Tree bark represents a large global carbon stock, comprising 2-20 % of woody biomass, and plays a distinct role in carbon and nutrient cycling. It is poorly understood how different abiotic and biotic drivers contribute to bark decomposition globally, or how these relationships play out in a changing climate. Here, we compile a global dataset to estimate the spatial variability of current bark decomposition rates (k values) and employ three machine learning approaches to project the response of k in future climate scenarios. We find that, globally, bark decomposes 2.9 times faster than the rate previously estimated for tree trunks (including bark). Mean k values increase latitudinally from 0.093 year-1 in the boreal region to 1.339 year-1 in the tropical region. Angiosperm bark decomposes faster (k = 0.450 year-1) than gymnosperm bark (0.198 year-1). Climate, especially mean annual temperature and mean annual precipitation, and bark phosphorus concentration, are the predominant factors driving bark decomposition. Four leading future climate scenarios predict bark decomposition to slow down in warm and wet regions and accelerate in cold and dry regions. Our results highlight the importance of considering bark decomposition in carbon cycling models.
Root exudation is a key trait involved in the plant carbon economy that exerts an important control over ecosystem functioning. Yet, our understanding of global patterns of root exudation and associated influencing factors remains limited. Here we assembled a comprehensive dataset of specific root exudation rate (that is, organic carbon exuded per unit root biomass), root traits and environmental covariates from in situ measurements and a literature synthesis. Our results show that non-woody plants have a higher specific root exudation rate than woody plants. Specific root exudation rate exhibits an increasing trend from high to low latitudes, and soil available phosphorus emerges as the primary influencing factor. We also show that specific root exudation rate primarily falls at the 'fast' end of the conservation axis of the root economics space, reflecting a trade-off between carbon allocation to root metabolism and tissue construction. These findings provide a basis for predictive understanding of plant impacts on belowground biogeochemical cycles.
Salinization-alkalization is increasing globally, yet its impact on cross-kingdom microbial correlations remains poorly understood. Herein, cross-kingdom correlations among rhizosphere archaea, bacteria, fungi, and viruses in farmland subjected to gradient saline-alkali stress were investigated. By analyzing changes in the structure and network of rhizosphere soil microbiomes in the typical saline-alkali region of Northeast China, we demonstrated that: (1) Rhizospheric archaeal and viral alpha-diversity increased significantly with saline-alkali stress intensity. Beta-diversity of all four kingdoms (archaea, bacteria, fungi, and viruses) was strongly influenced by both stress intensity and crop genotype, with community divergence primarily shaped by plant sodium concentration. (2) The composition of rhizosphere viral communities varied across crop types and stress levels, with dominant viruses belonging to Cressdnaviricota (infecting eukaryotes), Phixviricota and Uroviricota (both infecting prokaryotes). (3) Under increasing saline-alkali stress, co-occurrence networks exhibited a rise in both overall and cross-kingdom positive correlations, supporting the Stress Gradient Hypothesis (SGH). Notably, significant increases in positive correlations were observed between viruses and fungi, viruses and bacteria. In addition, the proportion of within-kingdom positive correlations declined, suggesting intensified intra-kingdom competition. These findings refine the traditional SGH at the microbial scale by incorporating viral communities, providing critical theoretical guidance for identifying key microbial species in ecological networks. This advances the design of targeted microbial inoculants to enhance saline-alkali soil remediation and broader ecosystem restoration.
Microplastics pollution constitutes a global environmental challenge. Although degradable microplastics can alter soil organic carbon (SOC) mineralization through the priming effect, the key drivers governing this process remain poorly understood, especially in the subsoil (> 30 cm). Combining a meta-analysis, an incubation experiment, and a random forest model, we found that dissolved organic carbon (DOC) from microplastics decomposition affected the priming effect by regulating microbial growth. Specifically, the highly biodegradable polyhydroxyalkanoate promoted microbial growth by increasing the DOC released from microplastics decomposition, thereby enhancing SOC mineralization via a positive priming effect (462 to 1198 mg CO2-C kg-1 soil). Polylactic acid with low biodegradability reduced carbon availability through the sorptive protection of soil DOC, consequently decreasing SOC mineralization by 149 to 268 mg CO2-C kg-1 soil. In the subsoil, fungi primarily used the microplastics-derived DOC as carbon and energy source through enzyme-mediated nutrient mining, which represents the main driving mechanism for the strong positive priming effect. In the topsoil, the preferential absorption and assimilation of microplastics-derived DOC by bacteria promoted the microbial necromass accumulation, thus attenuating the priming effect through microbial and mineral carbon pumps. These findings highlight the role of carbon availability (particularly DOC) in topsoil and subsoil on the microplastics-induced priming effect, and underscore the necessity of incorporating soil carbon status into assessments of microplastics pollution impacts on global carbon budgets and soil health.
