Responses of belowground communities to large aboveground herbivores: Meta‐analysis reveals biome‐dependent patterns and critical research gaps

The importance of herbivore–plant and soil biota–plant interactions in terrestrial ecosystems is amply recognized, but the effects of aboveground herbivores on soil biota remain challenging to predict. To find global patterns in belowground responses to vertebrate herbivores, we performed a meta‐analysis of studies that had measured abundance or activity of soil organisms inside and outside field exclosures (areas that excluded herbivores). Responses were often controlled by climate, ecosystem type, and dominant herbivore identity. Soil microfauna and especially root‐feeding nematodes were negatively affected by herbivores in subarctic sites. In arid ecosystems, herbivore presence tended to reduce microbial biomass and nitrogen mineralization. Herbivores decreased soil respiration in subarctic ecosystems and increased it in temperate ecosystems, but had no net effect on microbial biomass or nitrogen mineralization in those ecosystems. Responses of soil fauna, microbial biomass, and nitrogen mineralization shifted from neutral to negative with increasing herbivore body size. Responses of animal decomposers tended to switch from negative to positive with increasing precipitation, but also differed among taxa, for instance Oribatida responded negatively to herbivores, whereas Collembola did not. Our findings imply that losses and gains of aboveground herbivores will interact with climate and land use changes, inducing functional shifts in soil communities. To conceptualize the mechanisms behind our findings and link them with previous theoretical frameworks, we propose two complementary approaches to predict soil biological responses to vertebrate herbivores, one focused on an herbivore body size gradient, and the other on a climate severity gradient. Major research gaps were revealed, with tropical biomes, protists, and soil macrofauna being especially overlooked.     

Aggravated phosphorus limitation on biomass production under increasing nitrogen loading: a meta‐analysis

Nitrogen (N) and phosphorus (P), either individually or in combination, have been demonstrated to limit biomass production in terrestrial ecosystems. Field studies have been extensively synthesized to assess global patterns of N impacts on terrestrial ecosystem processes. However, to our knowledge, no synthesis has been done so far to reveal global patterns of P impacts on terrestrial ecosystems, especially under different nitrogen (N) levels. Here, we conducted a meta‐analysis of impacts of P addition, either alone or with N addition, on aboveground (AGB) and belowground biomass production (BGB), plant and soil P concentrations, and N : P ratio in terrestrial ecosystems. Overall, our meta‐analysis quantitatively confirmed existing notions: (i) colimitation of N and P on biomass production and (ii) more P limitation in tropical forest than other ecosystems. More importantly, our analysis revealed new findings: (i) P limitation on biomass production was aggravated by N enrichment and (ii) plant P concentration was a better indicator of P limitation than soil P availability. Specifically, P addition increased AGB and BGB by 34% and 13%, respectively. The effect size of P addition on biomass production was larger in tropical forest than grassland, wetland, and tundra and varied with P fertilizer forms, P addition rates, or experimental durations. The P‐induced increase in biomass production and plant P concentration was larger under elevated than ambient N. Our findings suggest that the global limitation of P on biomass production will become severer under increasing N fertilizer and deposition in the future.     

Multi‐symbiotic systems: functional implications of the coexistence of grass–endophyte and legume–rhizobia symbioses

