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Background: Interactions between plants and diverse root-associated fungi are essential drivers of forest ecosystem dynamics. The symbiosis is potentially dependent on multiple ecological factors/processes such as host/symbiont specificity, background soil microbiome, inter-root dispersal of symbionts, and fungus–fungus interactions within roots. Nonetheless, it has remained a major challenge to reveal the mechanisms by which those multiple factors/processes determine the assembly of root-associated fungal communities. Based on the framework of joint species distribution modeling, we examined 1,615 root-tips samples collected in a cool-temperate forest to reveal how root-associated fungal community structure was collectively formed through filtering by host plants, associations with background soil fungi, spatial autocorrelation, and symbiont–symbiont interactions. In addition, to detect fungi that drive the assembly of the entire root-associated fungal community, we inferred networks of direct fungus–fungus associations by a statistical modeling that could account for implicit environmental effects. Results: The fine-scale community structure of root-associated fungi were best explained by the statistical model including the four ecological factors/processes. Meanwhile, among partial models, those including background soil fungal community structure and within-root fungus–fungus interactions showed the highest performance. When fine-root distributions were examined, ectomycorrhizal fungi tended to show stronger associations with background soil community structure and spatially autocorrelated patterns than other fungal guilds. In contrast, the distributions of root-endophytic fungi were inferred to depend greatly on fungus–fungus interactions. An additional statistical analysis further suggested that some endophytic fungi, such as Phialocephala and Leptodontidium, were placed at the core positions within the web of direct associations with other root-associated fungi. Conclusion: By applying emerging statistical frameworks to intensive datasets of root-associated fungal communities, we demonstrated background soil fungal community structure and fungus–fungus associations within roots, as well as filtering by host plants and spatial autocorrelation in ecological processes, could collectively drive the assembly of root-associated fungi. We also found that basic assembly rules could differ between mycorrhizal and endophytic fungi, both of which were major components of forest ecosystems. Consequently, knowledge of how multiple ecological factors/processes differentially drive the assembly of multiple fungal guilds is indispensable for comprehensively understanding the mechanisms by which terrestrial ecosystem dynamics are organized by plant–fungal symbiosis. [ABSTRACT FROM AUTHOR]

Highlights: Urban trees in urban environments are commonly exposed to stresses, contributing to their health decline; Ectomycorrhizal fungi improve the vigour and resilience of Tilia seedlings, but their performance is dependent on soil pH. Inoculation promotes growth in height, biomass production, mycorrhization status, nutrient uptake, and proline accumulation, and these effects dependent on fungi species and soil pH; Inoculation with EcM can be a strategy to produce healthier saplings that when transplanted to the urban environment will have a better performance. Colonization by Ectomycorrhizal (EcM) fungi is key for the health and performance of plants under different stress scenarios, such as those faced by trees in urban environments. Because urban environments can be lacking EcM fungi, we here assessed the benefits of inoculating Tilia tomentosa seedlings in a pre-transplantation nursery context with the EcM fungi Lactarius deliciosus and Paxillus involutus, using substrates of different pH and facing water-stress. P. involutus had a more evident positive effect in T. tomentosa seedlings and had a good performance in both acidic and alkaline substrate. In acidic substrate the fungus increased the plant height by 0.91-fold, increased the mycorrhization rate by 3.23-fold, expansion rate by 5.03-fold and formation of secondary roots by 0.46-fold, compared to the non-inoculated control. This species also improved the phosphorus content of leaves, which revealed a promotion of nutrient uptake. In alkaline substrate P. involutus increased root dry weight by 3.92-fold and the mycorrhization parameters. In contrast, L. deliciosus only had a positive effect in the improvement of mycorrhization and expansion rates and phosphorus content in the root, effects visible only in alkaline substrate. When exposed to water-stress the increase of proline content was visible in acidic substrate for both fungi, L. deliciosus and P. involutus, and in alkaline substrate for the fungus P. involutus, a response indicative of the enhancement of defenses in stressing scenarios such as water scarcity. We conclude that fungal inoculation improves the vigour and resilience of Tilia seedlings and that it is of utmost importance to select a suitable EcM fungus and to consider the soil pH of the transplanting site. The inoculation approach can be a valuable tool to produce robust seedlings which may have a better performance when transplanted to the challenging urban environment. [ABSTRACT FROM AUTHOR]

The genus Cortinarius (Agaricales, Basidiomycota) is one of the most species-rich fungal genera, with thousands of species reported. Cortinarius species are important ectomycorrhizal fungi and form associations with many vascular plants globally. Until recently Cortinarius was the single genus of the family Cortinariaceae, despite several attempts to provide a workable, lower-rank hierarchical structure based on subgenera and sections. The first phylogenomic study for this group elevated the old genus Cortinarius to family level and the family was split into ten genera, of which seven were described as new. Here, by careful re-examination of the recently published phylogenomic dataset, we detected extensive gene-tree/species-tree conflicts using both concatenation and multispecies coalescent approaches. Our analyses demonstrate that the Cortinarius phylogeny remains unresolved and the resulting phylogenomic hypotheses suffer from very short and unsupported branches in the backbone. We can confirm monophyly of only four out of ten suggested new genera, leaving uncertain the relationships between each other and the general branching order. Thorough exploration of the tree space demonstrated that the topology on which Cortinarius revised classification relies on does not represent the best phylogenetic hypothesis and should not be used as constrained topology to include additional species. For this reason, we argue that based on available evidence the genus Cortinarius should not (yet) be split. Moreover, considering that phylogenetic uncertainty translates to taxonomic uncertainty, we advise for careful evaluation of phylogenomic datasets before proposing radical taxonomic and nomenclatural changes. [ABSTRACT FROM AUTHOR]

