Forest Age and Plant Species Composition Determine the Soil Fungal Community Composition in a Chinese Subtropical Forest.

Seasonal variations of soil fungal diversity and communities in subalpine coniferous and broadleaved forests

To investigate the response mechanisms of soil bacterial and fungal communities to the changes of preci-pitation in a desert steppe of Ningxia, we conducted a three-year precipitation control experiment following completely randomized design. There were five treatments, natural precipitation (T0), 50% less in precipitation (T1), 25% less in precipitation (T2), 25% more in precipitation (T3) and 50% more in precipitation (T4). By using Illumina high-throughput sequencing and bioinformatics analysis, we investigated the effects of increased and decreased precipitation on soil bacterial and fungal communities, and examined the correlations between soil physicochemical properties, plant communities and soil bacterial and fungal communities. The result showed that the richness of soil bacteria and fungi was highest in the T4 treatment. In addition, the relative abundance of Chloroflexi, the predominant phyla of soil bacteria was more sensitive to precipitation change. However, the relative abundance of only Ascomycota, a rare fungal taxon, responded to precipitation changes. Results of redundancy analysis showed that the first two axes accounted for 92.8% and 87.4% of the total variance for soil bacterial and fungal community composition, respectively. Precipitation and soil pH were the most important environmental factors driving changes in soil bacterial diversity and community composition. On the one hand, precipitation had a direct positive effect on bacterial diversity and community composition. On the other hand, precipitation changed pH by affecting soil moisture, which in turn had a significant indirect effect on bacterial diversity and community composition. Plant community biomass, plant species richness, and soil pH were the most influential environmental factors affecting fungal diversity and community composition. Precipitation had no direct effect on soil fungal community, but had a significant indirect effect by changing plant community richness and soil pH. The response mechanisms of bacterial and fungal communities in soil differed significantly under different precipitation regimes in the desert grasslands of Ningxia.

Whether and how seasonality of environmental variables impacts the spatial variability of soil fungal communities remain poorly understood. We assessed soil fungal diversity and community composition of five Chinese zonal forests along a latitudinal gradient spanning 23°N to 42°N in three seasons to address these questions. We found that soil fungal diversity increased linearly or parabolically with latitude. The seasonal variations in fungal diversity were more distinguishable in three temperate deciduous forests than in two subtropical evergreen forests. Soil fungal diversity was mainly correlated with edaphic factors such as pH and nutrient contents. Both latitude and its interactions with season also imposed significant impacts on soil fungal community composition (FCC), but the effects of latitude were stronger than those of season. Vegetational properties such as plant diversity and forest age were the dominant factors affecting FCC in the subtropical evergreen forests while edaphic properties were the dominant ones in the temperate deciduous forests. Our results indicate that latitudinal variation patterns of soil fungal diversity and FCC may differ among seasons. The stronger effect of latitude relative to that of season suggests a more important influence by the spatial than temporal heterogeneity in shaping soil fungal communities across zonal forests.

Discriminating the effects of agricultural land management practices on soil fungal communities