The coexistence of symbionts with different functional roles in co‐occurring plants is highly probable in terrestrial ecosystems. Analyses of how plants and microbes interact above‐ and belowground in multi‐symbiotic systems are key to understand community structure and ecosystem functioning. We performed an outdoor experiment in mesocosms to investigate the consequences of the interaction of a provider belowground symbiont of legumes (nitrogen‐fixing bacteria) and a protector aerial fungal symbiont of grasses (Epichloё endophyte) on nitrogen dynamics and aboveground net primary productivity. Four plants of Trifolium repens (Trifolium, a perennial legume) either inoculated or not with Rhizobium leguminosarum, grew surrounded by 16 plants of Lolium multiflorum (Lolium, an annual grass), with either low or high levels of the endophyte Neotyphodium occultans. After five months, we quantified the number of nodules in Trifolium roots, shoot biomass of both plant species, and the contribution of atmospheric nitrogen fixation vs. soil nitrogen uptake to above ground nitrogen in each plant species. The endophyte increased grass biomass production (+ 16%), and nitrogen uptake from the soil – the main source for the grass. Further, it reduced the nodulation of neighbour Trifolium plants (?50%). Notably, due to a compensatory increase in nitrogen fixation per nodule, this reduced neither its atmospheric nitrogen fixation – the main source of nitrogen for the legume – nor its biomass production, both of which were doubled by rhizobial inoculation. In consequence, the total amount of nitrogen in aboveground biomass and aboveground productivity were greatest in mesocosms with both symbionts (i.e. high rhizobia + high endophyte). These results show that, in spite of the deleterious effect of the endophyte on the establishment of the rhizobia–legume symbiosis, the coexistence of these symbionts, leading to additive effects on nitrogen capture and aboveground productivity, can generate complementarity on the functioning of multi‐symbiotic systems.     

Effects of climate warming on carbon fluxes in grasslands— A global meta‐analysis

Climate warming will affect terrestrial ecosystems in many ways, and warming‐induced changes in terrestrial carbon (C) cycling could accelerate or slow future warming. So far, warming experiments have shown a wide range of C flux responses, across and within biome types. However, past meta‐analyses of C flux responses have lacked sufficient sample size to discern relative responses for a given biome type. For instance grasslands contribute greatly to global terrestrial C fluxes, and to date grassland warming experiments provide the opportunity to evaluate concurrent responses of both plant and soil C fluxes. Here, we compiled data from 70 sites (in total 622 observations) to evaluate the response of C fluxes to experimental warming across three grassland types (cold, temperate, and semi‐arid), warming methods, and short (≤3 years) and longer‐term (>3 years) experiment lengths. Overall, our meta‐analysis revealed that experimental warming stimulated C fluxes in grassland ecosystems with regard to both plant production (e.g., net primary productivity (NPP) 15.4%; aboveground NPP (ANPP) by 7.6%, belowground NPP (BNPP) by 11.6%) and soil respiration (Rs) (9.5%). However, the magnitude of C flux stimulation varied significantly across cold, temperate and semi‐arid grasslands, in that responses for most C fluxes were larger in cold than temperate or semi‐arid ecosystems. In semi‐arid and temperate grasslands, ecosystem respiration (Reco) was more sensitive to warming than gross primary productivity (GPP), while the opposite was observed for cold grasslands, where warming produced a net increase in whole‐ecosystem C storage. However, the stimulatory effect of warming on ANPP and Rs observed in short‐term studies (≤3 years) in both cold and temperate grasslands disappeared in longer‐term experiments (>3 years). These results highlight the importance of conducting long‐term warming experiments, and in examining responses across a wide range of climate.     

Functional composition of plant communities determines the spatial and temporal stability of soil microbial properties in a long‐term plant diversity experiment

Stable provisioning of ecosystem functions and services is crucial for human well‐being in a changing world. Two essential ecological components driving vital ecosystem functions in terrestrial ecosystems are plant diversity and soil microorganisms. In this study, we tracked soil microbial basal respiration and biomass over a time period of 12 years in a grassland biodiversity experiment (the Jena Experiment) and examined the role of plant diversity and plant functional group composition for the spatial and temporal stability of soil microbial properties (basal respiration and biomass) in bulk‐soil. Spatial and temporal stability were calculated as the inverse coefficient of variation (CV^?1) of soil microbial respiration and biomass measured from soil samples taken over space and time, respectively. We found that 1) plant species richness consistently increased soil microbial properties after a time lag of four years since the establishment of the experimental plots, 2) plant species richness had minor effects on the spatial stability of soil microbial properties, whereas 3) the functional composition of plant communities significantly affected spatial stability of soil microbial properties, with legumes and tall herbs reducing both the spatial stability of microbial respiration and biomass, while grasses increased the latter, and 4) the effect of plant diversity on temporal stability of soil microbial properties turned from being negative to neutral, suggesting that the recovery of soil microbial communities from former arable land‐use takes more than a decade. Our results highlight the importance of plant functional group composition for the spatial and temporal stability of soil microbial properties, and hence for microbially‐driven ecosystem processes, such as decomposition and element cycling, in temperate semi‐natural grassland.     

