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1.
Most of the temperate conifers associate with ectomycorrhizal fungi, but their roots also harbour a wide range of endophytes. We focused on ascomycetes associating with basidiomycetous ectomycorrhizas of Norway spruce in a temperate montane forest in central Europe and found that the majority of the co-associated fungi belonged to the Rhizoscyphus ericae aggregate (REA), being dominated by Meliniomyces variabilis. We further tested the ability of representative isolates to colonize spruce root tips and European blueberry (Vaccinium myrtillus) hair roots in an agar system as well as their effect on blueberry growth in a peat-agar system. M. variabilis intracellularly colonized spruce (Picea abies) root tip cortex, formed ericoid mycorrhizas in blueberry and enhanced blueberry shoot and root growth in comparison with non-inoculated plants. Our findings suggest that spruce ectomycorrhizas may represent selective niches for ericoid mycorrhizal fungi in habitats lacking suitable ericaceous hosts.  相似文献   

2.
Fine root acclimation to different environmental conditions is crucial for growth and sustainability of forest trees. Relatively small changes in fine root standing biomass (FRB), morphology or mycorrhizal symbiosis may result in a large change in forest carbon, nutrient and water cycles. We elucidated the changes in fine root traits and associated ectomycorrhizal (EcM) fungi in 12 Norway spruce stands across a climatic and N deposition gradient from subarctic‐boreal to temperate regions in Europe (68°N–48°N). We analysed the standing FRB and the ectomycorrhizal root tip biomass (EcMB, g m?2) simultaneously with measurements of the EcM root morphological traits (e.g. mean root length, root tissue density (RTD), N% in EcM roots) and frequency of dominating EcM fungi in different stands in relation to climate, soil and site characteristics. Latitude and N deposition explained the greatest proportion of variation in fine root traits. EcMB per stand basal area (BA) increased exponentially with latitude: by about 12.7 kg m?2 with an increase of 10° latitude from southern Germany to Estonia and southern Finland and by about 44.7 kg m?2 with next latitudinal 10° from southern to northern Finland. Boreal Norway spruce forests had 4.5 to 11 times more EcM root tips per stand BA, and the tips were 2.1 times longer, with 1.5 times higher RTD and about 1/3 lower N concentration. There was 19% higher proportion of root tips colonized by long‐distance exploration type forming EcM fungi in the southern forests indicating importance of EcM symbiont foraging strategy in fine root nutrient acquisition. In the boreal zone, we predict ca. 50% decrease in EcMB per stand BA with an increase of 2 °C annual mean temperature. Different fine root foraging strategies in boreal and temperate forests highlight the importance of complex studies on respective regulatory mechanisms in changing climate.  相似文献   

3.
Dipterocarps are one of the most important tree families in the lowland forests of Southeast Asia and are somewhat unusual among tropical trees in that they form ectomycorrhizal (EcM) symbiotic root‐inhabiting fungal associations. It has been hypothesized that dipterocarps have been partnered in this mutualistic association prior to the separation of Gondwana. Under many conditions EcMs form rapidly on dipterocarp seedlings through inocula present in the soil, although few studies have been conducted to provide evidence that they improve seedling establishment and performance. There are hundreds of EcM species associated with dipterocarps. Fungal fruit body surveys suggest that the most important families are Amanitaceae, Boletaceae, and Russulaceae, although Thelephoraceae also become numerically important when root tips are examined. EcM communities are affected by various biotic and abiotic factors, as well as anthropogenic perturbations, and the importance of these in structuring EcM communities is examined herein.  相似文献   

