Depth-dependent effects of ericoid mycorrhizal shrubs on soil carbon and nitrogen pools are accentuated under arbuscular mycorrhizal trees

Plant mycorrhizal associations influence the accumulation and persistence of soil organic matter and could therefore shape ecosystem biogeochemical responses to global changes that are altering forest composition. For instance, arbuscular mycorrhizal (AM) tree dominance is increasing in temperate forests, and ericoid mycorrhizal (ErM) shrubs can respond positively to canopy disturbances. Yet how shifts in the co-occurrence of trees and shrubs with different mycorrhizal associations will affect soil organic matter pools remains largely unknown. We examine the effects of ErM shrubs on soil carbon and nitrogen stocks and indicators of microbial activity at different depths across gradients of AM versus ectomycorrhizal (EcM) tree dominance in three temperate forest sites. We find that ErM shrubs strongly modulate tree mycorrhizal dominance effects. In surface soils, ErM shrubs increase particulate organic matter accumulation and weaken the positive relationship between soil organic matter stocks and indicators of microbial activity. These effects are strongest under AM trees that lack fungal symbionts that can degrade organic matter. In subsurface soil organic matter pools, by contrast, tree mycorrhizal dominance effects are stronger than those of ErM shrubs. Ectomycorrhizal tree dominance has a negative influence on particulate and mineral-associated soil organic matter pools, and these effects are stronger for nitrogen than for carbon stocks. Our findings suggest that increasing co-occurrence of ErM shrubs and AM trees will enhance particulate organic matter accumulation in surface soils by suppressing microbial activity while having little influence on mineral-associated organic matter in subsurface soils. Our study highlights the importance of considering interactions between co-occurring plant mycorrhizal types, as well as their depth-dependent effects, for projecting changes in soil carbon and nitrogen stocks in response to compositional shifts in temperate forests driven by disturbances and global change.     

Soil fertility shifts the relative importance of saprotrophic and mycorrhizal fungi for maintaining ecosystem stability

Soil microbial communities are essential for regulating the dynamics of plant productivity. However, how soil microbes mediate temporal stability of plant productivity at large scales across various soil fertility conditions remains unclear. Here, we combined a regional survey of 51 sites in the temperate grasslands of northern China with a global grassland survey of 120 sites to assess the potential roles of soil microbial diversity in regulating ecosystem stability. The temporal stability of plant productivity was quantified as the ratio of the mean normalized difference vegetation index to its standard deviation. Soil fungal diversity, but not bacterial diversity, was positively associated with ecosystem stability, and particular fungal functional groups determined ecosystem stability under contrasting conditions of soil fertility. The richness of soil fungal saprobes was positively correlated with ecosystem stability under high-fertility conditions, while a positive relationship was observed with the richness of mycorrhizal fungi under low-fertility conditions. These relationships were maintained after accounting for plant diversity and environmental factors. Our findings highlight the essential role of fungal diversity in maintaining stable grassland productivity, and suggest that future studies incorporating fungal functional groups into biodiversity–stability relationships will advance our understanding of their linkages under different fertility conditions.     

Arbuscular mycorrhizae and terrestrial ecosystem processes

Arbuscular mycorrhizal fungi (AMF; phylum Glomeromycota) are ubiquitous in terrestrial ecosystems. Despite their acknowledged importance in ecology, most research on AMF has focused on effects on individual plant hosts, with more recent efforts aimed at the level of the plant community. Research at the ecosystem level is less prominent, but potentially very promising. Numerous human‐induced disturbances (including global change and agro‐ecosystem management) impinge on AMF functioning; hence study of this symbiosis from the ecosystem perspective seems timely and crucial. In this paper, I discuss four (interacting) routes via which AMF can influence ecosystem processes. These include indirect pathways (through changes in plant and soil microbial community composition), and direct pathways (effects on host physiology and resource capture, and direct mycelium effects). I use the case study of carbon cycling to illustrate the potentially pervasive influence of AMF on ecosystem processes. A limited amount of published research on AMF ecology is suited for direct integration into ecosystem studies (because of scale mismatch or ill‐adaptation to the ‘pools and flux’ paradigm of ecosystem ecology); I finish with an assessment of the tools (experimental designs, response variables) available for studying mycorrhizae at the ecosystem scale.     