Soil fungi play an integral and essential role in maintaining soil ecosystem functions. The understanding of altitude variations and their drivers of soil fungal community composition and diversity remains relatively unclear. Mountains provide an open, natural platform for studying how the soil fungal community responds to climatic variability at a short altitude distance. Using the Illumina MiSeq high-throughput sequencing technique, we examined soil fungal community composition and diversity among seven vegetation types (dry valley shrub, valley-mountain ecotone broadleaved mixed forest, subalpine broadleaved mixed forest, subalpine coniferous-broadleaved mixed forest, subalpine coniferous forest, alpine shrub meadow, alpine meadow) along a 2582 m altitude gradient in the alpine–gorge region on the eastern Qinghai–Tibetan Plateau. Ascomycota (47.72%), Basidiomycota (36.58%), and Mortierellomycota (12.14%) were the top three soil fungal dominant phyla in all samples. Soil fungal community composition differed significantly among the seven vegetation types along altitude gradients. The α-diversity of soil total fungi and symbiotic fungi had a distinct hollow pattern, while saprophytic fungi and pathogenic fungi showed no obvious pattern along altitude gradients. The β-diversity of soil total fungi, symbiotic fungi, saprophytic fungi, and pathogenic fungi was derived mainly from species turnover processes and exhibited a significant altitude distance-decay pattern. Soil properties explained 31.27−34.91% of variation in soil fungal (total and trophic modes) community composition along altitude gradients, and the effects of soil nutrients on fungal community composition varied by trophic modes. Soil pH was the main factor affecting α-diversity of soil fungi along altitude gradients. The β-diversity and turnover components of soil total fungi and saprophytic fungi were affected by soil properties and geographic distance, while those of symbiotic fungi and pathogenic fungi were affected only by soil properties. This study deepens our knowledge regarding altitude variations and their drivers of soil fungal community composition and diversity, and confirms that the effects of soil properties on soil fungal community composition and diversity vary by trophic modes along altitude gradients in the alpine–gorge region.

Soil fungal communities play a critical role in ecosystem carbon (C) and nitrogen (N) cycling. Although the effect of plant invasions on ecosystem C and N cycling is well established, its impact on soil fungal communities is not fully understood. The objective of this study was therefore to understand the variations in soil fungal communities as affected by plant invasion, and the mechanisms that drive these changes. We examined the impacts of invasive Spartina alternifolia Loisel. (SA) on soil fungal abundance, diversity, community composition, trophic modes and functional groups in comparison with bare flat (BF) and native Suaeda salsa (Linn.) Pall. (SS), Scirpus mariqueter Tang et Wang (SM), and Phragmites australis (Cav.) Trin. ex Steud. (PA) communities in coastal salt marshes of eastern China, based on analyses of the quantitative polymerase chain reaction (qPCR) and Illumina MiSeq DNA sequences of fungal internal transcribed spacer (ITS) region. SA invasion increased the soil fungal abundance and diversity compared to BF, SS, SM, and PA soils. The increased soil fungal abundance and diversity were highly related to soil organic carbon (C) and nitrogen (N), water-soluble organic carbon (WSOC), litter C:N ratio, and root C:N ratio. Soil fungal community composition was shifted following SA invasion. Specifically, SA invasion significantly enhanced the relative abundance of Basidiomycota, and reduced the relative abundance of Ascomycota compared with BF, SS, SM, and PA soils. Additionally, SA invasion changed soil fungal trophic modes and functional groups. The relative abundance of saprotrophic fungi significantly increased, while the relative abundances of symbiotic and pathotrophic fungi decreased following SA invasion. Our data revealed that SA invasion altered soil fungal abundance, diversity, community composition, trophic modes and functional groups, which were primarily driven by the quality and quantity of plant residues, soil nutrition substrates, as well as soil physicochemical properties. The changes in soil fungal communities, especially their trophic modes and functional groups following SA invasion would greatly affect soil C and N decomposition and accumulation with potential feedback on climate change.

Different responses of soil bacterial and fungal communities to nitrogen deposition in a subtropical forest

