an extracted soil bacterial community in vitro. The chemical composition of the mycelial exudates was analysed using proton nuclear magnetic resonance spectrometry. Following the addition of exudates to a bacterial community extracted from soil, bacterial growth and vitality were determined using a bacterial vitality stain and fluorescence microscopy. Changes in community composition were also analysed at various times over the course of 3 days by terminal restriction fragment length polymorphism analysis, in combination with cloning and sequencing of 16S rRNA genes. Mycelial exudates increased bacterial growth and vitality and changed bacterial community composition. Several Gammaproteobacteria, including a taxon within the Enterobacteriaceae, increased in frequency of occurrence in response to AM mycelial exudates. This study is the first attempt to identify carbohydrates from the extraradical mycelium of an AM fungus, and demonstrates the direct effects of mycelial exudates on a soil bacterial community.     

亚热带丛枝菌根与外生菌根森林对土壤氮循环的影响

菌根真菌能促进植物获取氮素从而调节土壤氮循环过程,但不同类型菌根影响土壤氮循环的特征尚待更多研究.本试验选择中国亚热带典型次生林植被,设置丛枝菌根(AM)和外生菌根(ECM)树种优势样地,对比分析两种菌根类型森林土壤氮状态的差异,以探究菌根类型影响土壤氮循环的可能作用机制.结果 表明,AM与ECM森林的土壤总氮和铵态氮...     

Polyphosphate dynamics in mycorrhizal roots during colonization of an arbuscular mycorrhizal fungus

Inorganic polyphosphate (poly P) has been considered to be a translocatable form of phosphate (Pi) in arbuscular mycorrhizal fungi (AMF). Here we examined time-course changes in poly P content during the AMF colonization process. Onion (Allium cepa) plants were cultured with or without inoculation with Gigaspora margarita for 2-8 wk with periodic sampling. Poly P in the extracts, purified through gel filtration, was quantified by the reverse reaction of polyphosphate kinase. The length of poly P in mycorrhizal roots appeared to be shorter than in extraradical hyphae or in spores of the AMF, indicating that AMF depolymerize poly P before providing Pi to the host. The poly P content increased as colonization proceeded, and was highly correlated with the weight of the colonized roots. These results support the model that AMF supply Pi to the host through the poly P pool, and that the poly P content of a mycorrhizal root can be a good indicator of the Pi-supplying activity of AMF.     

Plant community and soil available nutrients drive arbuscular mycorrhizal fungal community shifts during alpine meadow degradation

Arbuscular mycorrhizal fungi (AMF) play an important role in maintaining the function and sustainability of grassland ecosystem, but they are also susceptible to environmental changes. In recent decades, alpine meadows on the Tibetan Plateau have experienced severe degradation due to the impact of human activities and climate change. But it remains unclear how degradation affects the AMF community, a group of functionally important root associated microorganisms, which potentially limit the development and application of microbial technologies in the restoration of degraded grasslands. In this study, we investigated AMF communities richness and composition in non-degraded (ND), moderately degraded (MD) and severely degraded (SD) alpine meadows on the Tibetan Plateau, and then explored their main biotic and abiotic determinants. Alpine meadow degradation significantly reduced plant community biomass, richness, soil organic carbon, total nitrogen, total phosphorus, available nitrogen and available phosphorus, but increased soil pH. AMF community composition and the iesdominant family and genera differed significantly among different degradation stages. Grassland degradation shifted the AMF community composition in favor of Claroideoglomus over Rhizophagus, and resulted in a marked loss of Glomeraceae and the dominance of Diversisporaceae. Alpine meadow degradation significantly increased AMF hyphal density and richness, likely working as a plant strategy to relieve nutrient deficiencies or loss as a result of degradation. The structural equation model showed that AMF community richness and composition were significantly influenced by plant community, followed by soil available nutrients. Soil available nutrients was the key contributor to the increased AMF hyphal density and richness during grassland degradation. Our findings identify the effects of alpine meadow degradation on AMF richness and highlight the importance of the plant community in shaping the AMF community during alpine meadow degradation. These results suggest that plant community restoration should be the primary goal for the ecological restoration of degraded alpine meadows, and these soil functional microorganisms should be simultaneously integrated into ecological restoration strategies and management.     