The potential bias of nitrogen deposition effects on primary productivity and biodiversity

Atmospheric nitrogen (N) deposition is composed of both inorganic nitrogen (IN) and organic nitrogen (ON), and these sources of N may exhibit different impacts on ecosystems. However, our understanding of the impacts of N deposition is largely based on experimental gradients of INs or more rarely ONs. Thus, the effects of N deposition on ecosystem productivity and biodiversity may be biased. We explored the differential impacts of N addition with different IN:ON ratios (0:10, 3:7, 5:5, 7:3, and 10:0) on aboveground net primary productivity (ANPP) of plant community and plant diversity in a typical temperate grassland with a long-term N addition experiment. Soil pH, litter biomass, soil IN concentration, and light penetration were measured to examine the potential mechanisms underlying species loss with N addition. Our results showed that N addition significantly increased plant community ANPP by 68.33%–105.50% and reduced species richness by 16.20%–37.99%. The IN:ON ratios showed no significant effects on plant community ANPP. However, IN-induced species richness loss was about 2.34 times of ON-induced richness loss. Soil pH was positively related to species richness, and they exhibited very similar response patterns to IN:ON ratios. It implies that soil acidification accounts for the different magnitudes of species loss with IN and ON additions. Overall, our study suggests that it might be reasonable to evaluate the effects of N deposition on plant community ANPP with either IN or ON addition. However, the evaluation of N deposition on biodiversity might be overestimated if only IN is added or underestimated if only ON is added.     

荒漠油蒿（Artemisia ordosica）根围AM真菌分布与土壤酶活性

于2007年10月在油蒿(Artemisia ordosica)集中分布区选取4个典型样地,分别从0～10cm、10～20cm、20～30cm、30～40cm和40～50cm土层采集根围土样,分离其丛枝菌根(arbuscular mycorrhiza,AM)真菌并测定了3种土壤酶活性.结果表明,在各样地0～50cm土层中油蒿根系AM真菌菌丝定殖率均很高,有典型的泡囊与丛枝结构.菌丝定殖率与泡囊定殖率呈显著正相关,但菌丝、泡囊和丛枝定殖率与AM真菌孢子密度和土壤酶活性之间都无相关性;孢子密度在不同样地及采样深度间差异明显,与土壤有机质、速效磷和速效氮含量都呈一定正相关关系.孢子密度与脲酶和酸性磷酸酶活性有显著或极显著相关关系,与碱性磷酸酶活性之间的相关性受到土壤pH的显著影响.孢子密度的峰值出现在0～10cm表层土,并随土壤剖面深度增加而降低.土壤酶活性在土壤垂直剖面显示与孢子密度同样的规律.不同样地间AM真菌分布及油蒿根系定殖率的差异表明,油蒿与AM真菌之间有良好共生性,对维护荒漠生态环境系统结构的完整性具有重要意义.     

Consequences of wildfire on ecosystem CO2 and water vapour fluxes in the Great Basin