Responses of terrestrial ecosystems to temperature and precipitation change: a meta‐analysis of experimental manipulation

Global mean temperature is predicted to increase by 2–7 °C and precipitation to change across the globe by the end of this century. To quantify climate effects on ecosystem processes, a number of climate change experiments have been established around the world in various ecosystems. Despite these efforts, general responses of terrestrial ecosystems to changes in temperature and precipitation, and especially to their combined effects, remain unclear. We used meta‐analysis to synthesize ecosystem‐level responses to warming, altered precipitation, and their combination. We focused on plant growth and ecosystem carbon (C) balance, including biomass, net primary production (NPP), respiration, net ecosystem exchange (NEE), and ecosystem photosynthesis, synthesizing results from 85 studies. We found that experimental warming and increased precipitation generally stimulated plant growth and ecosystem C fluxes, whereas decreased precipitation had the opposite effects. For example, warming significantly stimulated total NPP, increased ecosystem photosynthesis, and ecosystem respiration. Experimentally reduced precipitation suppressed aboveground NPP (ANPP) and NEE, whereas supplemental precipitation enhanced ANPP and NEE. Plant productivity and ecosystem C fluxes generally showed higher sensitivities to increased precipitation than to decreased precipitation. Interactive effects of warming and altered precipitation tended to be smaller than expected from additive, single‐factor effects, though low statistical power limits the strength of these conclusions. New experiments with combined temperature and precipitation manipulations are needed to conclusively determine the importance of temperature–precipitation interactions on the C balance of terrestrial ecosystems under future climate conditions.     

Global‐scale impacts of nitrogen deposition on tree carbon sequestration in tropical,temperate, and boreal forests: A meta‐analysis

Elevated nitrogen (N) deposition may increase net primary productivity in N‐limited terrestrial ecosystems and thus enhance the terrestrial carbon (C) sink. To assess the magnitude of this N‐induced C sink, we performed a meta‐analysis on data from forest fertilization experiments to estimate N‐induced C sequestration in aboveground tree woody biomass, a stable C pool with long turnover times. Our results show that boreal and temperate forests responded strongly to N addition and sequestered on average an additional 14 and 13 kg C per kg N in aboveground woody biomass, respectively. Tropical forests, however, did not respond significantly to N addition. The common hypothesis that tropical forests do not respond to N because they are phosphorus‐limited could not be confirmed, as we found no significant response to phosphorus addition in tropical forests. Across climate zones, we found that young forests responded more strongly to N addition, which is important as many previous meta‐analyses of N addition experiments rely heavily on data from experiments on seedlings and young trees. Furthermore, the C–N response (defined as additional mass unit of C sequestered per additional mass unit of N addition) was affected by forest productivity, experimental N addition rate, and rate of ambient N deposition. The estimated C–N responses from our meta‐analysis were generally lower that those derived with stoichiometric scaling, dynamic global vegetation models, and forest growth inventories along N deposition gradients. We estimated N‐induced global C sequestration in tree aboveground woody biomass by multiplying the C–N responses obtained from the meta‐analysis with N deposition estimates per biome. We thus derived an N‐induced global C sink of about 177 (112–243) Tg C/year in aboveground and belowground woody biomass, which would account for about 12% of the forest biomass C sink (1,400 Tg C/year).     