4.
Many temperate forests of the Northeastern United States and Europe have received significant anthropogenic acid and nitrogen (N) deposition over the last century. Although temperate hardwood forests are generally thought to be N-limited, anthropogenic deposition increases the possibility of phosphorus (P) limiting productivity in these forest ecosystems. Moreover, inorganic P availability is largely controlled by soil pH and biogeochemical theory suggests that forests with acidic soils (i.e., <pH 5) are particularly vulnerable to P limitation. Results from previous studies in these systems are mixed with evidence both for and against P limitation. We hypothesized that shifts in mycorrhizal colonization and community structure help temperate forest ecosystems overcome an underlying P limitation by accessing mineral and organic P sources that are otherwise unavailable for direct plant uptake. We examined arbuscular mycorrhizal (AM) and ectomycorrhizal (EcM) communities and soil microbial activity in an ecosystem-level experiment where soil pH and P availability were manipulated in mixed deciduous forests across eastern Ohio, USA. One year after treatment initiation, AM root biomass was positively correlated with the most available P pool, resin P, while AM colonization was negatively correlated. In total, 15,876 EcM root tips were identified and assigned to 26 genera and 219 operational taxonomic units (97% similarity). Ectomycorrhizal richness and root tip abundance were negatively correlated with the moderately available P pools, while the relative percent of tips colonized by Ascomycetes was positively correlated with soil pH. Canonical correspondence analysis revealed regional, but not treatment, differences in AM communities, while EcM communities had both treatment and regional differences. Our findings highlight the complex interactions between mycorrhizae and the soil environment and further underscore the fact that mycorrhizal communities do not merely reflect the host plant community.  相似文献   

5.
杨浩  史加勉  郑勇 《生态学报》2024,44(7):2734-2744
森林生态系统在全球碳(C)储量中占据极为重要的地位。菌根真菌广泛存在于森林生态系统中,在森林生态系统C循环过程中发挥重要的作用。阐述了不同菌根类型真菌在森林生态系统C循环过程中的功能,对比了温带/北方森林与热带/亚热带森林中菌根真菌介导的C循环研究方面新近取得的研究结果。发现温带和北方森林的外生菌根(EcM)植物对地上生物量C的贡献相对较小,然而是地下C储量的主要贡献者;以丛枝菌根(AM)共生为主的热带/亚热带森林地表生物量占比较高,表明AM植被对热带/亚热带森林地上生物量C的贡献相对较大。我们还就全球变化背景下,菌根真菌及其介导的森林生态系统C汇功能,以及不同菌根类型树种影响C循环的机制等进行了总结。菌根真菌通过影响凋落物分解、土壤有机质形成及地下根系生物量,进而影响整个森林生态系统的C循环功能。菌根介导的森林C循环过程很大程度上取决于(优势)树木的菌根类型和森林土壤中菌根真菌的群落结构。最后指出了当前研究存在的主要问题以及未来研究展望。本文旨在明确菌根真菌在森林生态系统C循环转化过程中的重要生态功能,有助于准确地评估森林生态系统C汇现状,为应对全球变化等提供重要的依据。  相似文献   

6.
The ability of ericoid and ectomycorrhizal fungi to utilize 14C-labelled lignin and O14CH3-labelled dehydropolymer of coniferyl alcohol as sole C sources has been assessed in pure culture studies. The results indicate that ericoid mycorrhizal fungi are more effective in degrading lignin than ectomycorrhizal fungi. Amongst the ectomycorrhizal fungi the facultative mycorrhizal fungus Paxillus involutus degraded lignin more readily than those which are normally considered to be obligately mycorrhizal fungi such as Suillus bovinus and Rhizopogon roseolus. The importance of these lignin degrading capabilities is discussed in relation to the predominance of specific mycorrhiza forms along a gradient of increasing organic matter and hence lignin content of soil.  相似文献   

7.
Mycorrhiza samples of neighbouring Quercus ilex and Erica arborea plants collected in a postcutting habitat were processed to see whether plants differing in mycorrhizal status harbour the same root endophytes. Three experiments were performed in parallel: (i) isolation, identification and molecular characterization of fungi from surface-sterilized roots of both plant species; (ii) re-inoculation of fungal isolates on axenic E. arborea and Q. ilex seedlings; (iii) direct inoculation of field-collected Q. ilex ectomycorrhizas onto E. arborea seedlings. About 70 and 150 fungal isolates were obtained from roots of Q. ilex and E. arborea, respectively. Among them, Oidiodendron species and five cultural morphotypes of sterile isolates formed typical ericoid mycorrhizas on E. arborea in vitro. Fungi with such mycorrhizal ability were derived from both host plants. Isolates belonging to one of these morphotypes (sd9) also exhibited an unusual pattern of colonization, with an additional extracellular hyphal net. Ericoid mycorrhizas were also readily obtained by direct inoculation of E. arborea seedlings with Q. ilex ectomycorrhizal tips. Polymerase chain-restriction fragment length polymorphism and random amplified polymorphic DNA analyses of the shared sterile morphotypes demonstrate, in the case of sd9, the occurrence of the same genet on the two host plants. These results indicate that ericoid mycorrhizal fungi associate with ectomycorrhizal roots, and the ecological significance of this finding is discussed.  相似文献   