Increased fire frequency in the Great Basin of North America's intermountain West has led to large‐scale conversion of native sagebrush (Artemisia tridentata Nutt.) communities to postfire successional communities dominated by native and non‐native annual species during the last century. The consequences of this conversion for basic ecosystem functions, however, are poorly understood. We measured net ecosystem CO₂ exchange (NEE) and evapotranspiration (ET) during the first two dry years after wildfire using a 4‐m diameter (16.4 m³) translucent static chamber (dome), and found that both NEE and ET were higher in a postfire successional ecosystem (?0.9–2.6 µ mol CO₂ m^?2 s^?1 and 0.0–1.0 mmol H₂O m^?2 s^?2, respectively) than in an adjacent intact sagebrush ecosystem (?1.2–2.3 µ mol CO₂ m^?2 s^?1 and ?0.1–0.8 mmol H₂O m^?2 s^?2, respectively) during relatively moist periods. Higher NEE in the postfire ecosystem appears to be due to lower rates of above‐ground plant respiration while higher ET appears to be caused by higher surface soil temperatures and increased soil water recharge after rains. These patterns disappeared or were reversed, however, when the conditions were drier. Daily net ecosystem productivity (NEP; g C m^?2 d^?1), derived from multiple linear regressions of measured fluxes with continuously measured climate variables, was very small (close to zero) throughout most of the year. The wintertime was an exception in the intact sagebrush ecosystem with C losses exceeding C gains leading to negative NEP while C balance of the postfire ecosystem remained near zero. Taken together, our results indicate that wildfire‐induced conversion of native sagebrush steppe to ecosystems dominated by herbaceous annual species may have little effect on C balance during relatively dry years (except in winter months) but may stimulate water loss immediately following fires.     

Short‐term effects of a severe drought on avian diversity and abundance in a Pampas Agroecosystem

Agroecosystems are naturally variable at different scales showing strong environmental variations through time and in space. Therefore, temporal dynamics should be taken into account to understand the species‐habitat relationship and provide information for biodiversity management. Droughts are climatic events that introduce variations in environmental conditions by reducing food resources and are increasing in severity and frequency due to global climate change. In 2008, a severe drought occurred in the argentine Pampas, which allowed us to test its short‐term effect on avian diversity patterns according to different land uses. Also, we could test how variations in net primary productivity affected bird populations and explore the usefulness of this ecological process as an indicator of ecosystem service supply applicable to different ecological contexts. We surveyed bird diversity in two consecutive years (2007–2008) in the Rolling Pampas of Argentina, and we assessed environmental attributes at two spatial scales. We explored the relationship between bird diversity and an ecosystem services provision index (ESPI) that uses a proxy of net primary productivity and its intra‐annual variation to test its adequacy in the changing conditions, we conducted our surveys. Results showed that drought affected negatively both species richness and abundance as a consequence of net primary productivity reductions. There was not a clear association of diversity changes with land use, and it is probable that the effect of drought is a complex combination of productivity, land use and spatial scale. ESPI proved robust in front of the environmental changes, and its predictive capacity was better at larger scales. These results are promising for the assessment of ecosystem services provision in a context of global climate changes. Abstract in Spanish is available with online material.     

森林土壤微生物对干旱和氮沉降的响应

干旱和氮沉降深刻影响着人类世森林生态系统的生命活动与物质循环,进而影响全球碳平衡、并反馈作用于气候变化。土壤微生物驱动元素的生物地球化学循环和关键土壤生态过程,在气候变化生物学研究方面具有核心地位和全球重要性。本文综述了干旱和氮沉降对森林土壤细菌和菌根真菌的影响。提出未来应加强全球变化多因子交互作用对土壤微生物多样性、活性与生态功能的研究;建立野外长期定位站,强化亚热带森林生态系统与全球变化研究;注重土壤生物之间互作及网络研究;利用微生物大数据建立相关的机理模型等。从认识微生物多样性和群落组成对全球变化的响应与适应,逐步发展为调控利用微生物群落服务于森林的优化管理、生态资源的合理保护与可持续利用,为充分发挥微生物减缓全球气候变化的作用提供理论基础。     

Interactive effects of global change drivers as determinants of the link between soil biodiversity and ecosystem functioning

