Influence of tree mycorrhizal type, tree species identity, and diversity on forest root-associated mycobiomes.

Understanding the complex interactions between trees and fungi is crucial for forest ecosystem management, yet the influence of tree mycorrhizal types, species identity, and diversity on tree-tree interactions and their root-associated fungal communities remains poorly understood. Our study addresses this gap by investigating root-associated fungal communities of different arbuscular mycorrhizal (AM) and ectomycorrhizal (EcM) tree species pairs (TSPs) in a subtropical tree diversity experiment, spanning monospecific, two-species, and multi-species mixtures, utilizing Illumina sequencing of the ITS2 region. The study reveals that tree mycorrhizal type significantly impacts the alpha diversity of root-associated fungi in monospecific stands. Meanwhile, tree species identity's influence is modulated by overall tree diversity. Tree-related variables and spatial distance emerged as major drivers of variations in fungal community composition. Notably, in multi-species mixtures, compositional differences between root fungal communities of AM and EcM trees diminish, indicating a convergence of fungal communities irrespective of mycorrhizal type. Interestingly, dual mycorrhizal fungal communities were observed in these multi-species mixtures. This research underscores the pivotal role of mycorrhizal partnerships and the interplay of biotic and abiotic factors in shaping root fungal communities, particularly in varied tree diversity settings, and its implications for effective forest management and biodiversity conservation.

Environmental modulation of plant mycorrhizal traits in the global flora.

Mycorrhizal symbioses are known to strongly influence plant performance, structure plant communities and shape ecosystem dynamics. Plant mycorrhizal traits, such as those characterising mycorrhizal type (arbuscular (AM), ecto-, ericoid or orchid mycorrhiza) and status (obligately (OM), facultatively (FM) or non-mycorrhizal) offer valuable insight into plant belowground functionality. Here, we compile available plant mycorrhizal trait information and global occurrence data ( ∼ 100 million records) for 11,770 vascular plant species. Using a plant phylogenetic mega-tree and high-resolution climatic and edaphic data layers, we assess phylogenetic and environmental correlates of plant mycorrhizal traits. We find that plant mycorrhizal type is more phylogenetically conserved than plant mycorrhizal status, while environmental variables (both climatic and edaphic; notably soil texture) explain more variation in mycorrhizal status, especially FM. The previously underestimated role of environmental conditions has far-reaching implications for our understanding of ecosystem functioning under changing climatic and soil conditions.

Herbarium specimens reveal that mycorrhizal type does not mediate declining temperate tree nitrogen status over a century of environmental change.

Rising atmospheric carbon dioxide concentrations (CO2 ) and atmospheric nitrogen (N) deposition have contrasting effects on ectomycorrhizal (EM) and arbuscular mycorrhizal (AM) symbioses, potentially mediating forest responses to environmental change. In this study, we evaluated the cumulative effects of historical environmental change on N concentrations and δ15 N values in AM plants, EM plants, EM fungi, and saprotrophic fungi using herbarium specimens collected in Minnesota, USA from 1871 to 2016. To better understand mycorrhizal mediation of foliar δ15 N, we also analyzed a subset of previously published foliar δ15 N values from across the United States to parse the effects of N deposition and CO2 rise. Over the last century in Minnesota, N concentrations declined among all groups except saprotrophic fungi. δ15 N also declined among all groups of plants and fungi; however, foliar δ15 N declined less in EM plants than in AM plants. In the analysis of previously published foliar δ15 N values, this slope difference between EM and AM plants was better explained by nitrogen deposition than by CO2 rise. Mycorrhizal type did not explain trajectories of plant N concentrations. Instead, plants and EM fungi exhibited similar declines in N concentrations, consistent with declining forest N status despite moderate levels of N deposition.

Peat loss collocates with a threshold in plant-mycorrhizal associations in drained peatlands encroached by trees.