ABSTRACTDeciphering the relationships between microbes and their host plants is critical for a better understanding of microbial diversity maintenance and community stability. Here, we investigated fungal diversity and community assembly in the phyllosphere and rhizosphere of 13 tree species in a subtropical common-garden experiment. The results showed that fungal community structures significantly differed across compartments (leaf, root, and soil) and different tree species. Higher α-diversity was observed in the phyllosphere than in the roots and rhizospheric soil. Fungal community composition (β-diversity) was significantly affected by both compartment and species identity. The fungal community compositions were significantly correlated with soil pH in the roots and the soils as well as with soil nitrate and leaf total phosphorus in the leaves. We found that fungal community assemblies were mainly driven by deterministic processes, regardless of compartments. Moreover, host preference analyses indicated that stronger plant/fungus preferences occurred in leaves than in roots and soils. Our results highlight the differences in tree mycobiome between aboveground and belowground compartments and have important implications for the promotion of biodiversity conservation and management sustainability for the subtropical forest.IMPORTANCE Subtropical mountain forests are widely distributed in Southern China and are characterized by high biodiversity. The interactions between plants and fungi play pivotal roles in biodiversity maintenance and community stability. Nevertheless, knowledge of fungal diversity and of the community assembly patterns of woody plants is scarce. Here, we investigated fungal diversity and community assembly in the phyllosphere and rhizosphere of 13 tree species in a common-garden experiment. We found that both compartment and plant identity influenced fungal diversity, community, and guild compositions, while deterministic processes mainly governed the fungal community assembly, especially in the rhizospheric fungal communities. Our results demonstrate that tree leaves represent stronger host/fungi preferences than do roots and soils. Together, our findings enhance the understanding of the roles of compartment and plant identity in structuring fungal communities as well as promote fungal diversity maintenance in subtropical mountain forest ecosystems.

Fungi play an essential role in regulating the functioning of terrestrial ecosystems and are sensitive to climate change factors. Climate change incidents, such as N deposition and altered precipitation, create abiotic stress regarding the water use efficiency of soil and nutrient limitation impacting the activity of soil fungi. This study aimed to examine the combined effects of N fertilization and altered precipitation on soil fungal diversity and composition in the desert steppe. In the present study, we carried out a field experiment to assess the soil fungal diversity and composition of the desert steppe in response to N fertilizer (0 or 35 kg N ha−1 year−1) and precipitation changes (control, − 50% precipitation, or + 50% precipitation) in the desert steppe. The study was initiated in 2012, and plant and soil samples were collected after 5 years (August, 2017) of field treatments. High-throughput sequencing was applied to estimate the fungal diversity and composition. The soil fungal communities were dominated by Ascomycota (87.85% ± 1.26%), which primarily drove the fungal community composition. Decreased precipitation promoted strong shifts in fungal community composition under both N fertilizer levels. Increased precipitation significantly reduced Shannon-Wiener indices by 9.96%. The increasing relative abundances of fungal functional groups (lichenized saprotroph, animaland plant pathogens) resulted in a marked shift in fungal community composition from decreased precipitation to increased precipitation, which is attributed to the important role of the Ascomycota phylum in fungal communities. Structural equation modeling (SEM) indicated that C4 biomass was the predominant factor determining the Shannon-Wiener index for these fungi. Direct altered precipitation, indirect soil pH, and C4 biomass together controlled soil fungal community composition, with altered precipitation as the main driver. The interactive effects of N fertilizer and altered precipitation on grassland plant density, biomass, and soil properties may play an essential role in determining fungal diversity and community composition. Precipitation is a primary limiting factor that influences fungal community composition. Effects of N fertilizer on soil fungal community composition are highly dependent on changes in precipitation.

Different fertilization regimes can substantially influence soil fungal community composition, yet fewer studies try to control for the effects of nitrogen input. Here, we investigated the impact of fertilization with equal nitrogen upon soil properties and soil fungal diversity and community composition in the North China Plain in a long-term field experiment. Long-term (32 years) fertilization regimes were applied with equal amounts of nitrogen: no chemical fertilizer or organic manure; chemical fertilization only; organic manure fertilization only, and; combination of 1/2 chemical fertilizer and 1/2 organic manure. Then we investigated the influence of these four fertilization regimes to soil properties, fungal diversity and community composition. The results showed that applying organic manure significantly influenced soil properties. Illumina MiSeq sequencing and its analysis revealed that organic manure fertilization significantly changed soil fungal alpha diversity, but chemical fertilization did not. Although soil fungal community composition did not differ significantly among all the fertilization regimes at the phylum and class levels, they did show differences in the abundance of dominant fungi. Yet at the genus level, soil fungal community composition, abundance, and beta diversity was affected by all fertilization regimes. Application of organic manure also reduced the abundance of soil-born fungal pathogens such as Fusarium. Our results suggest that long-term application of organic manure could markedly improve soil properties, altering soil fungal community composition and its diversity. Moreover, organic manure fertilization could limit soil-born fungal diseases, to further contribute to soil ecosystem sustainability.