Biodiversity, both aboveground and belowground, is negatively affected by global changes such as drought or warming. This loss of biodiversity impacts Earth's ecosystems, as there is a positive relationship between biodiversity and ecosystem functioning (BEF). Even though soils host a large fraction of biodiversity that underlies major ecosystem functions, studies exploring the relationship between soil biodiversity and ecosystem functioning (sBEF) as influenced by global change drivers (GCDs) remain scarce. Here we highlight the need to decipher sBEF relationships under the effect of interactive GCDs that are intimately connected in a changing world. We first state that sBEF relationships depend on the type of function (e.g., C cycling or decomposition) and biodiversity facet (e.g., abundance, species richness, or biomass) considered. Then, we shed light on the impact of single and interactive GCDs on soil biodiversity and sBEF and show that results from scarce studies studying interactive effects range from antagonistic to additive to synergistic when two individual GCDs cooccur. This indicates the need for studies quantitatively accounting for the impacts of interactive GCDs on sBEF relationships. Finally, we provide guidelines for optimized methodological and experimental approaches to study sBEF in a changing world that will provide more valuable information on the real impact of (interactive) GCDs on sBEF. Together, we highlight the need to decipher the sBEF relationship in soils to better understand soil functioning under ongoing global changes, as changes in sBEF are of immediate importance for ecosystem functioning.     

Dynamics within mutualism and the maintenance of diversity: inference from a model of interguild frequency dependence

Numerical models have suggested that the dynamics within mutualisms are not important for the maintenance of diversity. In this study it is demonstrated that the dynamics within mutualism can contribute to the maintenance of diversity within its participants, using a general model of frequency dependence between two mutualistically interacting guilds. Specifically, it is demonstrated that while mutualisms may exhibit positive feedback in density, there may be a negative feedback within a mutualism as a result of the change in composition within the interacting guild. Such a negative feedback results from an asymmetry in the delivery of benefit between participants of the mutualism that generates a negative interguild frequency dependence. This dynamic contributes to the maintenance of diversity within the interacting guilds. Conditions are identified for the maintenance of diversity and the maximization of benefit from mutualism within the context of the model. The utility of these conditions for testing hypotheses using data from the mutualistic interaction between plants and mycorrhizal fungi is then demonstrated.     

农业技术措施对AM真菌群落结构的影响研究进展

农业生态系统中AM真菌多样性丰富,并以独特的群落结构发挥其功能.寄主植物和环境因子对AM真菌群落结构具有重要影响,此外,农业技术措施对农业生态系统中AM真菌群落结构的影响也值得关注.本文系统总结了施肥、灌溉、轮作、间作、土壤耕作、化学药剂等农业技术措施对AM真菌群落结构的影响研究进展,分析了农业技术措施改变AM真菌群落结构的可能机制,探讨了提高农业生态系统中AM真菌多样性的可能途径,提出通过改进施肥体制及其配套技术、增加植物多样性和人工接种AM真菌等可提高农业生态系统中AM真菌多样性;并指出当前存在的问题和今后的研究方向.     

A gradient of nutrient enrichment reveals nonlinear impacts of fertilization on Arctic plant diversity and ecosystem function

Rapid environmental change at high latitudes is predicted to greatly alter the diversity, structure, and function of plant communities, resulting in changes in the pools and fluxes of nutrients. In Arctic tundra, increased nitrogen (N) and phosphorus (P) availability accompanying warming is known to impact plant diversity and ecosystem function; however, to date, most studies examining Arctic nutrient enrichment focus on the impact of relatively large (>25x estimated naturally occurring N enrichment) doses of nutrients on plant community composition and net primary productivity. To understand the impacts of Arctic nutrient enrichment, we examined plant community composition and the capacity for ecosystem function (net ecosystem exchange, ecosystem respiration, and gross primary production) across a gradient of experimental N and P addition expected to more closely approximate warming‐induced fertilization. In addition, we compared our measured ecosystem CO₂ flux data to a widely used Arctic ecosystem exchange model to investigate the ability to predict the capacity for CO₂ exchange with nutrient addition. We observed declines in abundance‐weighted plant diversity at low levels of nutrient enrichment, but species richness and the capacity for ecosystem carbon uptake did not change until the highest level of fertilization. When we compared our measured data to the model, we found that the model explained roughly 30%–50% of the variance in the observed data, depending on the flux variable, and the relationship weakened at high levels of enrichment. Our results suggest that while a relatively small amount of nutrient enrichment impacts plant diversity, only relatively large levels of fertilization—over an order of magnitude or more than warming‐induced rates—significantly alter the capacity for tundra CO₂ exchange. Overall, our findings highlight the value of measuring and modeling the impacts of a nutrient enrichment gradient, as warming‐related nutrient availability may impact ecosystems differently than single‐level fertilization experiments.     