Drainage-induced encroachment by trees may have major effects on the carbon balance of northern peatlands, and responses of microbial communities are likely to play a central mechanistic role. We profiled the soil fungal community and estimated its genetic potential for the decay of lignin and phenolics (class II peroxidase potential) along peatland drainage gradients stretching from interior locations (undrained, open) to ditched locations (drained, forested). Mycorrhizal fungi dominated the community across the gradients. When moving towards ditches, the dominant type of mycorrhizal association abruptly shifted from ericoid mycorrhiza to ectomycorrhiza at c. 120 m from the ditches. This distance corresponded with increased peat loss, from which more than half may be attributed to oxidation. The ectomycorrhizal genus Cortinarius dominated at the drained end of the gradients and its relatively higher genetic potential to produce class II peroxidases (together with Mycena) was positively associated with peat humification and negatively with carbon-to-nitrogen ratio. Our study is consistent with a plant-soil feedback mechanism, driven by a shift in the mycorrhizal type of vegetation, that potentially mediates changes in aerobic decomposition during postdrainage succession. Such feedback may have long-term legacy effects upon postdrainage restoration efforts and implication for tree encroachment onto carbon-rich soils globally.

Mycorrhizal type of woody plants influences understory species richness in British broadleaved woodlands.

Mature temperate woodlands are commonly dominated by ectomycorrhizal trees, whereas understory plants predominantly form arbuscular mycorrhizal associations. Due to differences in plant-fungus compatibility between canopy and ground layer vegetation the 'mycorrhizal mediation hypothesis' predicts that herbaceous plant establishment may be limited by a lack of suitable mycorrhizal fungal inoculum. We examined plant species data for 103 woodlands across Great Britain recorded in 1971 and in 2000 to test whether herbaceous plant species richness was related to the proportion of arbuscular mycorrhizal woody plants. We compared the effect of mycorrhizal type with other important drivers of woodland plant species richness. We found a positive effect of the relative abundance of arbuscular mycorrhizal woody plants on herbaceous plant species richness. The size of the observed effect was smaller than that of pH. Moreover, the effect persisted over time, despite many woodlands undergoing marked successional change and increased understorey shading. This work supports the mycorrhizal mediation hypothesis in British woodlands and suggests that increased abundance of arbuscular mycorrhizal woody plants is associated with greater understory plant species richness.

A tipping point in carbon storage when forest expands into tundra is related to mycorrhizal recycling of nitrogen.

Tundra ecosystems are global belowground sinks for atmospheric CO2 . Ongoing warming-induced encroachment by shrubs and trees risks turning this sink into a CO2 source, resulting in a positive feedback on climate warming. To advance mechanistic understanding of how shifts in mycorrhizal types affect long-term carbon (C) and nitrogen (N) stocks, we studied small-scale soil depth profiles of fungal communities and C-N dynamics across a subarctic-alpine forest-heath vegetation gradient. Belowground organic stocks decreased abruptly at the transition from heath to forest, linked to the presence of certain tree-associated ectomycorrhizal fungi that contribute to decomposition when mining N from organic matter. In contrast, ericoid mycorrhizal plants and fungi were associated with organic matter accumulation and slow decomposition. If climatic controls on arctic-alpine forest lines are relaxed, increased decomposition will likely outbalance increased plant productivity, decreasing the overall C sink capacity of displaced tundra.

Aboveground herbivores drive stronger plant species-specific feedback than belowground fungi to regulate tree community assembly.