Background: The colonization of land and the diversification of terrestrial plants is intimately linked to the evolutionary history of their symbiotic fungal partners. Extant representatives of these fungal lineages include mutualistic plant symbionts, the arbuscular mycorrhizal (AM) fungi in Glomeromycota and fine root endophytes in Endogonales (Mucoromycota), as well as fungi with saprotrophic, pathogenic and endophytic lifestyles. These fungal groups separate into three monophyletic lineages but their evolutionary relationships remain enigmatic confounding ancestral reconstructions. Their taxonomic ranks are currently fluid. Results: In this study, we recognize these three monophyletic linages as phyla, and use a balanced taxon sampling and broad taxonomic representation for phylogenomic analysis that rejects a hard polytomy and resolves Glomeromycota as sister to a clade composed of Mucoromycota and Mortierellomycota. Low copy numbers of genes associated with plant cell wall degradation could not be assigned to the transition to a plant symbiotic lifestyle but appears to be an ancestral phylogenetic signal. Both plant symbiotic lineages, Glomeromycota and Endogonales, lack numerous thiamine metabolism genes but the lack of fatty acid synthesis genes is specific to AM fungi. Many genes previously thought to be missing specifically in Glomeromycota are either missing in all analyzed phyla, or in some cases, are actually present in some of the analyzed AM fungal lineages, e.g. the high affinity phosphorus transporter Pho89. Conclusion: Based on a broad taxon sampling of fungal genomes we present a well-supported phylogeny for AM fungi and their sister lineages. We show that among these lineages, two independent evolutionary transitions to mutualistic plant symbiosis happened in a genomic background profoundly different from that known from the emergence of ectomycorrhizal fungi in Dikarya. These results call for further reevaluation of genomic signatures associated with plant symbiosis. [ABSTRACT FROM AUTHOR]

The genus Hebeloma is renowned as difficult when it comes to species determination. Historically, many dichotomous keys have been published and used with varying success rate. Over the last 20 years the authors have built a database of Hebeloma collections containing not only metadata but also parametrized morphological descriptions, where for about a third of the cases micromorphological characters have been analysed and are included, as well as DNA sequences for almost every collection. The database now has about 9000 collections including nearly every type collection worldwide and represents over 120 different taxa. Almost every collection has been analysed and identified to species using a combination of the available molecular and morphological data in addition to locality and habitat information. Based on these data an Artificial Intelligence (AI) machine-learning species identifier has been developed that takes as input locality data and a small number of the morphological parameters. Using a random test set of more than 600 collections from the database, not utilized within the set of collections used to train the identifier, the species identifier was able to identify 77% correctly with its highest probabilistic match, 96% within its three most likely determinations and over 99% of collections within its five most likely determinations. [ABSTRACT FROM AUTHOR]

Russula albonigra is considered a well-known species, morphologically delimited by the context of the basidiomata blackening without intermediate reddening, and the menthol-cooling taste of the lamellae. It is supposed to have a broad ecological range and a large distribution area. A thorough molecular analysis based on four nuclear markers (ITS, LSU, RPB2 and TEF1-α) shows this traditional concept of R. albonigra s. lat. represents a species complex consisting of at least five European, three North American, and one Chinese species. Morphological study shows traditional characters used to delimit R. albonigra are not always reliable. Therefore, a new delimitation of the R. albonigra complex is proposed and a key to the described European species of R. subgen. Compactae is presented. A lectotype and an epitype are designated for R. albonigra and three new European species are described: R. ambusta, R. nigrifacta, and R. ustulata. Different thresholds of UNITE species hypotheses were tested against the taxonomic data. The distance threshold of 0.5% gives a perfect match to the phylogenetically defined species within the R. albonigra complex. Publicly available sequence data can contribute to species delimitation and increase our knowledge on ecology and distribution, but the pitfalls are short and low quality sequences. [ABSTRACT FROM AUTHOR]

Background: A better understanding of non-structural carbohydrate (NSC) dynamics in trees under drought stress is critical to elucidate the mechanisms underlying forest decline and tree mortality from extended periods of drought. This study aimed to assess the contribution of ectomycorrhizal (ECM) fungus (Suillus variegatus) to hydraulic function and NSC in roots, stems, and leaves of Pinus tabulaeformis subjected to different water deficit intensity. We performed a continuous controlled drought pot experiment from July 10 to September 10, 2019 using P. tabulaeformis seedlings under 80, 40, and 20% of the field moisture capacity that represented the absence of non-drought, moderate drought, and severe drought stress, respectively. Results: Results indicated that S. variegatus decreased the mortality rate and increased height, root biomass, and leaf biomass of P. tabulaeformis seedlings under moderate and severe drought stress. Meanwhile, the photosynthetic rates, stomatal conductance, and transpiration rates of P. tabulaeformis were significantly increased after S. variegatus inoculation. Moreover, the inoculation of S. variegatus also significantly increased the NSC concentrations of all seedling tissues, enhanced the soluble sugars content, and increased the ratios of soluble sugars to starch on all tissues under severe drought. Overall, the inoculation of S. variegatus has great potential for improving the hydraulic function, increasing the NSC storage, and improving the growth of P. tabulaeformis under severe drought. Conclusions: Therefore, the S. variegatus can be used as a potential application strain for ecological restoration on arid regions of the Loess Plateau, especially in the P. tabulaeformis woodlands. [ABSTRACT FROM AUTHOR]