8.
Ectomycorrhizal fungi constitute an important component of forest ecosystems that enhances plant nutrition and resistance against stresses. Diversity of ectomycorrhizal (EcM) fungi is, however, affected by host plant diversity and soil heterogeneity. This study provides information about the influence of host plants and soil resources on the diversity of ectomycorrhizal fungal fruiting bodies from rainforests of the Democratic Republic of the Congo. Based on the presence of fungal fruiting bodies, significant differences in the number of ectomycorrhizal fungi species existed between forest stand types (p < 0.001). The most ectomycorrhizal species‐rich forest was the Gilbertiodendron dewevrei‐dominated forest (61 species). Of all 93 species of ectomycorrhizal fungi, 19 demonstrated a significant indicator value for particular forest stand types. Of all analysed edaphic factors, the percentage of silt particles was the most important parameter influencing EcM fungi host plant tree distribution. Both host trees and edaphic factors strongly affected the distribution and diversity of EcM fungi. EcM fungi may have developed differently their ability to successfully colonise root systems in relation to the availability of nutrients.  相似文献   

9.
Introduction of exotic plants change soil microbial communities which may have detrimental ecological consequences for ecosystems. In this study, we examined the community structure and species richness of ectomycorrhizal (EcM) fungi associated with exotic pine plantations in relation to adjacent native ectomycorrhizal trees in Iran to elucidate the symbiont exchange between distantly related hosts, i.e. Fagales (Fagaceae and Betulaceae) and Pinaceae. The combination of morphological and molecular identification approaches revealed that 84.6 % of species with more than one occurrence (at least once on pines) were shared with native trees and only 5.9 % were found exclusively on pine root tips. The community diversity of ectomycorrhizal fungi in the pine plantations adjacent to native EcM trees was comparable to their adjacent native trees, but the isolated plantations hosted relatively a species-poor community. Specific mycobionts of conifers were dominant in the isolated plantation while rarely found in the plantations adjacent to native EcM trees. These data demonstrate the importance of habitat isolation and dispersal limitation of EcM fungi in their potential of host range expansion. The great number of shared and possibly compatible symbiotic species between exotic Pinaceae and local Fagales (Fagaceae and Betulaceae) may reflect their evolutionary adaptations and/or ancestral compatibility with one another.  相似文献   

10.
Recent studies have shown that mycorrhizal trees can greatly influence soil microbial communities, which in turn play important roles in the function offorest ecosystems. However, there is lack of understanding how the composition of trees with different mycorrhizal types affects soil microbial communities. Here, we collected 1606 soil samples from a 25-ha subtropical forest plot to investigate how the proportion of arbuscular mycorrhizal (AM) versus ectomycorrhizal (EcM) trees mediated soil microbial assemblages. Results showed the alpha diversities of both soil fungal and bacterial communities were significantly positively correlated with the ratio of AM/EcM trees. The AM/EcM tree ratio was important to the fungal community assembly, whereas soil pH was key to the bacterial communities. The increase in the AM/EcM tree ratio decreased the importance of stochastic forces in assembling fungal communities, while it had no significant effect on the bacterial communities. The differential importance of the AM/EcM tree ratio to fungal and bacterial communities highlights the role of mycorrhiza-associated tree composition in regulating soil microbial communities. This finding suggests that forests with different AM/EcM tree ratios would have different soil microbial communities, potentially leading to differences in soil nutrient cycling and in return different tree diversity and forest productivity.  相似文献   