Ectomycorrhizal (EcM) tree species often become more dominant than arbuscular mycorrhizal (AM) tree species in temperate forests, but they generally coexist. Theory predicts that ecological feedback mediated by aboveground herbivory and/or belowground microbes could explain these dominance/coexistence patterns. An experimental test of how aboveground/belowground organisms associated with AM/EcM trees mediate ecological feedbacks has been lacking at the community-level. By establishing AM and EcM tree sapling assemblages in mesocosms and then introducing seedlings of each type in a reciprocal planting experiment, we compared seedling performance under varying sapling species (conspecifics, heterospecifics within the same and different mycorrhizal types), using traits that reflect either aboveground herbivory-mediated feedback or belowground fungal-mediated feedback or both. When examining seedling traits that reflect aboveground herbivory-mediated feedbacks (i.e., foliar damage), AM plants tended to experience less foliar damage and EcM plants more damage under conspecific versus heterospecific saplings within the same mycorrhizal types, and aboveground herbivory-mediated feedback was species-specific rather than mycorrhizal type-specific. Conversely, when examining traits that reflect belowground fungal-mediated feedbacks, both AM and EcM plant species often exhibited mycorrhizal type-specific feedbacks (e.g., greater aboveground biomass under the same versus different mycorrhizal-type saplings) rather than species-specific feedbacks. Furthermore, tree species affected by herbivory-mediated feedback were less affected by belowground feedback, indicating that the relative importance of the feedbacks varied among plant species. Analysis of plant-associated organisms verified that the feedback outcomes corresponded with species accumulation of belowground fungi (but not of aboveground herbivores). Thus, aboveground herbivores drive stronger plant species-specific feedback than belowground fungi to regulate temperate tree diversity.

Tree growth response to soil nutrients and neighborhood crowding varies between mycorrhizal types in an old-growth temperate forest.

Forest dynamics are shaped by both abiotic and biotic factors. Trees associating with different types of mycorrhizal fungi differ in nutrient use and dominate in contrasting environments, but it remains unclear whether they exhibit differential growth responses to local abiotic and biotic gradients where they co-occur. We used 9-year tree census data in a 25-ha old-growth temperate forest in Northeast China to examine differences in tree growth response to soil nutrients and neighborhood crowding between tree species associating with arbuscular mycorrhizal (AM), ectomycorrhizal (EM), and dual-mycorrhizal (AEM) fungi. In addition, we tested the role of individual-level vs species-level leaf traits in capturing differences in tree growth response to soil nutrients and neighborhood crowding across mycorrhizal types. Across 25 species, soil nutrients decreased AM tree growth, while neighborhood crowding reduced both AM and EM tree growth, and neither soil nor neighbors impacted AEM tree growth. Across mycorrhizal types, individual-level traits were stronger predictors of tree growth than species-level traits. However, most traits poorly mediated tree growth response to soil nutrients and neighborhood crowding. Our findings indicate that mycorrhizal types strongly shape differences in tree growth response to local soil and crowding gradients, and suggest that including plant-mycorrhizae associations in future work offers great potential to improve our understanding of forest dynamics.

Plant nutrient-acquisition strategies drive topsoil microbiome structure and function.

Plant nutrient-acquisition strategies drive soil processes and vegetation performance, but their effect on the soil microbiome remains poorly understood. This knowledge is important to predict the shifts in microbial diversity and functions due to increasing changes in vegetation traits under global change. Here we documented the topsoil microbiomes of 145 boreal and temperate terrestrial sites in the Baltic region that broadly differed in vegetation type and nutritional traits, such as mycorrhizal types and symbiotic nitrogen-fixation. We found that sites dominated by arbuscular mycorrhizal (AM) vegetation harbor relatively more AM fungi, bacteria, fungal saprotrophs, and pathogens in the topsoil compared with sites dominated by ectomycorrhizal (EM) plants. These differences in microbiome composition reflect the rapid nutrient cycling and negative plant-soil feedback in AM soils. Lower fungal diversity and bacteria : fungi ratios in EM-dominated habitats are driven by monodominance of woody vegetation as well as soil acidification by EM fungi, which are associated with greater diversity and relative abundance of carbohydrate-active enzymes. Our study suggests that shifts in vegetation related to global change and land use may strongly alter the topsoil microbiome structure and function.

FungalRoot: global online database of plant mycorrhizal associations.