菌根真菌对土壤呼吸以及凋落物分解的影响

土壤呼吸是植物固定的碳由陆地生态系统进入大气的主要途径之一; 凋落物分解是养分循环的重要环节。陆地植物的90%以上可同菌根真菌形成共生关系, 菌根真菌对于植物获取环境中的养分具有重要的作用。然而, 其对土壤呼吸和凋落物分解的影响却经常在生态系统对环境变化的响应研究中被忽视。本文系统地综述了国内外相关研究进展, 对菌根真菌如何影响土壤呼吸和凋落物分解这两个过程及这种影响如何受到环境变化的制约做了全面的分析, 并对以往研究中存在的问题以及未来的研究方向提出了展望。     

亚热带森林凋落叶分解过程土壤节肢动物群落的变化特征

土壤动物群落结构和多样性可能随凋落物分解进程和基质质量的变化不断改变。为了解亚热带森林凋落叶分解过程中土壤节肢动物群落变化特征,以四川盆地亚热带森林麻栎（Quercus acutissima）和柳杉（Cryptomeria fortunei）凋落叶为对象,于2011-2015年采用分解袋法研究了2种凋落叶分解过程中土壤节肢动物组成、结构和多样性动态变化。整个研究期间,柳杉和麻栎凋落叶分解袋中共捕获土壤节肢动物3855只,分属于16目51科,且均以等节跳科和棘跳科为优势类群;麻栎凋落叶中土壤节肢动物的个体密度随分解进程呈现增加趋势,在分解的1079天达最高值后降低,而柳杉凋落叶则在分解的156天急剧增加后快速降低,2种凋落叶中土壤节肢动物类群数量具有相似的动态变化过程;2种凋落叶中土壤节肢动物总体以菌食性数量比例最高,腐食性最低,且随凋落叶分解进程,植食性土壤节肢动物占比明显下降,菌食性则上升;非度量多维尺度（NMDS）分析显示,2种凋落叶中土壤节肢动物群落组成具有显著差异,聚类分析表明,2种凋落叶土壤节肢动物群落结构相似性随分解进程不断降低。亚热带森林凋落叶分解过程中土壤节肢动物群落组成、结构和多样性受凋落叶类型影响。     

New perspectives on forest soil carbon and nitrogen cycling processes: Roles of arbuscular mycorrhizal versus ectomycorrhizal tree species

Nearly all tree species develop symbiotic relationships with either arbuscular mycorrhizal (AM) or ectomycorrhizal (EM) fungi to acquire nutrients from soils, and hence influence soil carbon (C) and nitrogen (N) cycles in terrestrial ecosystems. It is crucial to understand the differences in soil C and N cycles between AM and EM forests and the underlying mechanisms. In this review, we first compared the differences in the soil C and N cycles between AM and EM forests, and synthesized the underlying mechanisms from perspectives of the inputs, stabilization, and outputs of soil C and N in forest ecosystems. We also compared the responses of soil C and N cycles between AM and EM forests to global changes. In this field, one major research priority is comparing the structure and function (including the soil C and N cycles) between AM and EM forest ecosystems to provide theoretical basis and solid data for improving forest productivity and ecosystem services. The second research focus is deepening the understanding of the effects of interactions between aboveground litter and belowground mycorrhiza and free-living microbes on soil C and N cycles to reveal the potential underlying mechanisms in forests with different mycorrhizal symbioses. Third, the research methodology and new techniques need refining and applying to explicitly focus on scaling up the fine-scale measurements to better expound and predict the C and N cycles in forest ecosystems. Finally, more studies on the stability of soil organic matter among different mycorrhizal forests are needed to precisely assess responses of the structure and function of forest ecosystems to global changes.     