Background: Achlorophyllous orchids are mycoheterotrophic plants, which lack photosynthetic ability and associate with fungi to acquire carbon from different environmental sources. In tropical latitudes, achlorophyllous forest orchids show a preference to establish mycorrhizal relationships with saprotrophic fungi. However, a few of them have been recently found to associate with ectomycorrhizal fungi and there is still much to be learned about the identity of fungi associated with tropical orchids. The present study focused on mycorrhizal diversity in the achlorophyllous orchid C. inverta, an endangered species, which is endemic to southern China. The aim of this work was to identify the main mycorrhizal partners of C. inverta in different plant life stages, by means of morphological and molecular methods. Results: Microscopy showed that the roots of analysed C. inverta samples were extensively colonized by fungal hyphae forming pelotons in root cortical cells. Fungal ITS regions were amplified by polymerase chain reaction, from DNA extracted from fungal mycelia isolated from orchid root samples, as well as from total root DNA. Molecular sequencing and phylogenetic analyses showed that the investigated orchid primarily associated with ectomycorrhizal fungi belonging to a narrow clade within the family Ceratobasidiaceae, which was previously detected in a few fully mycoheterotrophic orchids and was also found to show ectomycorrhizal capability on trees and shrubs. Russulaceae fungal symbionts, showing high similarity with members of the ectomycorrhizal genus Russula, were also identified from the roots of C. inverta, at young seedling stage. Ascomycetous fungi including Chaetomium, Diaporthe, Leptodontidium, and Phomopsis genera, and zygomycetes in the genus Mortierella were obtained from orchid root isolated strains with unclear functional role. Conclusions: This study represents the first assessment of root fungal diversity in the rare, cryptic and narrowly distributed Chinese orchid C. inverta. Our results provide new insights on the spectrum of orchid-fungus symbiosis suggesting an unprecedented mixed association between the studied achlorophyllous forest orchid and ectomycorrhizal fungi belonging to Ceratobasidiaceae and Russulaceae. Ceratobasidioid fungi as dominant associates in the roots of C. inverta represent a new record of the rare association between the identified fungal group and fully mycoheterotrophic orchids in nature. [ABSTRACT FROM AUTHOR]

In this study, the mitogenomes of two Paxillus species were assembled, annotated and compared. The two mitogenomes of Paxillus involutus and P. rubicundulus comprised circular DNA molecules, with the size of 39,109 bp and 41,061 bp, respectively. Evolutionary analysis revealed that the nad4L gene had undergone strong positive selection in the two Paxillus species. In addition, 10.64 and 36.50% of the repetitive sequences were detected in the mitogenomes of P. involutus and P. rubicundulus, respectively, which might transfer between mitochondrial and nuclear genomes. Large-scale gene rearrangements and frequent intron gain/loss events were detected in 61 basidiomycete species, which revealed large variations in mitochondrial organization and size in Basidiomycota. In addition, the insertion sites of the basidiomycete introns were found to have a base preference. Phylogenetic analysis of the combined mitochondrial gene set gave identical and well-supported tree topologies, indicating that mitochondrial genes were reliable molecular markers for analyzing the phylogenetic relationships of Basidiomycota. This study is the first report on the mitogenomes of Paxillus, which will promote a better understanding of their contrasted ecological strategies, molecular evolution and phylogeny of these important ectomycorrhizal fungi and related basidiomycete species. [ABSTRACT FROM AUTHOR]

Context Quercus suber L. plays a key ecological and socio-economical role in the Iberian Peninsula. Symbiotic ectomycorrhizal fungi-ECM are crucial partners of several tree species, and assessing the efficacy of bioinoculants at nursery stage helps devising tools to increase plant resilience. Aims The aim of this study was to compare the effects of two inocula formulations of mixed ECM fungi and bacteria on the quality of seedlings produced in two forest nurseries, differing in environmental conditions and forest embedment. Methods Quercus suber L. seedlings were inoculated with a commercial product containing Pisolithus tinctorius (Pers) Coker & Couch, Scleroderma sp., and six bacterial species and with a non-commercial fungal and bacterial dual inoculum (Suillus granulatus (L.) Roussel + Mesorhizobium sp.). Biometric and nutritional parameters and morphological quality indexes were determined on seedlings. The ECM community was assessed by denaturing gradient gel electrophoresis and cloning-sequencing. Results In both nurseries, the seedling quality index in inoculated was up to 2-fold higher than in non-inoculated seedlings. Plant biomass differed significantly among nurseries. The inoculum influenced the profile of the fungal community. S. granulatus and P. tinctorius persisted for 6 months in the inoculated seedlings. Conclusion The nursery ecosystem influenced plant growth. Inoculation treatments increased plant performance; however, the dual inoculum resulted in more consistent improvements of Q. suber at nursery stage, highlighting the importance of inocula selection. [ABSTRACT FROM AUTHOR]

Two new species in the genus Auritella (Inocybaceae) are described as new from tropical rainforest in Cameroon. Descriptions, photographs, line drawings, and a worldwide taxonomic key to the described species of Auritella are presented. Phylogenetic analysis of 28S rDNA and rpb2 nucleotide sequence data suggests at least five phylogenetic species that can be ascribed to Auritella occur in the region comprising Cameroon and Gabon and constitute a strongly supported monophyletic subgroup within the genus. Phylogenetic analysis of ITS data supports the conspecificity of numerous collections attributed to the two new species as well as the monophyly of Australian species of Auritella. This work raises the known number of described species of Auritella to thirteen worldwide, four of which occur in tropical Africa, one in tropical India, and eight in temperate and tropical regions of Australia. This is the first study to confirm an ectomycorrhizal status of Auritella using molecular data. [ABSTRACT FROM AUTHOR]

Background: Coniophora olivacea is a basidiomycete fungus belonging to the order Boletales that produces brown-rot decay on dead wood of conifers. The Boletales order comprises a diverse group of species including saprotrophs and ectomycorrhizal fungi that show important differences in genome size. Results: In this study we report the 39.07-megabase (Mb) draft genome assembly and annotation of C. olivacea. A total of 14,928 genes were annotated, including 470 putatively secreted proteins enriched in functions involved in lignocellulose degradation. Using similarity clustering and protein structure prediction we identified a new family of 10 putative lytic polysaccharide monooxygenase genes. This family is conserved in basidiomycota and lacks of previous functional annotation. Further analyses showed that C. olivacea has a low repetitive genome, with 2.91% of repeats and a restrained content of transposable elements (TEs). The annotation of TEs in four related Boletales yielded important differences in repeat content, ranging from 3.94 to 41.17% of the genome size. The distribution of insertion ages of LTRretrotransposons showed that differential expansions of these repetitive elements have shaped the genome architecture of Boletales over the last 60 million years. Conclusions: Coniophora olivacea has a small, compact genome that shows macrosynteny with Coniophora puteana. The functional annotation revealed the enzymatic signature of a canonical brown-rot. The annotation and comparative genomics of transposable elements uncovered their particular contraction in the Coniophora genera, highlighting their role in the differential genome expansions found in Boletales species. [ABSTRACT FROM AUTHOR]