A meta‐analysis of soil salinization effects on nitrogen pools,cycles and fluxes in coastal ecosystems

Salinity intrusion caused by land subsidence resulting from increasing groundwater abstraction, decreasing river sediment loads and increasing sea level because of climate change has caused widespread soil salinization in coastal ecosystems. Soil salinization may greatly alter nitrogen (N) cycling in coastal ecosystems. However, a comprehensive understanding of the effects of soil salinization on ecosystem N pools, cycling processes and fluxes is not available for coastal ecosystems. Therefore, we compiled data from 551 observations from 21 peer‐reviewed papers and conducted a meta‐analysis of experimental soil salinization effects on 19 variables related to N pools, cycling processes and fluxes in coastal ecosystems. Our results showed that the effects of soil salinization varied across different ecosystem types and salinity levels. Soil salinization increased plant N content (18%), soil NH₄⁺ (12%) and soil total N (210%), although it decreased soil NO₃^? (2%) and soil microbial biomass N (74%). Increasing soil salinity stimulated soil N₂O fluxes as well as hydrological NH₄⁺ and NO₂^? fluxes more than threefold, although it decreased the hydrological dissolved organic nitrogen (DON) flux (59%). Soil salinization also increased the net N mineralization by 70%, although salinization effects were not observed on the net nitrification, denitrification and dissimilatory nitrate reduction to ammonium in this meta‐analysis. Overall, this meta‐analysis improves our understanding of the responses of ecosystem N cycling to soil salinization, identifies knowledge gaps and highlights the urgent need for studies on the effects of soil salinization on coastal agro‐ecosystem and microbial N immobilization. Additional increases in knowledge are critical for designing sustainable adaptation measures to the predicted intrusion of salinity intrusion so that the productivity of coastal agro‐ecosystems can be maintained or improved and the N losses and pollution of the natural environment can be minimized.     

氮添加对中国陆地生态系统植物-土壤碳动态的影响

近年来,中国大气氮沉降水平不断增加,过量的活性氮输入深刻影响了我国陆地生态系统碳循环。虽然已有大量的研究报道了模拟氮添加实验对我国陆地生态系统碳动态的影响,但是由于复杂的地理条件和不同的施氮措施,关于植物和土壤碳库对氮添加的一般响应特征和机制仍存在广泛争议。因此,采用整合分析方法,收集整理了172篇已发表的中国野外氮添加试验结果,在全国尺度上探究氮添加对我国陆地生态系统植物和土壤碳动态的影响及其潜在机制。结果表明,氮添加显著促进了植物的碳储存,地上和地下生物量均显著增加,且地上生物量比地下生物量增加得多。同时,氮添加显著增加了凋落物质量,但对细根生物量没有显著影响。氮添加显著降低了植物叶片、凋落物和细根的碳氮比。总体上,氮添加显著增加了土壤有机碳含量并降低了土壤pH值,但对可溶性有机碳、微生物生物量碳和土壤呼吸的影响并不显著。在不同的地理条件下,土壤有机碳含量对氮添加的响应呈现增加、减少或不变的不同趋势。回归分析表明,地上生物量与土壤有机碳含量之间,以及微生物生物量碳与土壤有机碳含量之间呈负相关关系。虽然氮添加通过增加凋落物质量显著促进了植物碳输入,但同时也会通过刺激微生物降解来增加土...     

A cross‐system meta‐analysis reveals coupled predation effects on prey biomass and diversity