Chemotropism in the arbuscular mycorrhizal fungus Glomus mosseae

In this work, we report the occurrence of chemotropism in the arbuscular mycorrhizal (AM) fungus Glomus mosseae. Fungal hyphae were able to respond to host-derived signals by reorienting their growth towards roots and to perceive chemotropic signals at a distance of at least 910 microm from roots. In order to reach the source of chemotropic signals, hyphal tips crossed interposed membranes emerging within 1 mm from roots, eventually establishing mycorrhizal symbiosis. The specificity of chemotropic growth was evidenced by hyphal growth reorientation and membrane penetration occurring only in experimental systems set up with host plants. Since pre-symbiotic growth is a critical stage in the life cycle of obligate AM fungal symbionts, chemotropic guidance may represent an important mechanism functional to host root location, appressorium formation and symbiosis establishment.     

外来植物紫茎泽兰入侵对土壤理化性质及丛枝菌根真菌(AMF)群落的影响

为了揭示外来植物紫茎泽兰入侵对入侵地土壤丛枝菌根真菌(AMF)群落及相关肥力的影响,比较测定了紫茎泽兰不同入侵程度土壤理化性质、AMF侵染率及AMF群落的差异。结果表明,紫茎泽兰入侵降低了土壤pH,使土壤中有机碳、全氮和速效钾含量分别增加83.0%,106.9%和111.0%;尽管对全磷含量没有显著影响,但有机磷含量呈升高的趋势,而速效磷呈降低的趋势。紫茎泽兰入侵降低了本地植物的AMF侵染率;随着入侵程度的加深,土壤中以膨胀无梗囊霉(Acauospora dilatata)为优势种的AMF群落结构逐渐转变为以近明球囊霉(Glomus claroideum )为优势种的结构,紫茎泽兰可在其根周选择培育近明球囊霉,而对其它AMF种,特别是对膨胀无梗囊霉则存在抑制作用;基于各AMF种多度的聚类分析表明,形成紫茎泽兰单优群落土壤中各AMF种多度与未入侵的本地植物群落及入侵程度较轻的紫茎泽兰与本地植物群落之间存在明显分歧。综合分析推断认为,紫茎法兰入侵改变了入侵地土壤理化性状,抑制AMF对土著植物的侵染,改变AMF群落,并在其根周选择培育近明球囊霉,这可能是紫茎泽兰入侵及扩张的重要途径之一。     

Spatial separation of litter decomposition and mycorrhizal nitrogen uptake in a boreal forest

Our understanding of how saprotrophic and mycorrhizal fungi interact to re-circulate carbon and nutrients from plant litter and soil organic matter is limited by poor understanding of their spatiotemporal dynamics. In order to investigate how different functional groups of fungi contribute to carbon and nitrogen cycling at different stages of decomposition, we studied changes in fungal community composition along vertical profiles through a Pinus sylvestris forest soil. We combined molecular identification methods with 14C dating of the organic matter, analyses of carbon:nitrogen (C:N) ratios and 15N natural abundance measurements. Saprotrophic fungi were primarily confined to relatively recently (< 4 yr) shed litter components on the surface of the forest floor, where organic carbon was mineralized while nitrogen was retained. Mycorrhizal fungi dominated in the underlying, more decomposed litter and humus, where they apparently mobilized N and made it available to their host plants. Our observations show that the degrading and nutrient-mobilizing components of the fungal community are spatially separated. This has important implications for biogeochemical studies of boreal forest ecosystems.     

Effect of soil bacteria on hyphal growth of the arbuscular mycorrhizal fungus Glomus claroideum

The effect of 46 bacterial strains isolated from tilled and non-tilled soils collected at 3 localities on the growth of intraradical hyphae of the arbuscular mycorrhizal (AM) fungusGlomus claroideum was demonstrated. A larger number of stimulatory bacterial isolates was obtained from tilled soils, but the bacteria showing the strongest stimulation of hyphal growth were isolated from a soil that had not been cultivated. Isolates obtained from hyphae of AM fungi showed no substantial stimulatory effects, but produced more uniform effects on hyphal growth than the isolates of bacteria obtained from soil. Bacterial cenoses present in 3 different soils differ significantly in their effects on AM fungi.     