Kombocles bakaiana gen. sp. nov. is described as new to science. This sequestrate, partially hypogeous fungus was collected around and within the stilt root system of an ectomycorrhizal (ECM) tree of the genus Uapaca (Phyllanthaceae) in a Guineo-Congolian mixed tropical rainforest in Cameroon. Molecular data place this fungus in Boletaceae (Boletales, Agaricomycetes, Basidiomycota) with no clear relationship to previously described taxa within the family. Macro- and micromorphological characters, habitat, and DNA sequence data are provided. Unique morphological features and a molecular phylogenetic analysis of 304 sequences across the Boletales justify the recognition of the new taxa. Kombocles bakaiana is the fourth sequestrate Boletaceae described from the greater African tropics, and the first to be described from Cameroon. [ABSTRACT FROM AUTHOR]

Jimtrappea guyanensis gen. sp. nov., Castellanea pakaraimophila gen. sp. nov., and Costatisporus cyanescens gen. sp. nov. are described as new to science. These sequestrate, hypogeous fungi were collected in Guyana under closed canopy tropical forests in association with ectomycorrhizal (ECM) host tree genera Dicymbe (Fabaceae subfam. Caesalpinioideae), Aldina (Fabaceae subfam. Papilionoideae), and Pakaraimaea (Dipterocarpaceae). Molecular data place these fungi in Boletaceae (Boletales, Agaricomycetes, Basidiomycota) and inform their relationships to other known epigeous and sequestrate taxa within that family. Macro- and micromorphological characters, habitat, and multi-locus DNA sequence data are provided for each new taxon. Unique morphological features and a molecular phylogenetic analysis of 185 taxa across the order Boletales justify the recognition of the three new genera. [ABSTRACT FROM AUTHOR]

Key message: The ICP Forests network can be a platform for large-scale mycorrhizal studies. Mapping and monitoring of mycorrhizas have untapped potential to inform science, management, conservation and policy regarding distributions, diversity hotspots, dominance and rarity, and indicators of forest changes. Context: A dearth of information about fungi at large scales has severely constrained scientific, forest management, fungal conservation and environmental policy efforts worldwide. Nonetheless, fungi fulfil critical functional roles in our changing environments and represent a considerable proportion of terrestrial biodiversity. Mycorrhizal fungi are increasingly viewed as a major functional guild across forest ecosystems, and our ability to study them is expanding rapidly. Aims: This study aimed to discuss the potential for starting a mycorrhizal monitoring programme built upon the existing forest monitoring network, raise questions, propose hypotheses and stimulate further discussion. Results: An overview of the state-of-the-art regarding forest ectomycorrhizal ecology raises questions and recommendations for scaling up mycorrhizal assessments aimed at informing a variety of stakeholders, with a new focus on conservation and policy. Conclusion: Fungal research and conservation are areas that can be informed by ICP Forests and may lead to useful spin-offs; research linked to long-term forest monitoring plots will enhance the relevance of science and conservation. [ABSTRACT FROM AUTHOR]

Background: Pedunculate oak, Quercus robur is an abundant forest tree species that hosts a large and diverse community of beneficial ectomycorrhizal fungi (EMFs), whereby ectomycorrhiza (EM) formation is stimulated by mycorrhiza helper bacteria such as Streptomyces sp. AcH 505. Oaks typically grow rhythmically, with alternating root flushes (RFs) and shoot flushes (SFs). We explored the poorly understood mechanisms by which oaks integrate signals induced by their beneficial microbes and endogenous rhythmic growth at the level of gene expression. To this end, we compared transcript profiles of oak microcuttings at RF and SF during interactions with AcH 505 alone and in combination with the basidiomycetous EMF Piloderma croceum. Results: The local root and distal leaf responses to the microorganisms differed substantially. More genes involved in the recognition of bacteria and fungi, defence and cell wall remodelling related transcription factors (TFs) were differentially expressed in the roots than in the leaves of oaks. In addition, interaction with AcH 505 and P. croceum affected the expression of a higher number of genes during SF than during RF, including AcH 505 elicited defence response, which was attenuated by co-inoculation with P. croceum in the roots during SF. Genes encoding leucine-rich receptor-like kinases (LRR-RLKs) and proteins (LRR-RLPs), LRR containing defence response regulators, TFs from bZIP, ERF and WRKY families, xyloglucan cell wall transglycolases/hydrolases and exordium proteins were differentially expressed in both roots and leaves of plants treated with AcH 505. Only few genes, including specific RLKs and TFs, were induced in both AcH 505 and co-inoculation treatments. Conclusion: Treatment with AcH 505 induces and maintains the expression levels of signalling genes encoding candidate receptor protein kinases and TFs and leads to differential expression of cell wall modification related genes in pedunculate oak microcuttings. Local gene expression response to AcH 505 alone and in combination with P. croceum are more pronounced when roots are in resting stages, possibly due to the fact that non growing roots re-direct their activity towards plant defence rather than growth. [ABSTRACT FROM AUTHOR]