11.
Revegetation following dam removal projects may depend on recovery of arbuscular mycorrhizal (AM) and ectomycorrhizal (EM) fungal communities, which perform valuable ecosystem functions. This study assessed the availability and function of AM and EM fungi for plants colonizing dewatered reservoirs following a dam removal project on the Elwha River, Olympic Peninsula, Washington, United States. Availability was assessed via AM fungal spore density in soils and EM root tip colonization of Salix sitchensis (Sitka willow) in an observational field study. The effect of mycorrhizal fungi from 4 sources (reservoir soils, commercial inoculum, and 2 mature plant community soils) on growth and nutrient status of S. sitchensis was quantified in a greenhouse study. AM fungal spores and EM root tips were present in all field samples. In the greenhouse, plants receiving reservoir soil inoculum had only incipient mantle formation, while plants receiving inoculum from mature plant communities had fully formed EM root tips. EM formation corresponded with alleviation of phosphorus stress in plants (lower shoot nitrogen:phosphorus). Thus, revegetating plants have access to AM and EM fungi following dam removal, and EM formation may be especially important for plant P uptake in reservoir soils. However, availability of mycorrhizal fungi declines with distance from established plant communities. Furthermore, EM fungal communities in recently dewatered reservoirs may not be as effective at forming beneficial mycorrhizae as those from mature plant communities. Whole soil inoculum from mature plant communities may be important for the success of revegetating plants and recovery of mycorrhizal fungal communities.  相似文献   

12.
Bell  Tina L.  Adams  Mark A. 《Plant Ecology》2004,171(1-2):35-52
As a potential means of monitoring functional properties of plantations of Pinus pinaster and Pinus radiata established as part of land rehabilitation in the wheatbelt of Western Australia, we examined aspects of the ecophysiology of ectomycorrhizal fungi associated with tree roots. A single species of ectomycorrhizal fungi, Rhizopogon roseolus, dominated the mycorrhizal flora. Sporocarps of Rhizopogon roseolus appeared with the onset of winter rains in May, increased in number and total biomass to peak in September, and decreased to negligible levels at the beginning of the summer drought in December. A greater number of sporocarps, and consequently a greater biomass of sporocarp tissue, was associated with roots of P. radiata than P. pinaster. A similar seasonal pattern of mycorrhizal root infection was determined by counts of individual ectomycorrhizal root tips from bimonthly collection of soil core samples. At the low rainfall (380 mm annually) site, greater numbers of live root tips were more strongly correlated with soil moisture than organic matter content of soil. In contrast, in wetter areas closer to Perth (800 mm annually), highest numbers of active root tips and greatest amounts of organic matter were both within 0–10 cm depths. Results suggest an overriding importance of soil moisture rather than nutrient status of the soil as the key determinant of spatial and temporal distribution of the fungus. Results from a range of assays determining enzyme activity of soil (protease, phosphomonoesterase, cellulase, L-asparaginase, L-glutaminase and β-glucosidase) surrounding mycorrhizal roots indicated seasonal patterns to be similar to those described for reproductive activity of mycorrhizal fungi. Factors responsible for patterns of seasonal activity and distribution of ectomycorrhizal roots are discussed in terms of managing systems in order to maximise tree growth and form while effectively restoring soil water balance.  相似文献   

13.
Linking roots and ectomycorrhizas (EcM) to individual host trees in the field is required to test whether individual trees support different ectomycorrhizal communities. Here we describe a method that identifies the source of EcM roots by PCR of polymorphic pine nuclear microsatellite loci using fluorescently labelled primers and high-throughput fragment analysis. ITS-PCR can also be performed on the same EcM DNA extract for fungal identification. The method was tested on five neighbouring Scots pine (Pinus sylvestris var scotica) trees in native woodland. Successful host tree identification from DNA extracts of EcM root tips was achieved for 93% of all root fragments recovered from soil cores. It was estimated that each individual mature pine sampled was colonised by between 15 and 19 EcM fungi. The most abundant fungal species were found on all five trees, and within the constraints of the sampling scheme, no differences between trees in EcM fungal community structure or composition were detected.  相似文献   

14.
Ericaceous dwarf shrubs including Calluna vulgaris and Vaccinium spp. occur both in open heathland communities and in forest ecosystems as understory vegetation. Ericaceous shrubs were once thought to form ericoid mycorrhizal associations with a relatively narrow range of ascomycetous fungi closely related to, and including, Rhizoscyphus ericae. However, perceptions have recently changed since the realization that a broader range of ascomycete fungi, and in some cases basidiomycete fungi, can also form associations with the roots of ericaceous plants. We used a combination of molecular approaches, including denaturing gradient gel electrophoresis, terminal restriction fragment length polymorphism, cloning and sequencing, to investigate the diversity of fungi associated with C. vulgaris roots collected across a heathland/native Scots pine forest vegetation gradient. We also determined differences in fungal community composition between roots of co-occurring C. vulgaris and Vaccinium myrtillus in the forest understory. Collectively, the data show that a large diversity of potentially ericoid mycorrhizal fungal taxa associate with roots of C. vulgaris and V. myrtillus, and that ascomycetes were about 2.5 times more frequent than basidiomycetes. The assemblages of fungi associated with C. vulgaris and V. myrtillus were different. In addition, the community of fungi associated with C. vulgaris hair roots was different for samples collected from the forest, open heathland and a transition zone between the two. This separation was partly, but not entirely, due to the occurrence of typical ectomycorrhizal basidiomycetes associated with the hair roots of C. vulgaris in the forest understory. These data demonstrate that forest understory ericaceous shrubs associate with a diverse range of ascomycete and basidiomycete taxa, including typical ectomycorrhizal basidiomycetes.  相似文献   