Predator diversity and abundance are under strong human pressure in all types of ecosystems. Whereas predator potentially control standing biomass and species interactions in food webs, their effects on prey biomass and especially prey biodiversity have not yet been systematically quantified. Here, we test the effects of predation in a cross‐system meta‐analysis of prey diversity and biomass responses to local manipulation of predator presence. We found 291 predator removal experiments from 87 studies assessing both diversity and biomass responses. Across ecosystem types, predator presence significantly decreased both biomass and diversity of prey across ecosystems. Predation effects were highly similar between ecosystem types, whereas previous studies had shown that herbivory or decomposition effects differed fundamentally between terrestrial and aquatic systems based on different stoichiometry of plant material. Such stoichiometric differences between systems are unlikely for carnivorous predators, where effect sizes on species richness strongly correlated to effect sizes on biomass. However, the negative predation effect on prey biomass was ameliorated significantly with increasing prey richness and increasing species richness of the manipulated predator assemblage. Moreover, with increasing richness of the predator assemblage present, the overall negative effects of predation on prey richness switched to positive effects. Our meta‐analysis revealed strong general relationships between predator diversity, prey diversity and the interaction strength between trophic levels in terms of biomass. This study indicates that anthropogenic changes in predator abundance and diversity will potentially have strong effects on trophic interactions across ecosystems. Synthesis The past centuries we have experienced a dramatic loss of top–predator abundance and diversity in most types of ecosystems. To understand the direct consequences of predator loss on a global scale, we quantitatively summarized experiments testing predation effects on prey communities in a cross‐system meta‐analysis. Across ecosystem types, predator presence significantly decreased both biomass and diversity of prey, and predation effects were highly similar. However, with increasing predator richness, the overall negative effects of predation on prey richness switched to positive ones. Anthropogenic changes in predator communities will potentially have strong effects on prey diversity, biomass, and trophic interactions across ecosystems.     

Aquatic insect subsidies influence microbial composition and processing of detritus in near-shore subarctic heathland

Insects are major conduits of resources moving from aquatic to terrestrial systems. While the ecological impacts of insect subsidies are well documented, the underlying mechanisms by which these resources change recipient ecosystems remain poorly understood. Most subsidy inputs enter terrestrial systems as detritus; thus, soil microbes will likely influence the processing of insect subsidies, with implications for plant community composition and net primary productivity (NPP). In a subarctic ecosystem near Lake Mývatn, Iceland where midge (Diptera: Chironomidae) deposition to land is high, we investigated how insect subsidies affected litter processing and microbial communities. We also evaluated how those belowground effects related to changes in inorganic nitrogen, plant composition and NPP. We simulated subsidies by adding midge carcasses to 1-m² heathland plots, where we measured effects on decomposition rates and the plant community. We then studied how fertilization treatments (control, KNO₃ and midge-carcass addition) affected graminoid biomass and inorganic nitrogen in greenhouse experiments. Lastly, we conducted a soil-incubation study with a phospholipid fatty acid analysis (PLFA) to examine how midge addition to heathland soils affected microbial respiration, biomass and composition. We found that midge addition to heathland soils increased litter decomposition and graminoid plant cover by 2.6× and 2×, respectively. Greenhouse experiments revealed similar patterns, with midge carcasses increasing graminoid biomass by at least 2× and NH₄⁺ concentrations by 7×. Our soil-incubation study found that midge carcasses elevated microbial respiration by 64%, microbial biomass by 43% and shifted microbial functional composition. Our findings indicate that insect subsidies can stimulate soil microbial communities and litter decomposition in subarctic heathlands, leading to increased NPP and changes in plant community composition.     

Microbial immobilization drives nitrogen cycling differences among plant species

In many terrestrial ecosystems nitrogen (N) limits productivity and plant community composition is influenced by N availability. However, vegetation is not only controlled by N; plant species may influence ecosystem N dynamics through positive or negative effects on N cycling. We examined four potential mechanisms of plant species effects on nitrogen (N) cycling. We found no species differences in gross ammonification suggesting there are no changes in the ecosystem N cycling rate between the soil organic matter pool (SOM) and the plant/microbial pool. We also found weak differences among plant species in gross nitrification, thus plant species only marginally change the relative sizes of the NH₄⁺ and NO₃^? pools. Next, more than 90% of mineralized N was microbially immobilized, and microbial N immobilization was positively correlated with root biomass. Finally, while species differed in extractable soil NO3^? concentration, these differences were not related to root biomass suggesting that microbial immobilization drives net N mineralization and soil NO₃^? levels. Our results indicate that plant species do not cause feedbacks on the N cycling rate among the three major ecosystem N pools over nine years. However, plant carbon (C) inputs to the soil control microbial N immobilization and thereby change N partitioning between the plant and microbial N pools. Furthermore our results suggest that the SOM pool can act as a strong bottleneck for N cycling in these systems.     