Despite the crucial ecological role of lactarioid taxa (Lactifluus, Lactarius) as common ectomycorrhiza formers in tropical African seasonal forests, their current diversity is not yet adequately assessed. During the last few years, numerous lactarioid specimens have been sampled in various ecosystems from Togo (West Africa). We generated 48 ITS sequences and aligned them against lactarioid taxa from other tropical African ecozones (Guineo-Congolean evergreen forests, Zambezian miombo). A Maximum Likelihood phylogenetic tree was inferred from a dataset of 109 sequences. The phylogenetic placement of the specimens, combined with morpho-anatomical data, supported the description of four new species from Togo within the monophyletic genus Lactifluus: within subgen. Lactifluus (L. flavellus), subgen. Russulopsis (L. longibasidius and L. pectinatus), and subgen. Edules (L. melleus). This demonstrates that the current species richness of the genus is considerably higher than hitherto estimated for African species and, in addition, a need to redefine the subgenera and sections within it. [ABSTRACT FROM AUTHOR]

Context: Increased summer drought is considered as a threat to the regeneration of Pinus sylvestris in the Central Alps. To a certain degree, seedlings are able to mitigate negative effects of drought by altering root/shoot ratios. But, seedlings may also enhance access to water and nutrients by cooperation with ectomycorrhizal fungi. Aims: We tested the importance of both mechanisms for drought resistance of P. sylvestris seedlings during early establishment and assessed whether differences occur between topsoil and deeper soil layers. Methods: Biomass allocation and colonisation of fine roots by ectomycorrhizal fungi were assessed in seedlings grown for 6 months in a common garden under different precipitation scenarios: constant drought (March-September), summer drought (June-September), and wet conditions. Results: Root/shoot ratios increased from 0.6 under wet conditions to 0.8 under drought conditions, irrespective of the onset of the drought (March vs. June). In both drought scenarios, seedlings had shorter roots in the topsoil, increased the number of root tips per root length in both soil layers, but did not alter the colonisation rate of root tips by ectomycorrhizal fungi. Conclusion: We conclude that plasticity in root architecture is an important mechanism for drought resistance of P. sylvestris during early seedling establishment. [ABSTRACT FROM AUTHOR]

Background: Mycoheterotrophic orchids are achlorophyllous plants that obtain carbon and nutrients from their mycorrhizal fungi. They often show strong preferential association with certain fungi and may obtain nutrients from surrounding photosynthetic plants through ectomycorrhizal fungi. Gastrodia is a large genus of mycoheterotrophic orchids in Asia, but Gastrodia species' association with fungi has not been well studied. We asked two questions: (1) whether certain fungi were preferentially associated with G. flavilabella, which is an orchid in Taiwan and (2) whether fungal associations of G. flavilabella were affected by the composition of fungi in the environment. Results: Using next-generation sequencing, we studied the fungal communities in the tubers of Gastrodia flavilabella and the surrounding soil. We found (1) highly diversified fungi in the G. flavilabella tubers, (2) that Mycena species were the predominant fungi in the tubers but minor in the surrounding soil, and (3) the fungal communities in the G. flavilabella tubers were clearly distinct from those in the surrounding soil. We also found that the fungal composition in soil can change quickly with distance. Conclusions: G. flavilabella was associated with many more fungi than previously thought. Among the fungi in the tuber of G. flavilabella, Mycena species were predominant, different from the previous finding that adult G. elata depends on Armillaria species for nutritional supply. Moreover, the preferential fungus association of G. flavilabella was not significantly influenced by the composition of fungi in the environment. [ABSTRACT FROM AUTHOR]

Context: The requirement for rebuilding forecrop stands besides replacement of meadow vegetation with forest plants and formation of soil humus is the presence of a compatible ectomycorrhizal (ECM) fungal community. Aims: This study aims to assess ectomycorrhizal fungi diversity associated with silver fir ( Abies alba Mill.) seedlings regenerating in silver fir stands and Scots pine forecrops. Methods: One-year-old seedlings were sampled in six study sites: three mature fir forests and three pine forests. ECM fungi were identified by polymerase chain reaction amplification and sequencing of the internal transcribed spacer of rDNA. Results: The mean mycorrhizal colonization exceeded 90 %. Thirty-six ectomycorrhizal taxa were identified in fir stands and 23 in pine forecrops; ten out of these species were common to both stands. The fungal communities were different between study sites ( R = 0.1721, p = 0.0001). Tomentella stuposa was the only species present at all sites. Conclusion: Silver fir seedlings in Scots pine forecrops supported smaller ECM fungal communities than communities identified in mature silver fir stands. Nevertheless, fungal colonization of seedling roots was similar in both cases. This suggests that pine stands afforested on formerly arable land bear enough ECM species to allow survival and growth of silver fir seedlings. [ABSTRACT FROM AUTHOR]

Sclerodermatoid fungi basidiomes were collected from northern Thailand and pure cultures were isolated. The morphology and molecular characteristics identified them as Astraeus odoratus, Phlebopus portentosus, Pisolithus albus and Scleroderma sinnamariense. This study investigated the in vitro ability of selected fungi to produce indole-3-acetic acid (IAA), to solubilize different toxic metal (Co, Cd, Cu, Pb, Zn)-containing minerals, and metal tolerance. The results indicated that all fungi are able to produce IAA in liquid medium. The optimum temperature for IAA production of all fungi was 30 °C, and the optimum concentration of L-tryptophan of Astraeus odoratus, Pisolithus albus and Scleroderma sinnamariense was 2 mg ml. The highest IAA yield (65.29 ± 1.17 μg ml) was obtained from Phlebopus portentosus after 40 days of cultivation in culture medium supplemented with 4 mg ml of L-tryptophan. The biological activity tests of fungal IAA showed that it can simulate coleoptile elongation, and increase seed germination and root length of tested plants. In addition, the metal tolerance and solubilizing activities varied for different minerals and fungal species. The presence of metal minerals affected fungal growth, and cobalt carbonate showed the highest toxicity. The solubilization index decreased when the concentration of metal minerals increased. Astraeus odoratus showed the lowest tolerance to metals. This is the first report of in vitro IAA production, solubilization of insoluble metal minerals and metal tolerance abilities of the tested fungi. [ABSTRACT FROM AUTHOR]