15.
Plant root associations with microbes such as mycorrhizal fungi or N-fixing bacteria enable ecosystems to tap pools of nitrogen (N) that might otherwise be inaccessible, including atmospheric N or N in large soil organic molecules. Such microbially assisted N-foraging strategies may be particularly important in late-successional retrogressive ecosystems where productivity is low and soil nutrients are scarce. Here, we use natural N-stable isotopic composition to constrain pathways of N supplies to different plant functional groups across a well-studied natural soil fertility gradient that includes a highly retrogressive stage. We demonstrate that ectomycorrhizal fungi, ericoid mycorrhizal fungi, and N-fixing bacteria support forest N supplies at all stages of ecosystem succession, from relatively young, N-rich/phosphorus (P)-rich sites, to ancient sites (ca. 500 ky) where both N supplies and P supplies are exceedingly low. Microbially mediated N sources are most important in older ecosystems with very low soil nutrient availability, accounting for 75–96% of foliar N at the oldest, least fertile sites. These isotopically ground findings point to the key role of plant–microbe associations in shaping ecosystem processes and functioning, particularly in retrogressive-phase forest ecosystems.  相似文献   

16.
We studied the effect of ectomycorrhizal fungi on bacterial communities colonizing roots of Douglas fir (Pseudotsuga menziesii). Mycorrhizal tips were cleaned of soil and separated based on gross morphological characteristics. Sequencing of the internal transcribed spacers of the nuclear rRNA gene cluster indicated that the majority of the tips were colonized by fungi in the Russulaceae, with the genera Russula and Lactarius comprising 70% of the tips. Because coamplification of organellar 16S rRNA genes can interfere with bacterial community analysis of root tips, we developed and tested a new primer pair that permits amplification of bacterial 16S rRNA genes but discriminates more effectively against organellar sequences than commonly used bacterial primer sets. We then used terminal restriction fragment length polymorphism (T-RFLP) and sequence analysis of the 16S rRNA gene to examine differences in bacterial communities associated with the mycorrhizal tips. Cluster analysis of T-RFLP profiles indicated that there were different bacterial communities among the root tips; however, the communities did not seem to be affected by the taxonomic identity of the ectomycorrhizal fungi. Terminal restriction fragment profiling and sequencing of cloned partial 16S rRNA genes indicated that most bacteria on the ectomycorrhizal tips were related to the Alphaproteobacteria and the Bacteroidetes group.  相似文献   

17.
Ericaceae associate with a wide spectrum of root mycobionts, but the most common are ascomycetous ericoid mycorrhizal fungi and dark septate endophytes (DSE), followed by basidiomycetous fungi and glomeracean arbuscular mycorrhizal fungi. We investigated distribution and morphological diversity of ericoid mycorrhizae (ErM), DSE associations, ectomycorrhizae (EcM) and arbuscular mycorrhizae (AM) in hair roots of six European native Rhododendron species and found that i) while EcM and AM were absent, ErM and DSE associations were simultaneously present in all screened plants; ii) their levels were negatively correlated, suggesting Ericaceae preference for certain root-fungus association in certain habitats; iii) the highest ErM colonization occurred at sites in southern and central Europe, while the highest DSE colonization was found in a subarctic site in northern Finland and in a subalpine site in the Carpathians, suggesting a latitudinal/altitudinal shift in Ericaceae root-fungus associations; iv) some mycelia could simultaneously form structures corresponding to ErM and DSE association, which occasionally resulted in a unique ectendomycorrhizal colonization comprising an intercellular parenchymatous net and intracellular hyphal coils. These results indicate frequent interactions between ErM fungi and DSE in roots of European rhododendrons and a morphological continuum between ErM and DSE associations. The new ectendomycorrhizal type deserves further investigation.  相似文献   