Ecological effects of atmospheric reactive nitrogen deposition on semi-natural terrestrial ecosystems

Evidence that enhanced reactive nitrogen deposition is affecting semi-natural terrestrial ecosystems comes from historic increases in plant tissue N concentrations, correlations between tissue N concentrations and present-day total atmospheric N deposition, changes in plant amino-acid composition and effects on N assimilation. The ecological significance of such changes in biomarkers is uncertain. This paper explores the ecological significance of reactive atmospheric N deposition through a review of previous experimental findings and new experimental evidence from an acidic and a calcareous grassland, both showing phosphorus limitation, and a N-limited Calluna vulgaris (L.) Hull heathland in upland Britain. Nitrogen addition in the range 0–20 g N m⁻² yr⁻¹ initially (years 0–4) increased the growth of Calluna and a decline in some subordinate species. In subsequent years, shoot extension was not stimulated, but winter injury was observed from 1993 onwards, suggesting a strong interaction between N supply and climatic conditions. By contrast, the grasslands showed a small decrease in the cover of higher plants in later years (6–7) of the experimental treatments (0–14 g N m⁻² yr⁻¹) and no growth stimulation. All N treatments reduced the bryophyte cover in the acidic grassland. There were marked effects on below-ground processes, including a sustained stimulation of N mineralization in the grassland soils, and an increase in the bacterial utilization of organic substrates in the heathland, as measured in BIOLOG plates. The results strongly suggest the importance of atmospheric N deposition on microbially driven processes in soils, and are discussed in relation to the scale of potential ecosystem changes and their reversibility by pollution abatement.     

丛枝菌根真菌形态结构、物种多样性和群落组成对氮沉降响应研究进展

工业革命以来,人类活动输入到生态系统中的氮迅速增加,已突破地球所能承受的氮循环阈值。过量氮沉降会造成生物多样性丧失等一系列危害,严重影响生态系统结构和功能。丛枝菌根(AM)真菌能够与大约70%-80%的陆地植物种类形成共生关系,在宿主植物养分吸收、抵抗外界不良环境压力、群落动态和物种共存、生物地球化学循环等方面具有重要的作用。探究AM真菌对氮沉降的响应对认识和把握菌根真菌缓解氮沉降的负面后果,维持生态系统的结构和功能具有重要意义。本文综述了AM真菌的形态结构、物种多样性和群落组成等对氮沉降的响应机制。前人研究表明氮沉降通常降低AM真菌的根系定殖率,减少根外菌丝密度和土壤孢子密度,改变菌丝生长的时间动态;降低AM真菌多样性,改变AM真菌群落组成。氮沉降主要通过缓解植物氮限制、降低植物对菌根的依赖性、减少植物对菌根的碳分配、改变根系和土壤中菌根生物量比率、在植物根内维持稳定的菌根真菌组成作为应对未来扰动的“保险”、改变土壤资源有效性及土壤酸度等直接和间接途径影响AM真菌结构和功能。我们建议在未来研究中整合多组学手段、开展学科交叉,聚焦复杂的生物互作体系对氮沉降的响应机制,以及AM真菌对氮沉降响应的生态后果。     

