Changes in ectomycorrhizal community structure on two containerized oak hosts across an experimental hydrologic gradient

Shifts in ectomycorrhizal (ECM) community structure were examined across an experimental hydrologic gradient on containerized seedlings of two oak species, Quercus montana and Quercus palustris, inoculated from a homogenate of roots from mature oak trees. At the end of one growing season, seedlings were harvested, roots were sorted by morphotype, and proportional colonization of each type was determined. DNA was subsequently extracted from individual root tips for polymerase chain reaction, restriction fragment length polymorphism, and rDNA sequencing of the ITS1/5.8S/ITS2 region to determine identities of fungal morphotypes. Twelve distinct molecular types were identified. Analysis of similarity showed that ECM fungal assemblages shifted significantly in composition across the soil moisture gradient. Taxa within the genus Tuber and the family Thelephoraceae were largely responsible for the changes in fungal assemblages. There were also significant differences in ECM community assemblages between the two oak host species. These results demonstrate that the structure of ECM fungal communities depends on both the abiotic and biotic environments and can shift with changes in soil moisture as well as host plant, even within the same genus.     

Influence of soil nutrients on ectomycorrhizal communities in a chronosequence of mixed temperate forests

Many factors associated with forests are collectively responsible for controlling ectomycorrhizal (ECM) fungal community structure, including plant species composition, forest structure, stand age, and soil nutrients. The objective of this study was to examine relationships among ECM fungal community measures, local soil nutrients, and stand age along a chronosequence of mixed forest stands that were similar in vegetation composition and site quality. Six combinations of age class (5-, 26-, 65-, and 100-year-old) and stand initiation type (wildfire and clearcut) were replicated on four sites, each representing critical seral stages of stand development in Interior Cedar-Hemlock (ICH) forests of southern British Columbia. We found significant relationships between ECM fungal diversity and both available and organic P; available P was also positively correlated with the abundance of two ECM taxa (Rhizopogon vinicolor group and Cenoccocum geophilum). By contrast, ECM fungal diversity varied unpredictably with total and mineralizable N or C to N ratio. We also found that soil C, N, available P, and forest floor depth did not exhibit strong patterns across stand ages. Overall, ECM fungal community structure was more strongly influenced by stand age than specific soil nutrients, but better correlations with soil nutrients may occur at broader spatial scales covering a wider range of site qualities.     

Post-fire, seasonal and annual dynamics of the ectomycorrhizal community in a Quercus ilex L. forest over a 3-year period

Two study plots, burned and control, were established in autumn 1998 in a Quercus ilex forest located in northern Spain, part of which had been affected by a low intensity fire in 1994. Soil samples for ectomycorrhizae (ECM) were taken over a 3-year period in each study plot in spring, summer, autumn and winter. ECM morphotypes were identified and the relative abundance of each morphotype in each soil sample calculated, along with species richness, Shannon diversity index and percentage of mycorrhization in each soil sample. The relative abundance of certain ECM morphotypes differed between burned and control plots, and the percentage of mycorrhizal tips was significantly lower in the burned than in the control plot. Nevertheless, there were no significant differences in the diversity, species richness or species composition of the ECM community in the burned and control plots. The dominant ECM morphotypes in both stands were Cenococcum geophilum and several thelephoroid fungi. Sphaerosporella brunnea and Pisolithus tinctorius thrived especially in the burned plot, whereas three ectomycorrhizal morphotypes assigned to the genus Hebeloma were especially abundant in the control plot. There was no significant variation in the relative abundance of the ECM morphotypes between seasons, but ECM community species richness was highest in autumn and lowest in summer. The percentage of mycorrhizal tips reached a maximum in winter, with its minimum in autumn. Collection of samples over the 3-year period also enabled us to detect a significant increase in percentage of ECM colonisation in the burned stand over time.     