Contrasting effects of ectomycorrhizal and arbuscular mycorrhizal tropical tree species on soil nitrogen cycling: the potential mechanisms and corresponding adaptive strategies

While it is increasingly recognized that ectomycorrhizal (ECM) and arbuscular mycorrhizal (AM) tree species vary in their effects on soil nitrogen (N) cycling, little is known about the mechanisms causing and how ECM and AM trees adapt to this variation. Using monoculture plots of six ECM and eight AM tropical trees planted in a common garden, we examined whether the contrasting effects of ECM and AM trees on soil N cycling could be explained by their differences in plant traits. Furthermore, rhizosphere effects on soil N transformations and soil exploration by fine roots were also measured to assess whether ECM and AM trees differed in N acquisition capacities. Results showed that soil NH₄⁺‐N concentration, net N mineralization and net nitrification rates were markedly lower, but soil C:N ratio was significantly higher beneath ECM trees than beneath AM trees. This more closed N cycling caused by ECM trees was attributed to their resource‐conservative traits, especially the poorer leaf litter decomposability compared with AM trees. To adapt to their induced lower soil N availability, ECM trees were found to have greater rhizosphere effects on NO₃^‐‐N concentration, net N mineralization and net nitrification rates to mine N, and higher soil exploration in terms of root length density to scavenge N from soils, indicating that these two strategies work in synergy to meet N demand of ECM trees. These findings suggest that ECM and AM trees have contrasting effects on soil N cycling owing to their differences in leaf litter decomposability and correspondingly possess different N acquisition capacities.     

红松混交林凋落物氮储量及分解释放对土壤氮的影响

2012年5—10月,采用直接收获法,研究了小兴安岭地区云冷杉红松混交林和枫桦红松混交林两种林型凋落物的未分解层(L层)、半分解层(F层)和腐殖质层(H层)以及土壤表层(S层)氮储量及凋落物分解释放对土壤氮影响。结果表明:研究期间两种林型凋落物现存量变化范围分别为19.43~27.25和21.25~24.28 t·hm-2,氮储量变化范围分别为287.21~418.22和274.81~351.21 kg·hm-2,各层氮含量大小次序均为LFHS;云冷杉红松混交林各层凋落物现存量及其氮储存量5月和9月达到峰值,每月氮储量从L~H层均增加,凋落物分解释放氮在F和H层易富集,输入到土壤中较少;枫桦红松混交林各层凋落物现存量及其氮储量5月和10月达到峰值,每月氮储量从L~H层均减少,氮在凋落物各层中均易迁移,输入到土壤中的氮比云冷杉红松混交林多;两种林型L、F、H层凋落物现存量以及H层氮含量与S层氮含量之间,L和F层凋落物现存量与H层氮含量之间均呈显著正相关。     