Simple SummaryThe gut fungi assist the host in various physiological activities, homeostasis, immune responses, and growth. The diversity and community composition of gut fungi are driven by multiple factors, including diet, environmental exposure, habitat type, and seasonal migration. Migratory birds have a peculiar life cycle, so it is interesting to understand the ecological function of their “gut fungal microbiome.” Birds are exposed to variable diets, environments, and habitats amid seasonal migration. The hooded crane is known as a long-distance migratory bird, inhabiting both wintering and stopover grounds during seasonal migration. During migratory seasons, it inhabits various habitats and is exposed to variable environments. This study analyzed the shifts between gut fungal diversity and the community composition of the hooded crane at both wintering and stopover sites amid seasonal migration. The gut fungal alpha diversity exhibited a more significant change during winter than in fall and spring. The gut fungal community composition exhibited significant shifts across winter, fall, and spring (ANOSIM, p = 0.001). The pathogenic diversity and relative abundance showed significant differences during winter at the wintering site relative to fall and spring at the stopover site. Moreover, the pathogenic fungal community composition was significantly different during fall, winter, and spring. This work contributes to present essential knowledge about the gut fungal microbiome of hooded cranes amid seasonal migration. This study also implicated that conservation measures for hooded crane conservation should be applied, as the risk of cross-transmission of potential fungal pathogens might increase during seasonal migration.The “gut fungal microbiome” maintains the immune system, homeostasis, and various physiological functions of an organism. Different factors shape and affect gut fungal diversity and community composition, such as environment, habitat type, food resources, and seasons during migration. Wild birds amid migration are exposed to different habitats with different environments, available food resources, and seasons, which may substantially impact their gut fungal community composition and diversity. The hooded crane (Grus monacha) is a known migratory bird that migrates over long distances and is exposed to varied habitats with different environments and food types. We investigated the differences in gut fungal diversity and community composition between wintering and stopover sites amid three migratory seasons. We deduced the gut fungal pathogenic diversity and community composition during winter, fall, and spring by using high throughput sequencing (Illumina Mi-seq), and the internal transcribed region 2 (ITS2) was examined. Samples were collected from Shengjin Lake in the winter and Lindian during the fall and spring. The dominant fungal phyla found across the three seasons were Ascomycota, Basidiomycota, Zygomycota, and Rozellomycota. The gut fungal alpha diversity showed significant shifts during winter at the wintering site compared with the fall and spring seasons at the stopover site. The fungal community composition exhibited a significant change across the three seasons (ANOSIM p = 0.001). The results also demonstrated that the diversity and relative abundance of potential pathogens also showed divergence in winter compared to fall and spring. This study provides the basis for understanding the discrepancy in gut fungal diversity and community composition during migratory seasons at both wintering and stopover grounds. It also suggests that conservation measures should be applied to the conservation of hooded cranes and other wild birds, as the risk of cross-infection increases during seasonal migration.