Overgrazing has caused widespread global concern. However, how grazing intensity affects ecosystem carbon dioxide (CO2) fluxes with wetness fluctuations in grasslands at a global scale remains poorly understood. Here we measured ecosystem CO2 fluxes covering the 7th to 11th years of a continuous grazing experiment in a typical steppe, and conducted a meta-analysis of 585 observations by collecting relevant data in global grasslands. Our experimental results showed that mean gross primary productivity (GPP) and ecosystem respiration (ER) were decreased under heavy grazing (HG) compared with control without grazing (CK), whereas net ecosystem productivity (NEP) was not different between HG and CK in the typical steppe. The NEP was higher under light grazing in the wetter years, but it was not different between LG and CK under drier years in the typical steppe. In contrast, grazing decreased GPP, ER and NEP, respectively, in global grasslands. Both our field experiment and the meta-analysis revealed that the response of NEP to light grazing was linearly correlated with the wetness index, which may be mediated by greater plant relative growth rate in wetter years. Higher wetness and aboveground biomass (AGB) increased the response of NEP to grazing in global grasslands. However, heavy grazing reduced NEP and AGB even under higher wetness indices in global grasslands, resulting from a loss of their resilience in long-term heavy grazing. These findings indicate that light grazing appears to be a promising management to promote plant relative growth rate and CO2 flux sequestration. Furthermore, major drivers influencing ecosystem CO2 fluxes in response to grazing were aboveground biomass, grazing intensity and wetness index. Overall, this study provides a global perspective on the response of ecosystem CO2 fluxes to grazing intensity and improves our knowledge of the factors influencing the response of ecosystem CO2 fluxes to grazing intensity. ### Competing Interest Statement The authors have declared no competing interest. the National Natural Science Foundation of China, U22A20559, 32260289, 32271656 Inner Mongolian Key Research and Development and Achievement Transformation Plan Project, 2025YFDZ0062 the Science and Technology Innovation Major Demonstration Project of Inner Mongolia, 2024JBGS0007 the Natural Science Foundation of Inner Mongolia, 2025MS03013, 2025MS03119
In this brief communication, we examined the effects of 4‐year experimental warming on plant species‐specific root exudation dynamics and their associated rhizosphere microbial communities in a temperate grassland ecosystem in Inner Mongolia, China. Our results revealed that Artemisia scoparia exhibited greater sensitivity in carbon and nitrogen exudation rates compared to Stipa krylovii . Furthermore, we identified species‐specific interactions between root exudates and rhizosphere microbiomes: S. krylovii primarily established stronger associations with fungal communities, whereas A. scoparia showed tighter linkages with bacterial communities. These findings underscore the need for future research to investigate how global warming may differently affect above‐ and belowground processes across plant functional groups in grassland ecosystems, particularly with respect to plant‐microbe‐soil feedback mechanisms. These findings suggest that the rhizosphere of different plants recruits different microbial groups to cope with climate warming. These species‐specific compensatory mechanisms could have important implications for nutrient cycling dynamics and ecosystem stability in grasslands under future climate warming.