The genus Astraeus is shown to be even more complex than recent studies have found. There are problems defining what the molecular fingerprint is of the generic type species. The present article, based on molecular and morphological information and the classical literature, attempts to throw further light on these important ectomycorrhizal fungi. Our studies go part way in an endeavour to unravel the taxonomy and systematics of this genus, necessitating the recognition of at least three new species. Potential nomenclatural problems are also outlined. [ABSTRACT FROM AUTHOR]

Background: Mycorrhizae, symbiotic interactions between soil fungi and tree roots, are ubiquitous in terrestrial ecosystems. The fungi contribute phosphorous, nitrogen and mobilized nutrients from organic matter in the soil and in return the fungus receives photosynthetically-derived carbohydrates. This union of plant and fungal metabolisms is the mycorrhizal metabolome. Understanding this symbiotic relationship at a molecular level provides important contributions to the understanding of forest ecosystems and global carbon cycling. Results: We generated next generation short-read transcriptomic sequencing data from fully-formed ectomycorrhizae between Laccaria bicolor and aspen (Populus tremuloides) roots. The transcriptomic data was used to identify statistically significantly expressed gene models using a bootstrap-style approach, and these expressed genes were mapped to specific metabolic pathways. Integration of expressed genes that code for metabolic enzymes and the set of expressed membrane transporters generates a predictive model of the ectomycorrhizal metabolome. The generated model of mycorrhizal metabolome predicts that the specific compounds glycine, glutamate, and allantoin are synthesized by L. bicolor and that these compounds or their metabolites may be used for the benefit of aspen in exchange for the photosynthetically-derived sugars fructose and glucose. Conclusions: The analysis illustrates an approach to generate testable biological hypotheses to investigate the complex molecular interactions that drive ectomycorrhizal symbiosis. These models are consistent with experimental environmental data and provide insight into the molecular exchange processes for organisms in this complex ecosystem. The method used here for predicting metabolomic models of mycorrhizal systems from deep RNA sequencing data can be generalized and is broadly applicable to transcriptomic data derived from complex systems. [ABSTRACT FROM AUTHOR]

• Introduction: Heavy metal pollution is a strong driver of above- and belowground communities and triggers evolutionary adaptation in organisms. This review provides an overview of our knowledge on the effects of toxic concentrations of metals on ectomycorrhizal populations and communities. • Discussion: Selection and adaptations within particular ectomycorrhizal species that colonise host plants in harsh environments is discussed. Among these adaptations, we focus on the metal exclusion strategy that is discovered in metal-tolerant ecotypes of Suillus species that thrive on metalliferous soils. Metal efflux in metal-tolerant ecotypes prevents metal overloading of cytoplasm and vacuole. At the same time, this metal-specific efflux system in Suillus seems to reduce the transfer of large quantities of metals towards the plant-fungus interface, without hampering normal nutrient transfer to the host plant. • Conclusion: The evolutionary adaptation in Suillus species contributes to the survival of host trees on metalliferous soils and might be exploited in phytostabilisation strategies for heavy metal-contaminated soils. [ABSTRACT FROM AUTHOR]

Introduction: Knowledge of the factors that determine ectomycorrhizal fungal community structure is essential in many areas of practical significance including conservation, habitat restoration, prevention or amelioration of species invasions, and the prediction of responses to climate change. With respect to these goals, however, community ecology, in general, and ectomycorrhizal fungal community ecology, in particular, have been rather disappointing. Discussion: Indeed, some feel that community ecology is, at best, an esoteric discipline and, at worst, an inane one. But there is hope. As we apply what has been learned about other organisms concerning the relationships between functional traits and success (abundance), it may be possible to elucidate general principles that govern much of the structuring of ectomycorrhizal fungal communities. Conclusion: A hierarchical model of ectomycorrhizal fungal community structure is presented that involves abiotic filtering of immigrant propagules based on functional traits, followed by interspecific competition as ameliorated by disturbance and habitat partitioning, the outcomes of which are dependent on functional traits. [ABSTRACT FROM AUTHOR]

Background: Dryas octopetala is a widespread dwarf shrub in alpine and arctic regions that forms ectomycorrhizal (ECM) symbiotic relationships with fungi. In this study we investigated the fungal communities associated with roots of D. octopetala in alpine sites in Norway and in the High Arctic on Svalbard, where we aimed to reveal whether the fungal diversity and species composition varied across the Alpine and Arctic regions. The internal transcribed spacer (ITS) region of nuclear ribosomal DNA was used to identify the fungal communities from bulk root samples obtained from 24 plants. Results: A total of 137 operational taxonomic units (OTUs) were detected (using 97% similarity cut off during sequence clustering) and well-known ECM genera such as Cenococcum, Cortinarius, Hebeloma, Inocybe and Tomentella occurred frequently. There was no decrease in fungal diversity with increasing latitude. The overall spatial heterogeneity was high, but a weak geographical structuring of the composition of OTUs in the root systems was observed. Calculated species accumulation curves did not level off. Conclusions: This study indicates that the diversity of fungi associated with D. octopetala does not decrease in high latitude arctic regions, which contrasts observations made in a wide spectrum of other organism groups. A high degree of patchiness was observed across root systems, but the fungal communities were nevertheless weakly spatially structured. Non-asymptotical species accumulation curves and the occurrence of a high number of singletons indicated that only a small fraction of the fungal diversity was detected. [ABSTRACT FROM AUTHOR]

Background: Geographic selection mosaics, in which species exert different evolutionary impacts on each other in different environments, may drive diversification in coevolving species. We studied the potential for geographic selection mosaics in plant-mycorrhizal interactions by testing whether the interaction between bishop pine (Pinus muricata D. Don) and one of its common ectomycorrhizal fungi (Rhizopogon occidentalis Zeller and Dodge) varies in outcome, when different combinations of plant and fungal genotypes are tested under a range of different abiotic and biotic conditions. Results: We used a 2 x 2 x 2 x 2 factorial experiment to test the main and interactive effects of plant lineage (two maternal seed families), fungal lineage (two spore collections), soil type (lab mix or field soil), and non-mycorrhizal microbes (with or without) on the performance of plants and fungi. Ecological outcomes, as assessed by plant and fungal performance, varied widely across experimental environments, including interactions between plant or fungal lineages and soil environmental factors. Conclusion: These results show the potential for selection mosaics in plant-mycorrhizal interactions, and indicate that these interactions are likely to coevolve in different ways in different environments, even when initially the genotypes of the interacting species are the same across all environments. Hence, selection mosaics may be equally as effective as genetic differences among populations in driving divergent coevolution among populations of interacting species. [ABSTRACT FROM AUTHOR]