Dominant forest tree mycorrhizal type mediates understory plant invasions

Forest mycorrhizal type mediates nutrient dynamics, which in turn can influence forest community structure and processes. Using forest inventory data, we explored how dominant forest tree mycorrhizal type affects understory plant invasions with consideration of forest structure and soil properties. We found that arbuscular mycorrhizal (AM) dominant forests, which are characterised by thin forest floors and low soil C : N ratio, were invaded to a greater extent by non‐native invasive species than ectomycorrhizal (ECM) dominant forests. Understory native species cover and richness had no strong associations with AM tree dominance. We also found no difference in the mycorrhizal type composition of understory invaders between AM and ECM dominant forests. Our results indicate that dominant forest tree mycorrhizal type is closely linked with understory invasions. The increased invader abundance in AM dominant forests can further facilitate nutrient cycling, leading to the alteration of ecosystem structure and functions.     

Temporal variation in temperature and rainfall differentially affects ectomycorrhizal colonization at two contrasting sites

We examined the roles that seasonal shifts in precipitation and temperature played in the ectomycorrhizal (ECM) colonization of pinyon pine ( Pinus edulis Engelm.) at two contrasting sites in northern Arizona. Pinyons growing in ash and cinder soils experienced much greater water and nutrient stress than pinyons growing nearby in sandy-loam soils. Over a one year period, we obtained monthly measurements of ECM colonization, root zone soil moisture and temperature, and air temperature and precipitation. Four major patterns emerged. Firstly, although climate as measured by ambient temperature and precipitation did not vary between the two sites, soil temperature was significantly higher and soil moisture significantly lower at the cinder site than at the sandy-loam site. Secondly, ECM colonization was significantly higher at the cinder site for 5 of 12 months. Thirdly, although nearly 70% of the variation in ECM colonization of pinyons growing in cinder soil was predicted by a combination of soil moisture and soil temperature, these same variables had little predictive power for pinyons growing in sandy-loam soils. Air temperature and precipitation were also significantly correlated with ECM colonization at the cinder site but not the sandy-loam site. Fourthly, a watering experiment showed that ECM colonization significantly increased with supplemental water at the cinder site, but not at the sandy-loam site. Thus, in two sites that did not differ in plant community or climate, ectomycorrhizas in cinder soils were far more sensitive to changes in moisture and temperature than ectomycorrhizas in sandy-loam soils.     

Regional scale gradients of climate and nitrogen deposition drive variation in ectomycorrhizal fungal communities associated with native Scots pine

Ectomycorrhizal fungi commonly associate with the roots of forest trees where they enhance nutrient and water uptake, promote seedling establishment and have an important role in forest nutrient cycling. Predicting the response of ectomycorrhizal fungi to environmental change is an important step to maintaining forest productivity in the future. These predictions are currently limited by an incomplete understanding of the relative significance of environmental drivers in determining the community composition of ectomycorrhizal (ECM) fungi at large spatial scales. To identify patterns of community composition in ECM fungi along regional scale gradients of climate and nitrogen deposition in Scotland, fungal communities were analysed from 15 seminatural Scots pine (Pinus sylvestris L.) forests. Fungal taxa were identified by sequencing of the ITS rDNA region using fungal‐specific primers. Nonmetric multidimensional scaling was used to assess the significance of 16 climatic, pollutant and edaphic variables on community composition. Vector fitting showed that there was a strong influence of rainfall and soil moisture on community composition at the species level, and a smaller impact of temperature on the abundance of ectomycorrhizal exploration types. Nitrogen deposition was also found to be important in determining community composition, but only when the forest experiencing the highest deposition (9.8 kg N ha^?1 yr^?1) was included in the analysis. This finding supports previously published critical load estimates for ectomycorrhizal fungi of 5–10 kg N ha^?1 yr^?1. This work demonstrates that both climate and nitrogen deposition can drive gradients of fungal community composition at a regional scale.     

Nutrient and Carbon Additions to the Microbial Soil Community and its Impact on Tree Seedlings in a Boreal Spruce Forest

We have added glucose and nutrients to manipulate soil microbial activity and nutrient availability in a boreal spruce forest to study the performance of birch and spruce seedlings in relation to the soil microbial community. The proportion of aboveground biomass in the seedlings was largest in plots amended with extra nutrients, while ectomycorrhizal (ECM) colonisation was low in these plots. ECM appeared beneficial for growth of both species, but only at low levels of colonisation (<25% ECM colonised root-tips). The soil microbial biomass, as determined by total PLFA, was largest in plots treated with glucose and there was a significant negative relationship between birch seedling size and levels of total PLFA in soil. This could be taken to suggest that poor seedling growth was due to nutrient limitation caused by microbial assimilation. However, the treatment response of the birch seedlings was generally weak, and spruce often showed no response at all to the addition of nutrients and glucose. The most consistent parameter for the variation in plant performance, as well as for the microbial soil community, was the block-effect. This suggests a strong spatial structure in the soil microbial community, and that this structure was robust with respect to our treatments even though they continued over a 3-year period.     