Climate change alters biogenic volatile organic compound (BVOC) emissions, yet a quantitative understanding of the interactive effects of warming and drought across plant types and physiological mechanisms remains limited. Here, we quantify the impacts of warming and drought on isoprene and monoterpene (MT) emissions through a meta-analysis of field and greenhouse experiments, complemented by machine learning simulations, and map their spatial distribution. Globally, warming increased isoprene and MT emissions by 107% and 60%, respectively, while drought reduced them by 27% and 33%. Combined warming and drought increased MT emissions by 37%. Neither warming nor drought had a significant effect on sesquiterpene (SQT) emissions. Warming-induced increases in isoprene emissions were positively correlated with changes in photosynthetic electron transport rate (Jf), whereas MT responses declined at higher mean annual temperatures. Under drought, although decoupled from photosynthesis, the responses of both compounds remained positively associated with those of stomatal conductance (gs). Spatially, global 1°C warming elicited the strongest responses in isoprenoid emissions at high altitudes/latitudes (e.g., Siberian Plateau, Arctic, and Tibetan Plateau). Under a low-warming scenario (SSP1-1.9), responses resembled historical trends, while a high-warming scenario (SSP5-8.5) amplified isoprene emissions in C₄ vegetation-dominated regions and suppressed MT responses in boreal coniferous forests. Our findings reveal distinct physiological controls: isoprene exhibits thermal resilience, MT remains temperature-sensitive, and both compounds are regulated by gs during drought. These response patterns vary significantly with plant functional type, experimental design and duration, and initial climate conditions. These mechanistic insights can inform the selection of plant speciesand adaptation strategies for vulnerable ecosystems, thereby helping to mitigate air quality and climate feedbacks under future warming and drought conditions.
Soil detritivores represent a major portion of terrestrial biodiversity and biomass. Their feeding activity accelerates the turnover of organic matter and nutrients, thereby enhancing energy and material flows within soil food webs. Yet, global environmental changes are increasingly disrupting terrestrial ecosystems, threatening soil detritivores and their ecological functions. We hypothesize that global environmental changes will result in a decline in the feeding activity of soil detritivores. To test this, we conduct a global meta-analysis, synthesizing 650 observations from 55 studies. Our results show that global environmental changes reduce the feeding activity of soil detritivores by 47.8% on average. Among GECs, climate change (- 59.8%), chemical pollution (- 57.6%), fire (- 49.1%), and land-use intensification (- 34%) exert the most pronounced detrimental effects. For climate change, drought (- 68.9%) suppresses the feeding activity of soil detritivores to a far greater extent than warming (- 25.4%). Notably, insecticides (- 98.9%), fungicides (- 59.7%), and heavy metals (- 59.5%) are particularly harmful within chemical pollutants. The negative effects of land-use intensification are predominantly driven by mineral fertilization (- 45.6%), whereas grazing (- 20.3%) and tillage (- 11.8%) have minor effects. The magnitude of reductions in soil detritivore feeding activity is strongly regulated by ecosystem type, soil properties (soil organic carbon and pH), and detritivore species richness and abundance. These findings suggest that global environmental change-induced declines in soil detritivore feeding activity may further impair energy transfer within soil food webs, with far-reaching implications for key ecosystem functioning in a rapidly changing global environment.
Trees associated with arbuscular mycorrhizal (AM) or ectomycorrhizal (ECM) fungi create distinct soil environments that influence organic matter accumulation, decomposition, and persistence. As forest composition changes globally, understanding how shifts in mycorrhizal dominance affect soil carbon (C) cycling becomes increasingly important. Priming effects, where fresh C inputs alter the decomposition of existing soil organic matter, represent a critical but poorly understood mechanism, through which tree mycorrhizal associations may regulate soil C storage and loss. To address the role of mycorrhizal associations in regulating priming effects, we collected soils from forest plots differing in their dominance of mycorrhizal-associated tree types in a subtropical forest and quantified priming effects with 13C-labeled glucose. In the organic topsoil, priming effects were positive, but strongly decreased (from 19 to 2.3 μg C g-1 soil after 28 days of incubation) as ECM trees became more dominant. This reduction was linked to higher particulate organic carbon, a greater fungal-to-bacterial ratio, and more genes for degrading recalcitrant C. Conversely, in the mineral subsoil, priming effects were generally negative, with AM-dominated plots (-14 μg C g-1 soil) showing stronger negative priming than in ECM-dominated plots (-3.2 μg C g-1 soil after 28 days of incubation). This is primarily because AM dominated soils have greater availability of mineral-associated organic matter and nitrate content, as well as higher microbial carbon use efficiency. Tree mycorrhizal associations thus regulate priming effects through depth-dependent changes in C stability, nitrogen availability, and microbial functional activity. We conclude that forecasting forest C dynamics in response to mycorrhizal compositional shifts must account for these soil depth-specific mechanisms.