短期氮素添加和模拟放牧对青藏高原高寒草甸生态系统呼吸的影响

氮沉降和放牧是影响草地碳循环过程的重要环境因子,但很少有研究探讨这些因子交互作用对生态系统呼吸的影响。在西藏高原高寒草甸地区开展了外源氮素添加与刈割模拟放牧实验,测定了其对植物生物量分配、土壤微生物碳氮和生态系统呼吸的影响。结果表明:氮素添加显著促进生态系统呼吸,而模拟放牧对其无显著影响,且降低了氮素添加的刺激作用。氮素添加通过提高微生物氮含量和土壤微生物代谢活性,促进植物地上生产,从而增加生态系统的碳排放;而模拟放牧降低了微生物碳含量,且降低了氮素添加的作用,促进根系的补偿性生长,降低了氮素添加对生态系统碳排放的刺激作用。这表明,放牧压力的存在会抑制氮沉降对高寒草甸生态系统碳排放的促进作用,同时外源氮输入也会缓解放牧压力对高寒草甸生态系统生产的负面影响。     

Mechanisms of plant species impacts on ecosystem nitrogen cycling

Plant species are hypothesized to impact ecosystem nitrogen cycling in two distinctly different ways. First, differences in nitrogen use efficiency can lead to positive feedbacks on the rate of nitrogen cycling. Alternatively, plant species can also control the inputs and losses of nitrogen from ecosystems. Our current understanding of litter decomposition shows that most nitrogen present within litter is not released during decomposition but incorporated into soil organic matter. This nitrogen retention is caused by an increase in the relative nitrogen content in decomposing litter and a much lower carbon‐to‐nitrogen ratio of soil organic matter. The long time lag between plant litter formation and the actual release of nitrogen from the litter results in a bottleneck, which prevents feedbacks of plant quality differences on nitrogen cycling. Instead, rates of gross nitrogen mineralization, which are often an order of magnitude higher than net mineralization, indicate that nitrogen cycling within ecosystems is dominated by a microbial nitrogen loop. Nitrogen is released from the soil organic matter and incorporated into microbial biomass. Upon their death, the nitrogen is again incorporated into the soil organic matter. However, this microbial nitrogen loop is driven by plant‐supplied carbon and provides a strong negative feedback through nitrogen cycling on plant productivity. Evidence supporting this hypothesis is strong for temperate grassland ecosystems. For other terrestrial ecosystems, such as forests, tropical and boreal regions, the data are much more limited. Thus, current evidence does not support the view that differences in the efficiency of plant nitrogen use lead to positive feedbacks. In contrast, soil microbes are the dominant factor structuring ecosystem nitrogen cycling. Soil microbes derive nitrogen from the decomposition of soil organic matter, but this microbial activity is driven by recent plant carbon inputs. Changes in plant carbon inputs, resulting from plant species shifts, lead to a negative feedback through microbial nitrogen immobilization. In contrast, there is abundant evidence that plant species impact nitrogen inputs and losses, such as: atmospheric deposition, fire‐induced losses, nitrogen leaching, and nitrogen fixation, which is driven by carbon supply from plants to nitrogen fixers. Additionally, plants can influence the activity and composition of soil microbial communities, which has the potential to lead to differences in nitrification, denitrification and trace nitrogen gas losses. Plant species also impact herbivore behaviour and thereby have the potential to lead to animal‐facilitated movement of nitrogen between ecosystems. Thus, current evidence supports the view that plant species can have large impacts on ecosystem nitrogen cycling. However, species impacts are not caused by differences in plant quantity and quality, but by plant species impacts on nitrogen inputs and losses.     