凋落物多样性及组成对凋落物分解和土壤微生物群落的影响

凋落物分解是生态系统营养物质循环的核心过程,而土壤微生物群落在凋落物分解过程中扮演着极其重要且不可替代的角色。随着生物多样性的丧失日益严峻,探讨凋落物多样性及组成对凋落物分解和土壤微生物群落的影响,不仅有助于了解凋落物分解的内在机制,而且可为退化草原生态系统的恢复提供参考。以内蒙古呼伦贝尔草原退化恢复群落中的草本植物为研究对象,依据植物多度、盖度、频度和物种的重要值及其在群落中的恢复程度筛选出排序前4的羊草（Leymus chinensis）、茵陈蒿（Artemisia capillaris）、麻花头（Serratula centauroides）、二裂委陵菜（Potentilla bifurca）的凋落物为实验材料,通过设置3种凋落物多样性水平（1,2,4）,包括11种凋落物组合（单物种凋落物共4种,两物种凋落物混合共6种,四物种凋落物混合共1种）,利用磷脂脂肪酸（PLFA）方法来研究分解60 d后凋落物多样性及组成对凋落物分解和土壤微生物群落的影响。结果表明：（1）凋落物物种多样性仅对C残余率具有显著影响,表现在两物种混合凋落物C残余率显著低于单物种凋落物,而凋落物组成对所观测的4个凋落物分解参数（质量、C、N残余率以及C/N）均具有显著影响;（2）凋落物物种多样性对细菌（B）含量具有显著影响,而凋落物组成对真菌（F）含量具有显著影响,两者对F/B以及微生物总量均无显著影响;（3）冗余分析结果表明凋落物组成与凋落物分解相关指标（凋落物质量、C、N残余率及C/N）和土壤微生物（真菌、细菌含量）的相关关系高于凋落物多样性。（4）进一步建立结构方程模型（Structural Equation Model,SEM）发现,凋落物初始C含量对凋落物质量、C、N残余率及C/N有显著正的直接影响;凋落物木质素含量对凋落物质量、C、N残余率有显著正的直接影响;凋落物初始N含量对N残余率有显著正的直接影响,而对C残余率及C/N有显著负的直接影响;凋落物初始C/N对凋落物质量、N残余率有显著正的直接影响,而对C/N有显著负的直接影响。此外,凋落物初始C、N、木质素含量及C/N均对真菌含量具有显著正影响,并可通过真菌对凋落物质量分解产生显著负的间接影响。以上结果表明该退化恢复区域优势种凋落物分解以初始C、木质素为主导,主要通过土壤真菌影响凋落物的分解进程,这将减缓凋落物的分解速率进而减慢草原生态系统的进程。这些结果为凋落物多样性及组成对自身分解和土壤微生物群落的影响提供了实验依据,也为进一步分析凋落物分解内在机制以及草原生态系统的恢复提供了数据参考。     

凋落物分解过程中土壤微生物群落的变化

凋落物分解是生态系统碳循环和营养物质循环的关键过程, 受多种因素共同影响。土壤微生物是影响凋落物分解的重要因素, 其群落组成在一定程度上依赖于所处植物群落的特征。因此, 研究分解过程中微生物群落组成的变化及其对植物多样性的响应, 有利于对凋落物分解机制的理解。本文采用分解袋野外原位分解的方法, 对凋落物分解过程中微生物群落的变化及其对所处森林环境中树木的种类和遗传多样性的响应进行了研究。结果表明: (1)凋落物分解183天后, 土壤中微生物群落的多样性降低, 并且森林群落的物种多样性与微生物群落多样性呈负相关关系; (2)凋落物分解前后, 土壤中真菌和细菌群落的磷脂脂肪酸(PLFA)量均有所增加, 说明凋落物分解为微生物生存和繁殖提供了养分; (3)地形因素是影响微生物群落变化最显著的因素, 可解释微生物群落变化的29.55%; 其次是凋落物的基质质量, 可以解释15.39%; 最后是森林群落的多样性, 可以解释8.45%; 这3种因素共同解释率为2.97%。综上所述, 与森林群落的植物多样性相比, 样地的地形因素与凋落物的基质质量对微生物群落的影响更显著。     

Leguminous plants significantly increase soil nitrogen cycling across global climates and ecosystem types

Leguminous plants are an important component of terrestrial ecosystems and significantly increase soil nitrogen (N) cycling and availability, which affects productivity in most ecosystems. Clarifying whether the effects of legumes on N cycling vary with contrasting ecosystem types and climatic regions is crucial for understanding and predicting ecosystem processes, but these effects are currently unknown. By conducting a global meta-analysis, we revealed that legumes increased the soil net N mineralization rate (R_min) by 67%, which was greater than the recently reported increase associated with N deposition (25%). This effect was similar for tropical (53%) and temperate regions (81%) but was significantly greater in grasslands (151%) and forests (74%) than in croplands (−3%) and was greater in in situ incubation (101%) or short-term experiments (112%) than in laboratory incubation (55%) or long-term experiments (37%). Legumes significantly influenced the dependence of R_min on N fertilization and experimental factors. The R_min was significantly increased by N fertilization in the nonlegume soils, but not in the legume soils. In addition, the effects of mean annual temperature, soil nutrients and experimental duration on R_min were smaller in the legume soils than in the nonlegume soils. Collectively, our results highlighted the significant positive effects of legumes on soil N cycling, and indicated that the effects of legumes should be elucidated when addressing the response of soils to plants.