Testing of ecological, biogeographical and phylogenetic hypotheses of mycorrhizal traits requires a comprehensive reference dataset about plant mycorrhizal associations. Here we present a database, FungalRoot, which summarizes publicly available data about vascular plant mycorrhizal type and intensity of root colonization by mycorrhizal fungi, accompanied with rich metadata. We compiled and digitized data about plant mycorrhizal colonization in nine widespread languages. The present version of the FungalRoot database contains 36 303 species-by-site observations for 14 870 plant species, tripling the previously available compiled information about plant mycorrhizal associations. Based on these data, we provide a recommended list of genus-level plant mycorrhizal associations, based on the majority of data for species and careful analysis of conflicting data. The majority of ectomycorrhizal and ericoid mycorrhizal plants are trees (92%) and shrubs (85%), respectively. The majority of arbuscular and nonmycorrhizal plant species are herbaceous (50% and 70%, respectively). Our publicly available database is a powerful resource for mycorrhizal scientists and ecologists. It features possibilities for dynamic updating and addition of data about plant mycorrhizal associations. The new database will promote research on plant and fungal biogeography and evolution, and on links between above- and belowground biodiversity and ecosystem functioning.

Conspecific distance-dependent seedling performance, and replacement of conspecific seedlings by heterospecifics in five hardwood, temperate forest species.

The mechanisms driving species diversity in the context of Janzen-Connell model are best understood by evaluating not only conspecific distance-dependent (CDD) seedling performance, but also replacement of conspecific seedlings by heterospecific seedlings beneath adult trees. We evaluated CDD and replacement as a log response ratio of seedling performance (height, age) directly beneath and at a distance from adult plants in a temperate forest, and examined the log response ratio of that between conspecifics and heterospecifics beneath adults for five hardwood species with different ecological traits (e.g., seed size, mycorrhizal type, relative abundance). CDD was greater in three small-seeded species with arbuscular mycorrhizae (AM) associations than it was in two large-seeded species with ectomycorrhizae (EM) associations. Replacement was also higher for small-seeded AM species compared to large-seeded EM species, resulting in a strong, positive relationship between CDD and replacement. The traits suggest that small-seeded AM seedlings are more likely to be replaced by heterologous seedlings beneath the adults than large-seeded EM seedlings, probably due to that the small-seeded AM species are more susceptible to attack by plant natural enemies (e.g., soil pathogens, leaf diseases). As a result, small-seeded AM species had lower relative abundances compared to large-seeded EM species. This study suggests that either seed size or associations with microorganisms play an important role in driving forest diversity by regulating replacement and CDD, although relative importance of the two traits (i.e., seed size, mycorrhizal type) remains unclear, because of the autocorrelation between the two traits for the five species studied.

Changes in mycorrhizal status and type in plant communities along altitudinal and ecological gradients-a case study from the Northern Urals (Russia).

The Ural Mountains (the Urals) are a mountain range on the border between the continents of Europe and Asia. The Urals extend about 2500 km from north to south and run from the coast of the Arctic Ocean to the steppe of northwestern Kazakhstan. In terms of mycorrhizal traits, the vegetation of the Urals has not been studied absolutely compared with the other mountain systems of Europe. Detailed research of vegetation at the Telpos-iz Ridge (Northern Urals, Russia) allowed us to analyze changes in mycorrhizal status (obligatory mycorrhizal, OM; facultative mycorrhizal, FM; non-mycorrhizal, NM) and type (arbuscular, AM; ectomycorrhiza, ECM; ericoid, ERM; ORM, orchid mycorrhiza; NM) both in the main vegetation types and the individual communities along the elevation and ecological gradients based on 165 releves. This is the first attempt to describe the mycorrhizal status of plant communities across different elevations and ecological conditions outside Europe and North America. OM species were most diverse in all vegetation types. Maximal share of NM species was found in the mires and may be explained by over logged and unfertile soils of these habitats. Arbuscular mycorrhizas dominated across all vegetation types except for mires, where specific ecological conditions result in the prevalence of ECM and NM species. We analyzed the mycorrhizal status and type of plant communities along the main ecological gradients and found a decisive role of elevation and soil nitrogen content. At the same time, it remains unclear which factor determines the distribution of ECM and AM communities which are most represented in the vegetation of the study area.

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.

Similar below-ground carbon cycling dynamics but contrasting modes of nitrogen cycling between arbuscular mycorrhizal and ectomycorrhizal forests.