Background: Small GTPases of the Rho family function as tightly regulated molecular switches that govern important cellular functions in eukaryotes. Several families of regulatory proteins control their activation cycle and subcellular localization. Members of the guanine nucleotide dissociation inhibitor (GDI) family sequester Rho GTPases from the plasma membrane and keep them in an inactive form. Results: We report on the characterization the RhoGDI homolog of Tuber borchii Vittad., an ascomycetous ectomycorrhizal fungus. The Tbgdi gene is present in two copies in the T. borchii genome. The predicted amino acid sequence shows high similarity to other known RhoGDIs. Real time PCR analyses revealed an increased expression of Tbgdi during the phase preparative to the symbiosis instauration, in particular after stimulation with root exudates extracts, that correlates with expression of Tbcdc42. In a translocation assay TbRhoGDI was able to solubilize TbCdc42 from membranes. Surprisingly, TbRhoGDI appeared not to interact with S. cerevisiae Cdc42, precluding the use of yeast as a surrogate model for functional studies. To study the role of TbRhoGDI we performed complementation experiments using a RhoGDI null strain of Dictyostelium discoideum, a model organism where the roles of Rho signaling pathways are well established. For comparison, complementation with mammalian RhoGDI1 and LyGDI was also studied in the null strain. Although interacting with Rac1 isoforms, TbRhoGDI was not able to revert the defects of the D. discoideum RhoGDI null strain, but displayed an additional negative effect on the cAMP-stimulated actin polymerization response. Conclusion: T. borchii expresses a functional RhoGDI homolog that appears as an important modulator of cytoskeleton reorganization during polarized apical growth that antecedes symbiosis instauration. The specificity of TbRhoGDI actions was underscored by its inability to elicit a growth defect in S. cerevisiae or to compensate the loss of a D. discoideum RhoGDI. Knowledge of the cell signaling at the basis of cytoskeleton reorganization of ectomycorrhizal fungi is essential for improvements in the production of mycorrhized plant seedlings used in timberland extension programs and fruit body production. [ABSTRACT FROM AUTHOR]

Background: Symbiotic ectomycorrhizal associations of fungi with forest trees play important and economically significant roles in the nutrition, growth and health of boreal forest trees, as well as in nutrient cycling. The ecology and physiology of ectomycorrhizal associations with Pinus sp are very well documented but very little is known about the molecular mechanisms behind these mutualistic interactions with gymnosperms as compared to angiosperms. Results: Using a micro-array approach, the relative abundance of 2109 EST transcripts during interaction of Pinus sylvestris roots with the ectomycorrhizal fungus was profiled. The results reveal significant differential expression of a total of 236 ESTs, 96 transcripts differentially abundant after 1 day of physical contact with the fungus, 134 transcripts after 5 days and only 6 after 15 days at early stages of mantle formation on emerging lateral roots. A subset of cell wall modification and stress related genes was further assessed by quantitative reverse transcription PCR at late stages of mycorrhizal development coinciding with Hartig net formation. The results reveal down regulation of gene transcripts involved in general defence mechanism (e.g. antimicrobial peptide) as well as those involved in cell wall modification (e.g. glycine rich protein, xyloglucan endo transglycosylase). Conclusion: This study constitutes the first attempt to characterize the transcriptome of the plant partner in the Pinus sylvestris -- Laccaria bicolor model system. We identified 236 ESTs which are potentially important for molecular regulation of a functional symbiotic association in conifer host. The results highlight similarities with other studies based on angiosperm model systems, nevertheless some differences were found in the timing and spatial scale of gene regulation during ectomycorrhiza development in gymnosperms. The present study has identified a number of potentially important molecular events responsible for the initiation and regulation of biochemical, physiological and morphological changes during development of a fully functional symbiosis that are relevant for gymnosperm hosts. [ABSTRACT FROM AUTHOR]

The geographic mosaic theory is fast becoming a unifying framework for coevolutionary studies. A recent experimental study of interactions between pines and mycorrhizal fungi in BMC Biology is the first to rigorously test geographical selection mosaics, one of the cornerstones of the theory. [ABSTRACT FROM AUTHOR]

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Background: Pinus massoniana Lamb. is an important afforestation tree species with high economic, ecological and medicinal values. Aluminum (Al) toxicity driven by soil acidification causes dieback of P. massoniana plantations. Previous studies showed that ectomycorrhizal fungi alleviate Al stress damages in Pinus, but the underlying molecular mechanisms and key genes induced by ectomycorrhizal fungi inoculation under Al stress in Pinus have not been explored. Herein, we applied Al stress for 60 days to P. massoniana seedlings inoculated with Suillus luteus (SL) and those non-inoculated. Then, we compared their growth parameters and transcriptome in order to detect candidate genes induced by SL conferring Al tolerance in P. massoniana. Result: Our results showed that SL inoculation confers Al stress tolerance in P. massoniana through improved growth performance, strong antioxidant enzyme activities and reduced malondialdehyde accumulation as compared to non-inoculated seedlings. Transcriptome sequencing further supported these findings as very few genes (51 genes) were transcriptionally altered by Al in SL inoculated plants as compared to non-inoculated plants (2140 genes). We identified three core genes (cox1, cox3 and Nd1) that were strongly up-regulated by Al in the SL inoculated plants but were down-regulated in the non-inoculated plants. We also identified 42 genes specifically regulated by SL inoculated plants under Al stress, which are involved in a wide range of biological processes such as antioxidative response, transporters, hormone signaling and plant pathogen infection responses. Conclusions: Altogether, our data suggest that SL inoculation induces priming of key stress response pathways and triggers specific genes that efficiently alleviate Al stress effects in P. massoniana. The candidate genes resources generated in this study are of utmost importance for functional characterization and molecular studies aiming at improving Al tolerance in plants. [ABSTRACT FROM AUTHOR]