Body size is a master functional trait of soil fauna, reflecting interactions among developmental, life-history, physiological, and ecological processes. Though recognized as a critical parameter, traditional manual measurement remains a major bottleneck due to its low efficiency and subjective error from different labors, hindering progress in large-scale, trait-based soil ecology. To address this gap, we developed FaunaAIM, a novel automated tool for high-throughput extraction of key morphological characteristics of soil fauna from images using machine learning. The workflow introduces an attention-based U-Net model integrated with Convolutional Block Attention Modules (CBAM) for individual segmentation, followed by morphological feature calculation using a series of morphological methods. The model achieved high segmentation accuracy (97.3
Deadwood is a critical component of ecosystems, contributing to long-term carbon sequestration, nutrient cycling, and biodiversity maintenance. As a spatially constrained and resource-rich habitat, its invertebrate communities are shaped by fine-scale environmental filtering and species interactions, with fauna-based facilitation playing a key role. Although wood-boring insects are widely recognized as key ecosystem engineers that modify deadwood structure and habitat provisioning, the ecological consequences of their physical tunneling activity are often confounded with concurrent biological processes. As a result, the independent role of structural modification in shaping deadwood macrofauna communities remains poorly understood. This study examined whether simulated longhorn beetle (Cerambycidae) boring, as the physical aspect of ecosystem engineering, influences the diversity and composition of soil macrofauna in deadwood. Using logs from two conifer species, i.e., Picea abies and Abies grandis, we created standardized mechanical drill holes to mimic beetle tunnels and incubated them in a common garden decomposition experiment, for two to three years under similar environmental conditions. Soil macrofauna composition in drilled and undrilled logs was compared to isolate the abiotic effects of cavity formation. Results showed that simulated tunnels altered species composition in some contexts, with taxa such as Isopoda and Diplopoda consistently responding positively to the presence of tunnels. Of particular interest was the positive effect of tunnel boring on the abundance and diversity of soil macrofauna in P. abies after 35 months, which seemed consistent with the positive effect on wood decomposition rate and water holding capacity. These findings provide experimental evidence that small-scale structural heterogeneity made by ecosystem engineers can benefit low-mobility and environmentally sensitive soil macrofauna species by offering microhabitat refuges, while the broader consequences for decomposition are context-dependent and likely shaped by the interaction between physical modification and the biological processes linked to beetle activity.
Background Soil respiration (Rs) is a highly sensitive and consequential process in terrestrial ecosystem carbon (C) cycling under global change scenarios, particularly warming and altered precipitation patterns. However, how warming and delayed rainfall interact to influence Rs and its components remains poorly understood.Methods Here, we combined a field factorial manipulation experiment with delta 13C natural abundance partitioning and a complementary meta-analysis to quantify responses of Rs and its components in grassland ecosystems.Results We found that warming and delayed rainfall exerted contrasting effects on Rs, with warming overall tending to stimulate Rs and delayed rainfall tending to suppress it. Crucially, results from the field experiment indicated that the strong negative effect of delayed rainfall could additively diminish or even reverse the positive net response of Rs to warming, a pattern further supported by the meta-analysis, revealing an average 9.7% reduction in Rs under their combined effects. This interaction was most pronounced during the mid-growing season, as warming exacerbated drought stress under delayed rainfall, potentially shifting the ecosystem from temperature limitation to water limitation. Moreover, such warming-induced drought drove differential responses of respiration components, where autotrophic respiration showed higher sensitivity than heterotrophic respiration.Conclusions Our findings demonstrate that delayed rainfall diminishes the warming-induced stimulation of Rs by exacerbating moisture deficits. Consequently, ignoring the interaction between warming and delayed rainfall may lead to overestimations of positive soil carbon-climate feedback under future compound hot-dry scenarios.