连续三年夜间增温和施氮对云杉外生菌根及菌根真菌多样性的影响

以西南亚高山针叶林建群种粗枝云杉(Picea asperata)为研究对象,采用红外加热模拟增温结合外施氮肥(NH₄NO₃ 25 g N m^-2 a^-1)的方法,研究连续3a夜间增温和施肥对云杉幼苗外生菌根侵染率、土壤外生菌根真菌生物量及其群落多样性的影响。结果表明:夜间增温对云杉外生菌根侵染率的影响具有季节性及根级差异。夜间增温对春季(2011年5月)云杉1级根,夏季(2011年7月)和秋季(2010年10月)云杉2级根侵染率影响显著。除2011年7月1级根外,施氮对云杉1、2级根侵染率无显著影响。夜间增温对土壤中外生菌根真菌的生物量和群落多样性无显著影响,施氮及增温与施氮联合处理使土壤中外生菌根真菌生物量显著降低,但却提高了外生菌根真菌群落的多样性。这说明云杉幼苗外生菌根侵染率对温度较敏感,土壤外生菌根真菌生物量及其群落多样性对施氮较敏感。这为进一步研究该区域亚高山针叶林地下过程对全球气候变化的响应机制提供了科学依据。     

Similar taxonomic richness but different communities of ectomycorrhizas in native forests and non-native plantation forests

This investigation sought to examine if there was a difference between the ectomycorrhizal (ECM) communities in plots of native oak and introduced Scots pine and Sitka spruce forest. The ECM communities in four plots of each forest type were described, from five soil cores collected in each plot, by morphotyping, internal transcribed spacer (ITS)-restriction fragment length polymorphism matching of mycorrhizas and sporocarps and ITS sequencing. Fifty-one distinct taxa were distinguished; 25 were identified to species level, 11 to genus and 15 remained unidentified. Seventy-one ECM species were recorded as sporocarps from the forest plots; most (43 species) were found in the Sitka spruce plots. The below-ground ECM communities of the different forest types did not differ significantly with respect to species richness of taxa on roots, but differed in species composition. Multivariate analysis produced a clear separation of the communities of the different forest types using below-ground data, but the above-ground sporocarp data did not separate the forest types. Moreover, results of a Mantel test found no relationship between the above- and below-ground similarity matrices. The oak plots had the most distinctive ECM community, with Laccaria amethystina and Elaphomyces granulatus being frequent. The Sitka spruce plots showed the lowest intra-forest type similarity and were often dominated by "nursery type" ectomycorrhizas. There was only 10% similarity between the above- and below-ground ECM species in these plots, different colonisation methods of ectomycorrhizal taxa and insufficient below-ground sampling being possible reasons for this disparity. Our results indicate that plantations of non-native Sitka spruce can support similar levels of ECM diversity as native forests.     

Effects of ectomycorrhizal colonization and nitrogen fertilization on morphology of root tips in a <Emphasis Type="Italic">Larix gmelinii</Emphasis> plantation in northeastern China

Root morphology is important in understanding root functions in forest ecosystems. However, the effects of ectomycorrhizal colonization and soil nutrient availability on root morphology is not clear. In this study, root morphology in relation to season, soil depth, soil nitrogen (N) availability, and mycorrhizal fungal colonization were investigated in a larch (Larix gmelinii) plantation in northeastern China. The first-order roots (or root tips) of larch were sampled four times in May, July, and September of 2005, and May of 2006 from two depths of upper soil layer (0–10 and 10–20 cm) in the control and the N-fertilized plots. The results showed that ectomycorrhizal (ECM) colonization rates for the first-order roots were reduced by 17% under N fertilization. The peak of root colonization rates occurred in summer and was positively correlated with soil temperature. ECM colonization significantly altered root morphology: root diameter was increased by 19 and 29%, root length shortened by 27 and 25%, and specific root length (SRL) reduced by 16 and 19% for the control and the N-fertilized plots, respectively. N fertilization led to decreased root length, but did not affect root diameter and SRL. In addition, effects of ECM colonization on root morphology varied with season and soil depth. The observed relationships among ECM fungal colonization, soil N availability, and root-tip morphology should improve our understanding of how root tips respond to environmental changes in soil in temperate forest ecosystems.     