Compared with ectomycorrhizal (ECM) forests, arbuscular mycorrhizal (AM) forests are hypothesized to have higher carbon (C) cycling rates and a more open nitrogen (N) cycle. To test this hypothesis, we synthesized 645 observations, including 22 variables related to below-ground C and N dynamics from 100 sites, where AM and ECM forests co-occurred at the same site. Leaf litter quality was lower in ECM than in AM trees, leading to greater forest floor C stocks in ECM forests. By contrast, AM forests had significantly higher mineral soil C concentrations, and this result was strongly mediated by plant traits and climate. No significant differences were found between AM and ECM forests in C fluxes and labile C concentrations. Furthermore, inorganic N concentrations, net N mineralization and nitrification rates were all higher in AM than in ECM forests, indicating 'mineral' N economy in AM but 'organic' N economy in ECM trees. AM and ECM forests show systematic differences in mineral vs organic N cycling, and thus mycorrhizal type may be useful in predicting how different tree species respond to multiple environmental change factors. By contrast, mycorrhizal type alone cannot reliably predict below-ground C dynamics without considering plant traits and climate.

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Species richness plays an important role in ecosystem stability and health. Mycorrhizal type is an important factor affecting ecological processes. How mycorrhizal types affect understory herb species richness and their responses to environmental changes remain largely unknown. We investigated the effects of mycorrhizal types on species richness and their responses to environmental change in understory herbaceous communities based on data of three mycorrhizal types of dominated trees (including 1604 arbuscular mycorrhiza (AM) trees, 4654 ectomycorrhiza (ECM) trees, and 5568 AM+ECM trees) and environmental factors in America. The results showed significant differences in species richness of herbaceous plant communities among different mycorrhizal types. Forests with higher dominance of AM plants tended to have higher herbaceous plant richness, supporting the mycorrhizal mediation hypothesis. The impacts of environmental factors (latitude, temperature, precipitation, nitrogen deposition, and soil characteristics) on species richness of herbaceous plant communities depended on mycorrhizal type of forests. The species richness of understory herbs in AM, ECM, and AM+ECM forests was mostly affected by nitrogen deposition, temperature, and soil pH, with the relative importance of 42.3%, 41.1% and 48.7%, respectively. Mycorrhizal types of dominant trees played a vital role in regulating the species richness of understory herbs and influenced their responses to environmental changes.

Factors associated with seedling establishment on logs of different fungal decay types-A seed-sowing experiment.

Wood decay fungi alter the abiotic and biotic properties of deadwood, which are important as nurse logs for seedling regeneration. However, the relationship between fungal decay type and seedling performance has not been evaluated experimentally. In this study, we examined the germination, growth, and survival of six arbuscular mycorrhizal (AM) and six ectomycorrhizal (ECM) tree species on three substrates (pine logs with brown and white rot and soil) by conducting seed-sowing experiments in a mixed forest dominated by Pinus densiflora and Quercus serrata. Analysis using ribosomal DNA internal transcribed spacer 1 (rDNA ITS1) sequencing revealed that the fungal community was significantly different across three substrates. The richness of operational taxonomic units (OTUs) of AM and ECM fungi was the largest on brown rot logs and soil, respectively. The substrate significantly affected the seedling performance when comparing wood decay types, but these were not consistent across the mycorrhizal status of the seedlings. Nevertheless, seedlings of some AM trees showed better growth and enhanced mycorrhizal colonization on brown rot logs than on white rot logs. The wood decay type influenced fungal communities in the logs and the performance of some seedling species partly by different mycorrhizal colonization rates. However, the effect was seedling species dependent and showed no apparent difference between AM and ECM trees.

Functional Potential of Soil Microbial Communities and Their Subcommunities Varies with Tree Mycorrhizal Type and Tree Diversity.