18.
Waiting for fungi: the ectomycorrhizal invasion of lowland heathlands   总被引:1,自引:1,他引:0  
1.  In England, the loss of lowland heathland, a habitat of global conservation importance, is primarily due to the invasion of birch and pine. This encroachment has been researched in depth from a plant perspective but little is known about the role of mycorrhizal fungi. In lowland heathlands the resident dwarf shrubs form ericoid mycorrhizas whereas invading trees form ectomycorrhizas. Therefore, tree encroachment into heathlands can be regarded as the replacement of a resident mycorrhizal community by an invading one.
2 . This study examined how fungi form mycorrhizas with Betula and Pinus in lowland heathlands. We addressed the question of whether there are mycorrhizal fungi that mediate invasion using a molecular ecology approach to compare the mycorrhizal inoculum potential of soil at three levels of invasion (uninvaded heathland, invaded heathland and woodland) and the fungi forming mycorrhizas on tree seedlings and trees across diverse sites.
3.  We show that in lowland heathlands: (i) seedlings have severely limited access to ectomycorrhizal fungi relative to woodlands, (ii) there are few keystone spore-dispersed ectomycorrhizal fungi that can mediate tree invasion, (iii) tree seedlings can remain non-mycorrhizal for at least one year when no inoculum is present, even near saplings, and (iv) mycorrhizal seedlings achieve greater biomass than non-mycorrhizal seedlings. Within uninvaded heathland we detected only Rhizopogon luteolus , Suillus variegatus , S. bovinus ( Pinus symbionts) and Laccaria proxima (primarily a Betula symbiont).
4. Synthesis . Overall, ectomycorrhizal inoculum in lowland heathlands is rare; most tree seedlings growing in heathland soil are not mycorrhizal due to limited spore dispersal, poorly developed spore banks and weak common mycorrhizal networks. These seedlings can persist awaiting mycorrhization to boost their growth.  相似文献   

19.
The genera Dicymbe and Aldina (Fabaceae) host ectomycorrhizal fungi (EcM) and are common in white sand forests (WSFs), a highly specialized habitat with a high level of plant endemism compared with terra-firme forests. In this study, we visited four times a 1-ha permanent plot established in a small patch of a WSF in the south of Colombia Amazonia. Forty-eight species of EcM fungi were recovered from sporocarps and 15 ITS species-level were detected from root tips. Seventeen species were new reports to Colombia and seven corresponded to undescribed species. These results confirm that this WSF supports a significant EcM fungal diversity. Most of the species found in this study have been previously reported to be associated with other legume and/or dipterocarp species from geographically distant forests. The long-distance occurrence combined with low host specificity, suggest the possibility of gene flow between geographically distant populations of EcM fungi in neotropical lowland rainforests.  相似文献   

20.
Four in vitro experiments were set up to verify the colonization potential of ectomycorrhizal (EcM) Cenococcum geophilum FR. (strain CGE-4), saprotrophic Geomyces pannorum (LINK) SIGLER & CARMICHAEL (GPA-1) and a frequent root-associated, potentially ericoid mycorrhiza (ErM)-forming Meliniomyces variabilis Hambleton & Sigler (MVA-1) in roots of Rhododendron and Vaccinium. A typical ErM fungus, Rhizoscyphus ericae (Read) Zhuang & Korf (RER-1), was included for comparison. All fungal strains intracellularly colonized rooted Vaccinium microcuttings: GPA-1 occasionally produced hyphal loops similar to ErM, MVA-1 and RER-1 exhibited a typical ErM colonization pattern. CGE-4 hyphae grew vigorously on and around newly formed roots and rarely penetrated turgescent rhizodermal cells forming intracellular loose loops. Rooting of Rhododendron sp. microcuttings was not promoted by any fungal strain except CGE-4, which also promoted the most vigorous growth of Rhododendron ponticum L. seedlings. The widespread EcM fungus C. geophilum has a potential to colonize non-EcM roots and support their development which may influence overall growth of ericaceous plants. As shown for G. pannorum, structures resembling ErM may be formed by fungi that are to date not regarded as ericoid mycorrhizal.  相似文献   

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