Temporal Changes in the Ectomycorrhizal Community in Two Soil Horizons of a Temperate Oak Forest

The species structure of an ectomycorrhizal (ECM) community was assessed monthly for 15 months in the two horizons (A1 and A2) of an oak temperate forest in northeastern France. Ectomycorrhizal species were identified each month by internal transcribed spacer sequencing. Seventy-five fungal symbionts were identified. The community was dominated by Tomentellaceae, Russulaceae, Cortinariaceae, and Boletales. Four species are abundant in the study site: Lactarius quietus, Tomentella sublilacina, Cenococcum geophilum, and Russula sp1. The relative abundance of each species varied depending on the soil horizon and over time. Some species, such as L. quietus, were present in the A1 and A2 horizons. C. geophilum was located particularly in the A2 horizon, whereas T. sublilacina was more abundant in A1. Some species, such as Clavulina sp., were detected in winter, while T. sublilacina and L. quietus were present all year long. Our results support the hypothesis that a rapid turnover of species composition of the ECM community occurs over the course of a month. The spatial and temporal unequal distribution of ECM species could be explained by their ecological preferences, driven by such factors as root longevity, competition for resources, and resistance to environmental variability.     

Ectomycorrhizal fungal community structure across a bog-forest ecotone in southeastern Alaska

We examined the ectomycorrhizal (ECM) fungal community across a bog-forest ecotone in southeastern Alaska. The bog and edge were both characterized by poorly drained Histosols and a continuous layer of Sphagnum species, ericaceous shrubs, Carex species, and shore pine [Pinus contorta Dougl. ex Loud. var. contorta]. The forest had better-drained Inceptisols and Spodosols, a tree community comprised of western hemlock [Tsuga heterophylla (Raf.) Sarg.], yellow cedar (Thuja plicata Donn ex D. Don.), Sitka spruce [Picea sitchensis (Bong.) Carr.] and shore pine, and an understorey of ericaceous shrubs and herbs. ECM root tip density (tips cm^–3 soil) was significantly greater in the forest than the edge or bog and ECM colonization was significantly different in all three plant communities. The below ground ECM fungal taxa were analyzed using molecular techniques (PCR-RFLP and DNA sequencing). Three ECM fungal taxa, Suillus tomentosus (Kauffman) Singer, Cenococcum geophilum Fr.:Fr, and a Russula species, differed in relative frequency, yet were among the four most frequent in all three plant communities. Although differences in ECM fungal richness were observed across plant communities, unequal sampling of ECM roots due to root density and colonization differences confounded richness comparisons. Using resampling procedures for creating taxon-accumulation curves as a function of sampled ECM roots revealed similarities in cumulative ECM fungal taxa richness across the ecotone.     

The functioning of ectomycorrhizal fungi in the field: linkages in space and time

Individual trees, either of the same or different species, can be linked spatially and temporally by the hyphae of ectomycorrhizal (ECM) fungi that allow carbon and nutrients to pass among them and promote forest establishment following disturbance. Spatial and temporal linkages between plants influence the function of ECM fungi in the field. Studies indicate that ECM linkages can reduce plant competition for resources, promote forest recovery, and influence the pattern of plant succession. The degree of influence depends on many factors, including the composition and arrangement of the vegetative community and soil and climatic conditions. Management practices that create intense disturbance and loss of organic matter or promote the introduction of non-ectomycorrhizal host species can decrease the ability of plants to form linkages with ECM fungi. Management practices that retain living trees and shrubs and input of organic matter provide the energy source and substrate necessary for ECM linkages. More research is needed to determine the degree to which ECM fungal linkages occur in the field and their role in ecosystem function and long-term health.     