Trophic and non‐trophic interactions influence the mechanisms underlying biodiversity–ecosystem functioning relationships under different abiotic conditions

Plant diversity effects on ecosystem functioning usually have been studied from a plant perspective. However, the mechanisms underlying biodiversity–ecosystem functioning relationships may also depend on positive or negative interactions between plants and other biotic and abiotic factors, which remain poorly understood. Here we assessed whether plant–herbivore and/or plant–detritivore interactions modify the biodiversity–ecosystem functioning relationship and the mechanisms underlying biodiversity effects, including complementarity and selection effects, biomass allocation, vertical distribution of roots, and plant survival using a microcosm experiment. We also evaluated to what extent trophic and non‐trophic interactions are affected by abiotic conditions by studying drought effects. Our results show that biotic and abiotic conditions influence the shape of the biodiversity–ecosystem function relationship, varying from hump‐shaped to linear. For instance, total biomass increased linearly with plant richness in the presence of detritivores, but not in the absence of detritivores. Moreover, detritivore effects on belowground plant productivity were highly context dependent, varying in the presence of herbivores. Plant interactions with soil biota, especially with herbivores, influenced the mechanisms underlying diversity effects. Herbivores increased plant complementarity and modified biomass allocation and vertical distribution of roots. Furthermore, biotic–abiotic interactions influenced plant productivity differently across plant functional groups. Our findings emphasize the importance of complex biotic interactions underlying biodiversity effects, and that these biotic interactions may change with abiotic conditions. Despite minor changes in productivity in the short‐term, soil biota‐induced changes in plant–plant interactions and plant survival are likely to have significant long‐term consequences for ecosystem functioning. Considering the context‐dependency of multichannel interactions may contribute to reconciling differences among observed patterns in biodiversity studies. Further, abiotic conditions modified the effects of biotic interactions, suggesting that changes in environmental conditions may not only affect ecosystems directly, but also change the biotic composition of and dynamics within ecosystems.     

Responses of terrestrial nitrogen pools and dynamics to different patterns of freeze‐thaw cycle: A meta‐analysis

Altered freeze‐thaw cycle (FTC) patterns due to global climate change may affect nitrogen (N) cycling in terrestrial ecosystems. However, the general responses of soil N pools and fluxes to different FTC patterns are still poorly understood. Here, we compiled data of 1519 observations from 63 studies and conducted a meta‐analysis of the responses of 17 variables involved in terrestrial N pools and fluxes to FTC. Results showed that under FTC treatment, soil NH₄⁺, NO₃^?, NO₃^? leaching, and N₂O emission significantly increased by 18.5%, 18.3%, 66.9%, and 144.9%, respectively; and soil total N (TN) and microbial biomass N (MBN) significantly decreased by 26.2% and 4.7%, respectively; while net N mineralization or nitrification rates did not change. Temperate and cropland ecosystems with relatively high soil nutrient contents were more responsive to FTC than alpine and arctic tundra ecosystems with rapid microbial acclimation. Therefore, altered FTC patterns (such as increased duration of FTC, temperature of freeze, amplitude of freeze, and frequency of FTC) due to global climate warming would enhance the release of inorganic N and the losses of N via leaching and N₂O emissions. Results of this meta‐analysis help better understand the responses of N cycling to FTC and the relationships between FTC patterns and N pools and N fluxes.     

Long‐term nitrogen loading alleviates phosphorus limitation in terrestrial ecosystems

Increased human‐derived nitrogen (N) deposition to terrestrial ecosystems has resulted in widespread phosphorus (P) limitation of net primary productivity. However, it remains unclear if and how N‐induced P limitation varies over time. Soil extracellular phosphatases catalyze the hydrolysis of P from soil organic matter, an important adaptive mechanism for ecosystems to cope with N‐induced P limitation. Here we show, using a meta‐analysis of 140 studies and 668 observations worldwide, that N stimulation of soil phosphatase activity diminishes over time. Whereas short‐term N loading (≤5 years) significantly increased soil phosphatase activity by 28%, long‐term N loading had no significant effect. Nitrogen loading did not affect soil available P and total P content in either short‐ or long‐term studies. Together, these results suggest that N‐induced P limitation in ecosystems is alleviated in the long‐term through the initial stimulation of soil phosphatase activity, thereby securing P supply to support plant growth. Our results suggest that increases in terrestrial carbon uptake due to ongoing anthropogenic N loading may be greater than previously thought.     