Soil microbial communities play crucial roles in the earth's biogeochemical cycles. Yet, their genomic potential for nutrient cycling in association with tree mycorrhizal type and tree-tree interactions remained unclear, especially in diverse tree communities. Here, we studied the genomic potential of soil fungi and bacteria with arbuscular (AM) and ectomycorrhizal (EcM) conspecific tree species pairs (TSPs) at three tree diversity levels in a subtropical tree diversity experiment (BEF-China). The soil fungi and bacteria of the TSPs' interaction zone were characterized by amplicon sequencing, and their subcommunities were determined using a microbial interkingdom co-occurrence network approach. Their potential genomic functions were predicted with regard to the three major nutrients carbon (C), nitrogen (N), and phosphorus (P) and their combinations. We found the microbial subcommunities that were significantly responding to different soil characteristics. The tree mycorrhizal type significantly influenced the functional composition of these co-occurring subcommunities in monospecific stands and two-tree-species mixtures but not in mixtures with more than three tree species (here multi-tree-species mixtures). Differentiation of subcommunities was driven by differentially abundant taxa producing different sets of nutrient cycling enzymes across the tree diversity levels, predominantly enzymes of the P (n = 11 and 16) cycles, followed by the N (n = 9) and C (n = 9) cycles, in monospecific stands and two-tree-species mixtures, respectively. Fungi of the Agaricomycetes, Sordariomycetes, Eurotiomycetes, and Leotiomycetes and bacteria of the Verrucomicrobia, Acidobacteria, Alphaproteobacteria, and Actinobacteria were the major differential contributors (48% to 62%) to the nutrient cycling functional abundances of soil microbial communities across tree diversity levels. Our study demonstrated the versatility and significance of microbial subcommunities in different soil nutrient cycling processes of forest ecosystems. IMPORTANCE Loss of multifunctional microbial communities can negatively affect ecosystem services, especially forest soil nutrient cycling. Therefore, exploration of the genomic potential of soil microbial communities, particularly their constituting subcommunities and taxa for nutrient cycling, is vital to get an in-depth mechanistic understanding for better management of forest soil ecosystems. This study revealed soil microbes with rich nutrient cycling potential, organized in subcommunities that are functionally resilient and abundant. Such microbial communities mainly found in multi-tree-species mixtures associated with different mycorrhizal partners can foster soil microbiome stability. A stable and functionally rich soil microbiome is involved in the cycling of nutrients, such as carbon, nitrogen, and phosphorus, and their combinations could have positive effects on ecosystem functioning, including increased forest productivity. The new findings could be highly relevant for afforestation and reforestation regimes, notably in the face of growing deforestation and global warming scenarios.

Species of fast bulk-soil nutrient cycling have lower rhizosphere effects: A nutrient spectrum of rhizosphere effects.

Tree roots not only acquire readily-usable soil nutrients but also affect microbial decomposition and manipulate nutrient availability in their surrounding soils, that is, rhizosphere effects (REs). Thus, REs challenge the basic understanding of how plants adapt to the environment and co-exist with other species. Yet, how REs vary among species in response to species-specific bulk soil nutrient cycling is not well-known. Here, we studied how plant-controlled microbial decomposition activities in rhizosphere soils respond to those in their corresponding bulk soils and whether these relations depend on species-specific nutrient cycling in the bulk soils. We targeted 55 woody species of different clades and mycorrhizal types in three contrasting biomes, namely a temperate forest, a subtropical forest, and a tropical forest. We found that microbial decomposition activities in rhizosphere soils responded linearly to those in their corresponding bulk soils at the species level. Thereafter, we found that REs (parameters in rhizosphere soils minus those in corresponding bulk soils) of microbial decomposition activities had negative linear correlations with microbial decomposition activities in corresponding bulk soils. A multiple factor analysis revealed that soil organic carbon, total nitrogen, and soil water content favored bulk soil decomposition activities in all three biomes, showing that the magnitude of REs varied along a fast-slow nutrient cycling spectrum in bulk soils. The species of fast nutrient cycling in their bulk soils tended to have smaller or even negative REs. Therefore, woody plants commonly utilize both positive and negative REs as a nutrient-acquisition strategy. Based on the trade-offs between REs and other nutrient-acquisition strategies, we proposed a push and pull conceptual model which can bring plant nutrient-acquisition cost and plant carbon economics spectrum together in the future. This model will facilitate not only the carbon and nutrient cycling but also the mechanisms of species co-existence in forest ecosystems.