Ectomycorrhizal root development in wet Alder carr forests in response to desiccation and eutrophication

Effects of desiccation and eutrophication on ectomycorrhizal (ECM) root development in wet Alder carr forests in The Netherlands were studied. In northwestern Europe, wet Alder carr forests are found mostly in peatlands and along streams, forming an important component of wetland ecosystems. The dominant tree species in wet Alder carr forests is Alnus glutinosa (L.) Gaertn. (Black alder), which associates with ectomycorrhizal fungi. During recent decades, wet Alder carr forests in Europe have declined because of desiccation and eutrophication, particularly in The Netherlands. In the present study, the number of root tips of A. glutinosa trees was highest in an undisturbed wet Alder carr forest in a peatland area. Eutrophication in the peatland area significantly inhibited ectomycorrhizal (ECM) root development of A. glutinosa. In the eutrophied forest, ECM root tips were observed only close to A. glutinosa trees growing on hummocks. The concentrations of nitrate and potassium in soil water of the eutrophied forest were significantly higher than in the undisturbed forest, while magnesium and iron concentrations and the pH were significantly lower. The number of ECM root tips of A. glutinosa in a desiccated forest along a stream was generally lower than in an undisturbed wet Alder carr forest on waterlogged soil in the same area. The sulphate concentration in soil water in the desiccated forest was significantly higher than in the forest on waterlogged soil. ECM root development of A. glutinosa may have been negatively affected by the chemical composition of the soil water.     

抚育间伐对油松人工林下大型真菌的影响

真菌在森林凋落物分解过程中起重要作用,研究间伐如何影响真菌进而影响凋落物分解,对深入了解间伐调控人工林凋落物分解有重要意义。本文以抚育间伐后的中龄油松人工林为研究对象,设立对照（I）、轻度（II）、中度（III）和强度（IV）4种间伐强度,于2011年对间伐后林下大型真菌进行两次调查,分析了不同间伐强度下大型真菌的科的分布、优势种组成和生态指标（包括丰富度指数、多样性指数和均匀度指数）。结果表明：（1）8月、9月采集到的大型真菌分别为35种和25种,分属13个科和10个科;（2）在大型真菌出菇期,间伐改变了大型真菌的优势种组成,对照林下大型真菌优势种最初为外生菌根菌（粘盖乳牛肝菌 血红铆钉菇）后变为腐生菌（大盖小皮伞和脐顶小皮伞）,而间伐后林下优势种始终为腐生菌;（3）间伐影响大型真菌的生态指标,中度间伐林下大型真菌丰富度和多样性指数最高。总之,适度间伐不仅有利于提高林下大型真菌的丰富度与多样性,同时使其群落结构发生改变,群落优势种由外生菌根菌变为以分解凋落物为主的腐生菌,可促进凋落物的分解和养分循环。     

Does forest liming impact the enzymatic profiles of ectomycorrhizal communities through specialized fungal symbionts?

Liming (Ca–Mg soil amendment) is a forestry practice used to correct soil acidification and restore health and productivity in declining stands. Liming is known to modify tree mineral nutrition beyond the sole Ca and Mg. We hypothesized that liming also modifies the very functioning of the tree absorbing system (that is the ectomycorrhizal fine roots) in a way that facilitates the mobilization of mineral nutrients, particularly those entrapped in soil organic matter. This hypothesis has been tested here in beech and Norway spruce stands in North-Eastern France. In autumn, we compared the ectomycorrhizal community structure and the enzymatic profiles of ectomycorrhizal root tips in limed and untreated plots by measuring the activities of eight enzymes related to the degradation of soil organic matter. The results show that the ectomycorrhizal community responds to the Ca–Mg amendment and to the resulting soil modifications by modified enzyme activity profiles and ability to mobilize nutrients from soil organic matter. The effects of liming on the belowground functioning of the tree stands result essentially from specialized ECM fungal species such as Clavulina cristata (with strong glucuronidase activity), Lactarius subdulcis (with strong laccase activity) or Xerocomus pruinatus (with strong leucine aminopeptidase activity).     