Soil fungi are a diverse group of organisms extremely crucial to forest nutrient cycling and carbon (C) storage. The elevational pattern of soil microbial diversity has been widely studied, but how soil properties, elevation, and their association affect fungal community in subalpine forests remain to be explored. Here, soil fungal community diversity was investigated using high-throughput sequencing along an elevation gradient (3500–4300 m a.s.l.) in the Abies georgei var. smithii forests, a typical subalpine forest type in the Segila Mountains of Southeast Tibet. Elevation significantly affected the soil properties. Available phosphorus (P), total nitrogen (N), and soil organic C (SOC) increased whereas pH decreased with elevation. A U-shaped pattern for fungal diversity was found in topsoil while a slight monotonically decreasing pattern in subsoil layer across the elevation gradient. Basidiomycota, Ascomycota, and Mortierellomycota were the top three dominant phyla, and ectomycorrhizal fungi were the dominant functional fungi group in the A. georgei forest soils. Significant negative correlations were observed between fungal OTUs richness and NO 3 - ( r = −0.431, p < 0.01), indicating that soil N availability is an important factor for fungal diversity. Based on redundancy analysis, soil factors explained 32.1% of the total variations in composition structure of fungal community. Soil pH, moisture, and fertilities (i.e., SOC, TN, TP, NO 3 - ) were independent factors affecting the fungal community composition. Collectively, the alterations in soil environmental factor across elevations are essential in shaping the soil fungal diversity and community composition in the A. georgei forests in Segila Mountains, Southeast Tibet. • Elevation significantly increased soil organic C and nutrients while decreased pH. • Topsoil fungal richness showed a U-shaped pattern with increasing elevation. • Ectomycorrhizal fungi (EcM) were the dominant functional group across elevation. • NO 3 - was the prominent driving factor on fungal diversity in the A. georgei forests.

The differences in soil fungal communities in four agricultural areas growing wheat (Triticum aestivum), rapeseed (Brassica napus), and barley (Hordeum vulgare) in the Qinghai Province, namely the Dulan (DL), Gonghe (GH), Huzhu (HZ), and Datong (DT) counties, were investigated using high-throughput sequencing. The region showed highly significant effects on soil pH, organic matter, ammonium nitrogen, nitrate nitrogen, total phosphate, effective phosphate, total sulfur, and effective sulfur (p &lt; 0.01). The crop type resulted in highly significant (p &lt; 0.01) variations in total phosphate and effective phosphate. Principal coordinates analysis and nonmetric multidimensional scaling revealed significant differences in soil fungal diversity and fungal community composition in the soils of three crops or four regions (p &lt; 0.05). Although the soils of the four regions or three crops had similar dominant phyla, classes, and genera, these taxa differed in terms of their relative abundance. Four, 12, 15, and 16 biomarkers with significant linear discriminant analysis effect sizes were identified in the HZ, DL, GH, and DT groups, respectively. A total of 36, 12, and eight significant biomarkers were observed in the wheat, rapeseed, and barley soils, respectively. In addition, altitude and soil physicochemical properties had significant relationships with fungal diversity and community composition (p &lt; 0.05, p &lt; 0.01).

Impacts of invasive plants on soil fungi and on above- and belowground plant diversity in temperate forests

Atmospheric nitrogen (N) deposition and phosphorus (P) addition both can change soil bacterial and fungal community structure with a consequent impact on ecosystem functions. However, which factor plays an important role in regulating responses of bacterial and fungal community to N and P enrichments remains unclear. We conducted a manipulative experiment to simulate N and P inputs (10 g N · m−2 · yr−1 NH4NO3 or 10 g P · m−2 · yr−1 NaH2PO4) and compared their effects on soil bacterial and fungal species richness and community composition. The results showed that the addition of N significantly increased NH4+ and Al3+ by 99.6% and 57.4%, respectively, and consequently led to a decline in soil pH from 4.18 to 3.75 after a 5-year treatment. P addition increased Al3+ and available P by 27.0% and 10-fold, respectively, but had no effect on soil pH. N addition significantly decreased bacterial species richness and Shannon index and resulted in a substantial shift of bacterial community composition, whereas P addition did not. Neither N nor P addition changed fungal species richness, Shannon index, and fungal community composition. A structural equation model showed that the shift in bacterial community composition was related to an increase in soil acid cations. The principal component scores of soil nutrients showed a significantly positive relationship with fungal community composition. Our results suggest that N and P additions affect soil bacterial and fungal communities in different ways in subtropical forest. These findings highlight how the diversity of microbial communities of subtropical forest soil will depend on future scenarios of anthropogenic N deposition and P enrichment, with a particular sensitivity of bacterial community to N addition.