Soil microorganisms are pivotal in sustaining ecosystem multifunctionality, yet their influence depends not only on diversity but also on community assembly and interactions. How these dimensions jointly operate along elevational gradients across different regions remains poorly understood. Here, we conducted a comparative study in two alpine grasslands (Haibei: 3200-4200 m; Damxung: 4400-5200 m) on the Tibetan Plateau, based on 11 elevational sites (33 soil samples) to assess elevational patterns of microbial diversity, community assembly, and interactions, and to clarify their roles in regulating soil multifunctionality. Overall, bacterial diversity decreased with elevation in Haibei but remained relatively stable in Damxung, whereas fungal diversity showed only weak elevational variation in both regions. Assembly processes of bacterial communities in Haibei transitioned from drift to heterogeneous selection and ultimately to homogeneous selection, whereas in Damxung they were consistently dominated by homogeneous selection; fungal communities in both regions were primarily governed by stochastic processes. Network analyses based on a composite network complexity index revealed a consistent decrease in network complexity with elevation in Haibei, whereas fungal network complexity in Damxung exhibited a non-monotonic elevational pattern. Multiple multifunctionality indices consistently indicated that soil multifunctionality was constrained at high elevations in Haibei but was promoted in Damxung. Mechanistically, network complexity was a more robust predictor of soil multifunctionality than diversity, with assembly indirectly influencing soil multifunctionality by restructuring networks. Importantly, the dominant regulatory pathways diverged. The bacterial communities in Haibei were most strongly associated with assembly-mediated network effects, whereas in Damxung, the network complexity of fungal communities covaried with higher soil multifunctionality and was concomitant with plant-derived carbon inputs. Our results validate an integrative framework of assembly-driven, network-mediated functional maintenance, highlighting microbial mechanisms underpinning alpine ecosystem multifunctionality across environmental gradients.
The effects of nitrogen (N) deposition on forest soil organic carbon (SOC) are largely unclear, likely due to the divergent responses of particulate (POC) and mineral-associated carbon (MAOC). Conventional understory inorganic N (UIN) additions neglect canopy processes and the impacts of organic N, potentially misevaluating N deposition effects. This study was conducted in a long-term N addition experiment established in a Moso bamboo forest, which included six treatments combining canopy and understory N additions with organic (urea + glycine) and inorganic (NH4NO3) forms at a rate of 50 kg N·ha−1·yr−1. Litterbags were installed for a two-year decomposition experiment and collected at quarterly intervals, together with concurrent soil sampling under litterbags at 0–10 cm depth. We aimed to examine the effects of canopy vs. understory N addition and organic vs. inorganic N form on soil POC and MAOC concentrations. Our results showed that canopy N additions significantly reduced POC (−15.9%) but did not affect MAOC (P > 0.05). Conversely, understory N additions significantly increased POC (+30.9%) and decreased MAOC (−28.9%). Canopy N additions decreased POC by enhancing peroxidase activity and fungal diversity (FuD), while understory N additions promoted POC by inhibiting litter decomposition. Additionally, understory N addition-induced soil acidification decreased soil Ca2+ concentration, microbial carbon use efficiency, and bacterial necromass C, as well as the release of litter water-soluble compounds, thereby inhibiting MAOC. Moreover, nitrogen forms (organic vs. inorganic) had no effect on SOC fractions. Our findings underscore that canopy and understory N addition approaches differentially regulate SOC fractions by altering litter decomposition–microbial–mineral interactions, and the understory approach may overestimate soil POC gain and MAOC loss driven by atmospheric N deposition.
Nitrogen cycling regulates terrestrial ecosystem productivity and carbon sequestration, yet its response to climate warming remains uncertain. Here, we compiled the most comprehensive dataset to date, integrating 7,941 observations from 413 field warming experiments worldwide with random forest regression and Community Land Model (CLM) simulations. Field warming significantly accelerated nitrogen cycling, increasing N2O emissions (+24.7%), mineralization (+25.8%), nitrification (+51.7%), and denitrification (+41.1%). Soil inorganic nitrogen also increased, while plant nitrogen remained largely unchanged. Elevated natural abundance of 15N indicated that warming alleviates nitrogen limitation and promotes more open nitrogen cycles. Soil moisture, ecosystem type, and warming magnitude were key drivers. N2O emission and nitrification further intensified with increased warming magnitude in random forest analyses. In contrast, CLM5-BGC simulated weak responses in N2O emissions and nitrification and negative changes in nitrogen mineralization, substantially diverging from field observations. These discrepancies highlight the omission of microbial processes and the oversimplification of large-scale ecosystem feedbacks, respectively. Uniquely, this study provides a direct comparison among empirical data, random forest regression, and CLM simulations, revealing discrepancies and their potential causes. Collectively, our findings demonstrate that terrestrial nitrogen cycling is more responsive to climate warming than previously recognized and underscore the importance of integrating multiple analytical approaches to synthesize cross-scale ecological data.