Species composition of an ectomycorrhizal fungal community along a local nutrient gradient in a boreal forest

Soil abiotic factors are considered to be important in determining the distribution of ectomycorrhizal (ECM) fungal species; however, there are few field data to support this. Here, we relate ECM species distributions to changes in soil chemistry along a short (90-m), natural nutrient gradient. The ECM community was characterized, using morphological and molecular techniques, in soil samples collected at 10-m intervals. There were pronounced changes in ECM fungal community structure along the transect, with many taxa showing discrete distributions. Although there was a change of host from Pinus to Picea along the gradient, host-specific fungi did not account for the observed change in community structure. Ordination analyses showed that community structure was strongly correlated with soil characteristics, in particular extractable ammonium and base saturation. However, autocorrelation among soil parameters makes it difficult to isolate the effects of individual parameters. The distinctive changes in soil and vegetation along the transect used in this study provided an exceptional opportunity to examine the local-scale impact of natural spatial heterogeneity on an ECM fungal community.     

神农架国家公园林线过渡带土壤真菌多样性

林线过渡带是陆地生态系统对气候变化响应的敏感区域,研究林线过渡带土壤真菌的群落结构和形成机制,对于预测气候变化对土壤养分循环和维持陆地生态系统功能的影响具有重要意义。利用Illumina高通量测序技术分析了神农架国家公园林线上下的灌木林和针叶林的土壤真菌群落结构和多样性。结果表明,在真菌物种组成上,两种植被类型的土壤优势菌门、属和种类不同,针叶林和灌木林的优势菌门分别是担子菌门(Basidiomycota)和接合菌门(Zygomycota)。除趋势对应分析(DCA)和不相似性检验(Dissimilarity test)表明两种林型的土壤真菌群落结构组成存在显著差异,且针叶林土壤真菌Shannon指数、Chao值和Richness指数均显著(P0.05)高于灌木林。典范对应分析(CCA)和Mantel检验显示土壤真菌群落结构与土壤p H、植物多样性、土壤温度和土壤湿度存在显著相关性。因此,林线过渡带上下的土壤真菌群落结构和多样性存在显著差异,土壤p H、植物多样性、土壤湿度和土壤温度可能是影响土壤真菌群落结构的重要因素。     

Ectomycorrhizal fungal communities associated to Nothofagus species in Northern Patagonia

Ectomycorrhizal fungi constitute an important component of soil biota in Nothofagus forests in Patagonia. However, ectomycorrhizal fungal community is poorly known in this region. Here, we assess biodiversity and community compositions of ectomycorrhizal fungal species associated with Nothofagus dombeyi, N. obliqua and N. alpina. We selected three monospecific Nothofagus forest sites for each species within the boundaries of the Lanin National Park in Northern Patagonia. Ectomycorrhizal fungal species were identified based on morphotyping and rDNA (ITS and 28S rDNA) sequence analysis using both universal and taxon-specific primers. Contrary to previous studies on congeneric host trees, our results showed no significant differences among Nothofagus forest types in terms of fungal biodiversity and community composition. However, altitude had a strong effect on the structure of the ectomycorrhizal fungal community associated with Nothofagus spp.     

Fungal community dynamics in relation to substrate quality of decaying Norway spruce ( Picea abies [L.] Karst.) logs in boreal forests

Decaying wood plays an important role in forest biodiversity, nutrient cycling and carbon balance. Community structure of wood-inhabiting fungi changes with mass loss of wood, but the relationship between substrate quality and decomposers is poorly understood. This limits the extent to which these ecosystem services can be effectively managed. We studied the fungal community and physico-chemical quality (stage of decay, dimensions, density, moisture, C : N ratio, lignin and water or ethanol extractives) of 543 Norway spruce logs in five unmanaged boreal forest sites of southern Finland. Fungi were identified using denaturing gradient gel electrophoresis and sequencing of DNA extracted directly from wood samples. Macroscopic fruiting bodies were also recorded. Results showed a fungal community succession with decreasing wood density and C : N ratio, and increasing moisture and lignin content. Fungal diversity peaked in the most decayed substrates. Ascomycetes typically colonized recently fallen wood. Brown-rot fungi preferred the intermediate decay stages. White-rot fungi represented approximately one-fifth of sequenced species in all decay phases excluding the final phase, where ectomycorrhizal (ECM) fungi became dominant. Lignin content of logs with white-rot fungi was low, and ECM fungi were associated with substrates containing abundant nitrogen. Macroscopic fruiting bodies were observed for only a small number of species detected with molecular techniques.