Nitrate reduction capacity is limited by belowground plant recovery in a 32‐year‐old created salt marsh

Human activities have decreased global salt marsh surface area with a subsequent loss in the ecosystem functions they provide. The creation of marshes in terrestrial systems has been used to mitigate this loss in marsh cover. Although these constructed marshes may rapidly recover ecosystem structure, biogeochemical processes may be slow to recover. We compared denitrification and dissimilatory nitrate reduction to ammonium (DNRA) rates between a 32‐year‐old excavation‐created salt marsh (CON‐2) and a nearby natural reference salt marsh (NAT) to assess the recovery of ecosystem function. These process rates were measured at 5 cm increments to a depth of 25 cm to assess how plant rooting depth and organic matter accumulation impact N‐cycling. We found that, for both marshes, denitrification and DNRA declined with depth with the highest rates occurring in the top 10 cm. In both systems, N‐retention by DNRA accounted for upwards of 75% of nitrate reduction, but denitrification and DNRA rates were nearly 2× and 3× higher in NAT than CON‐2, respectively. Organic matter was 6× lower in CON‐2, likely due to limited plant belowground biomass production. However, there was no response to glucose additions, suggesting that the microbial functional community, not substrate limitation, limited nitrate reduction recovery. Response ratios showed that denitrification in CON‐2 recovered in surficial sediments where belowground biomass was highest, even though biomass recovery was minimal. This indicates that although recovery of ecosystem function was constrained, it occurred on a faster trajectory than that of ecosystem structure.     

植物与土壤微生物在调控生态系统养分循环中的作用

陆地生态系统的地上、地下是相互联系的。植物与土壤微生物作为陆地生态系统中的重要组成部分, 它们之间的相互作用是生态系统地上、地下结合的重要纽带。该文首先介绍了植物在养分循环中对营养元素的吸收、积累和归还等作用, 阐述了土壤微生物对养分有效性及土壤质量具有重要的作用。其次, 重点综述了植物与土壤微生物之间相互依存、相互竞争的关系。植物通过其凋落物与分泌物为土壤微生物提供营养, 土壤微生物作为分解者提供植物可吸收的营养元素, 比如共生体菌根真菌即可使植物根与土壤真菌达到互惠。然而, 植物的养分吸收与微生物的养分固持同时存在, 因而两者之间存在对养分的竞争。通过植物多样性对土壤微生物多样性的影响分析, 以及土壤微生物直接或间接作用于植物多样性和生产力的分析, 探讨了植物物种多样性与土壤微生物多样性之间的内在联系。针对当前植物与土壤微生物对养分循环的调控机制的争论, 提出植物凋落物是调节植物与土壤微生物养分循环的良好媒介, 植物与土壤微生物的共同作用对维持整个生态系统的稳定性具有重要意义。也指出了目前在陆地生态系统地上、地下研究中存在的不足和亟待解决的问题。     

Abiotic drivers and plant traits explain landscape‐scale patterns in soil microbial communities

The controls on aboveground community composition and diversity have been extensively studied, but our understanding of the drivers of belowground microbial communities is relatively lacking, despite their importance for ecosystem functioning. In this study, we fitted statistical models to explain landscape‐scale variation in soil microbial community composition using data from 180 sites covering a broad range of grassland types, soil and climatic conditions in England. We found that variation in soil microbial communities was explained by abiotic factors like climate, pH and soil properties. Biotic factors, namely community‐weighted means (CWM) of plant functional traits, also explained variation in soil microbial communities. In particular, more bacterial‐dominated microbial communities were associated with exploitative plant traits versus fungal‐dominated communities with resource‐conservative traits, showing that plant functional traits and soil microbial communities are closely related at the landscape scale.