This study investigates the contributions of microbial necromass carbon (MNC) and mineral-associated organic carbon (MAOC) to soil organic carbon (SOC) pools in coastal wetlands and explores the mechanisms underlying their stabilization. Using amino sugar biomarkers and chemical extractions across three coastal wetland soil profiles (0-100 cm), we found that MAOC contributed an average of 74.9% of SOC, suggesting the importance of mineral protection in long-term carbon storage. Although MNC represented only 20.9% of the MAOC pool, on average 70.9% of total MNC was associated with minerals, indicating a coupled interaction between the "microbial carbon pump" and the "mineral carbon pump". The findings further reveal distinct mineral stabilization pathways for fungal and bacterial necromass. Clay and amorphous aluminum oxides were found to enhance the stabilization of both MAOC and fungal necromass, while exchangeable calcium supported the protection of bacterial necromass. In contrast, iron oxides played a comparatively minor role. Cross-ecosystem comparisons pointed to a potential saturation threshold for MAOC accumulation, although no such saturation was observed for mineral associated MNC in this study. These findings highlight the ecosystem-specific nature of mineral-organic matter interactions and provide new insights into the role of MNC in the long-term SOC stabilization in coastal wetlands.
Leaf traits influence biotic interactions and ecosystem functions in forests. However, biogeographic drivers of leaf traits remain highly uncertain, limiting their integration into global vegetation models. Using a global dataset of forest plots, we show that community canopy leaf traits align along three dimensions: leaf economy, leaf density, and nitrogen-to-phosphorus (N:P) ratio. Changes in these trait dimensions across forest communities were driven by different variables: leaf economy was primarily shaped by the proportion of deciduous trees, leaf density by soil available P and temperature, and leaf N:P ratio by temperature. These three leaf trait dimensions together explained 60.2% of the variation in ecosystem-scale forest maximum photosynthesis, with the leaf N:P ratio showing the strongest association, followed by the dimensions of leaf economy and leaf density. Forests with moderate leaf N:P ratios and more acquisitive traits had higher photosynthesis than other forests. Our findings highlight the potential of community canopy leaf traits in predicting biogeographic variation in ecosystem-scale forest functioning.
The soils of alpine meadows on the Tibetan Plateau act as significant carbon reservoirs and are particularly vulnerable to warming. Nevertheless, the long-term (≽20 years) warming impact on SOC composition and deep soil dynamics in alpine meadows remains unclear. This study explored the effects of two decades of warming using open-top chambers on various aspects of alpine meadow ecosystems, including plant community composition and biomass, soil physicochemical characteristics, microbial communities, and SOC content in both bulk soil and its fractions. Prolonged warming had no impact on plant-derived C inputs, as indicated by both unchanged above- and below-ground biomass, but it reduced the light fraction carbon (LF-C) in the surface soil layer (0–10 cm) by 25
The rhizospheric and endophytic microbiota, recognized as the plant's second genome, have gained increasing attention. Root hairs, as pivotal structures for nutrient uptake and exudate secretion, are essential regulators of plant-microbe interactions. However, how root microbiota respond to root hair traits and their potential compensatory mechanisms remains poorly understood. Here, the maize inbred line B73 and its root hairless mutant were compared based on a field experiment. This study integrated rhizosphere and root endosphere microbial community characterization with transcriptomic sequencing and untargeted metabolomic analysis to detect genes with distinct expression patterns and differential metabolites in mutant versus wild-type maize. The findings revealed that root hair absence significantly impairs maize growth, resulting in pronounced differences in the endophytic and rhizospheric microbial communities. Notably, the absence of root hairs particularly impacted endophytic nitrogen-fixing bacteria and even the amino acid composition in root exudates. The change pattern of endophytic nitrogen-fixing bacteria was highly consistent with ferredoxin nitrate reductase genes, further corroborated by RNA-seq, indicating these bacteria partially compensate for the insufficient nitrogen absorption caused by the loss of root hair. Mortierella abundance declined by over 80% in the rhizosphere of hairless root mutant, which strongly associated with many downregulated root exudates. Exogenous inoculation with Mortierella efficiently restored these root metabolites. Mortierella inoculation effectively mitigated the poor growth of the mutant. This study highlights the compensatory role of rhizosphere microorganisms in alleviating the adverse effects of plant gene mutations and highlights the significant influence of specific rhizosphere microorganisms on crop growth.