Background: Ectomycorrhizal fungi (ECM) play a central role in nutrient cycling in boreal and temperate forests, but their role in the soil food web remains little understood. One of the groups assumed to live as specialised mycorrhizal feeders are Protura, but experimental and field evidence is lacking. We used a combination of three methods to test if Protura are specialized mycorrhizal feeders and compared their trophic niche with other soil invertebrates. Using pulse labelling of young beech and ash seedlings we analysed the incorporation of 13C and 15N into Acerentomon gallicum. In addition, individuals of Protura from temperate forests were collected for the analysis of neutral lipid fatty acids and natural variations in stable isotope ratios. Results: Pulse labelling showed rapid incorporation of root-derived 13C, but no incorporation of root-derived 15N into A. gallicum. The transfer of 13C from lateral roots to ectomycorrhizal root tips was high, while it was low for 15N. Neutral lipid fatty acid (NLFA) analysis showed high amounts of bacterial marker (16:1ω7) and plant marker (16:0 and 18:1ω9) fatty acids but not of the fungal membrane lipid 18:2ω6,9 in A. gallicum. Natural variations in stable isotope ratios in Protura from a number of temperate forests were distinct from those of the great majority of other soil invertebrates, but remarkably similar to those of sporocarps of ECM fungi. Conclusions: Using three in situ methods, stable isotope labelling, neutral lipid fatty acid analysis and natural variations of stable isotope ratios, we showed that Protura predominantly feed on mycorrhizal hyphae via sucking up hyphal cytoplasm. Predominant feeding on ectomycorrhizal mycelia by Protura is an exception; the limited consumption of ECM by other soil invertebrates may contribute to carbon sequestration in temperate and boreal forests. [ABSTRACT FROM AUTHOR]

Background: Below-ground linkage between plant and fungal communities is one of the major drivers of terrestrial ecosystem dynamics. However, we still have limited knowledge of how such plant–fungus associations vary in their community-scale properties depending on fungal functional groups and geographic locations. Methods: By compiling a high-throughput sequencing dataset of root-associated fungi in eight forests along the Japanese Archipelago, we performed a comparative analysis of arbuscular mycorrhizal, ectomycorrhizal, and saprotrophic/endophytic associations across a latitudinal gradient from cool-temperate to subtropical regions. Results: In most of the plant–fungus networks analyzed, host–symbiont associations were significantly specialized but lacked "nested" architecture, which has been commonly reported in plant–pollinator and plant–seed disperser networks. In particular, the entire networks involving all functional groups of plants and fungi and partial networks consisting of ectomycorrhizal plant and fungal species/taxa displayed "anti-nested" architecture (i.e., negative nestedness scores) in many of the forests examined. Our data also suggested that geographic factors affected the organization of plant–fungus network structure. For example, the southernmost subtropical site analyzed in this study displayed lower network-level specificity of host–symbiont associations and higher (but still low) nestedness than northern localities. Conclusions: Our comparative analyses suggest that arbuscular mycorrhizal, ectomycorrhizal, and saprotrophic/endophytic plant–fungus associations often lack nested network architecture, while those associations can vary, to some extent, in their community-scale properties along a latitudinal gradient. Overall, this study provides a basis for future studies that will examine how different types of plant–fungus associations collectively structure terrestrial ecosystems. [ABSTRACT FROM AUTHOR]

Key message: Selection of the best salt-tolerant combination ofCasuarinasp. and arbuscular mycorrhizal fungi (AMF) is one of the key criteria for successful setup of saline land rehabilitation program.Context: Land salinization is a serious problem worldwide that mainly leads to soil degradation and reduces crop productivity. These degraded areas could be rehabilitated by planting salt-tolerant species like Casuarina glauca Sieb. and Casuarina equisetifolia L. These are pioneer plants, able to form symbiotic associations with arbuscular mycorrhizal fungi (AMF), ectomycorrhizal fungi (EMF), and nitrogen-fixing bacteria.Aims: The aim of this study was to select the highest salt-tolerant combination of Casuarina/AMF that can be used for the rehabilitation of lands degraded by salinity.Methods: C. equisetifolia and C. glauca were grown in sandy sterile soil in the greenhouse and inoculated separately with Rhizophagus fasciculatus (Thaxt.) C. Walker & A. Schüßler, Rhizophagus aggregatus (N.C. Schenck & G.S. Sm.) C. Walker, and Rhizophagus intraradices (N.C. Schenck & G.S. Sm.) C. Walker & A. Schüßler. After confirming the establishment of a symbiosis, the plants were watered with gradually increasing concentrations of saline solution. After harvest, size and biomass of the seedlings, root colonization by AMF, and AMF metabolic activities were evaluated.Results: A larger growth was obtained in the two species when the individuals were inoculated with R. fasciculatus. Root colonization rates did not differ among fungal species, but fungal metabolic activities were higher in mycorrhizal roots of C. glauca plants inoculated with R. fasciculatus.Conclusion: Among the three mycorrhizal fungi, R. fasciculatus was more efficient in association with Casuarinaceae species under salt stress. Our results suggest that selection of appropriate fungal strains is crucial to improve plant performance in saline soils. [ABSTRACT FROM AUTHOR]

The article reports on the findings of a study about the benefits of ectomycorrhizal fungi (EMF) to trees and other plants, conducted by O. Franklin and colleagues, which suggest that feedback loops with EMF may maintain the nitrogen limitation situation for plant growth in boreal forests.