禾草内生真菌对其它微生物的影响研究进展

禾草内生真菌在宿主植物的茎叶等地上组织中普遍存在,不仅能够提高禾草对生物与非生物逆境的抗性,而且能够对周围环境中的不同微生物类群产生影响。主要总结了禾草Neotyphodium/Epichlo内生真菌对病原真菌、丛枝菌根真菌和土壤微生物的影响及其作用机理。发现禾草内生真菌普遍存在对病原真菌的抑制作用,而对丛枝菌根真菌存在不对称的竞争作用,且因种类而异。禾草内生真菌对土壤微生物群落的作用则会随着土壤类型和时间等外界因素发生变化。禾草内生真菌对不同类群微生物的影响机制主要包括:通过生态位竞争、抑菌物质分泌、诱导抗病性等对病原真菌造成影响;通过根系化学物质释放、营养元素调节、侵染条件差异等对丛枝菌根真菌造成影响;通过根际沉积物和凋落物等对土壤微生物群落造成影响。禾草内生真菌产生的生物碱能提高宿主植物对包括昆虫在内草食动物采食的抗性,影响病原菌的侵入、定殖和扩展;根组织分泌物中包含次生代谢产物能够抑制菌根真菌、土传病原真菌及其它土壤微生物的侵染与群落组成;也可能通过次生代谢物影响禾草的其它抗性。因此,禾草内生真菌在植物-微生物系统中的作用应该给予更多的关注和深入研究。     

氮沉降对杉木人工林土壤有机碳矿化和土壤酶活性的影响

为探讨氮沉降对亚热带森林土壤有机碳矿化及土壤酶活性的影响规律,在杉木人工林中开展了野外模拟N沉降试验。试验设计为4种处理,分别为N0（对照）、N1（60 kg N?hm-2?a-1）、N2（120 kg N?hm-2?a-1）和N3（240 kg N?hm-2?a-1）,每处理重复3次。通过28 d的培养后发现,各土层有机碳日均矿化量随培养时间的延长呈下降趋势,而有机碳累计矿化量则逐步增加。不同氮沉降处理下各土层有机碳累计矿化量总体趋势表现为：随着氮沉降量的增加而降低,日均矿化量降低幅度以N1最大,其次是N0和N2,N3降幅最小。相同N沉降处理下,参与土壤碳循环的6种主要酶（蔗糖酶、纤维素酶、淀粉酶、β-葡糖苷酶、多酚氧化酶、过氧化物酶）活性、土壤有机碳日均矿化量和有机碳累计矿化量均随土层加深而降低。氮沉降对6种土壤酶活性的影响存在差异,对纤维素酶和多酚氧化酶具有促进作用,而对淀粉酶和过氧化物酶表现出一定的抑制作用;中-低氮沉降（N1、N2）对蔗糖酶无影响,而对β-葡糖苷酶具有促进作用,高氮沉降（N3）促进了蔗糖酶活性,但抑制了β-葡糖苷酶活性。表层土壤中,土壤有机碳累积矿化量与土壤纤维素酶、β-葡糖苷酶、过氧化物酶活性呈显著正相关。因此,氮沉降促进了表层土壤纤维素酶、多酚氧化酶和蔗糖酶的活性,但在一定程度上抑制了淀粉酶和过氧化物酶,对土壤有机碳矿化也表现出明显的抑制作用。     

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。     

降水和氮沉降增加对草地土壤酶活性的影响

为探究降水和氮沉降增加对草地生态系统土壤酶活性的影响,于2014年生长季在内蒙古温带典型羊草草原开展了野外原位控制实验。试验共设置降水(对照,W0,自然降水;W15,增加15%的年均降水量)、施氮(对照,CK,0 kg N hm~(-2)a~(-1);低氮,LN,25 kg N hm~(-2)a~(-1);中氮,MN,50 kg N hm~(-2)a~(-1);高氮,HN,100 kg N hm~(-2)a~(-1))及其交互作用等8个不同的处理水平来模拟降水和氮沉降增加的全球变化情景,分别定量探讨了不同水、氮添加条件下草地表层土壤中与氮循环相关的蛋白酶,脲酶,硝酸还原酶,亚硝酸还原酶活性的月动态变化及其与土壤理化性质之间的相关性。研究结果表明:在自然降水条件下,不同施氮水平蛋白酶、脲酶和硝酸还原酶活性无显著差异,亚硝酸还原酶活性相比于对照显著降低;在增加降水条件下,不同施氮水平对蛋白酶和硝酸还原酶活性未产生显著性影响,高氮水平显著降低脲酶和亚硝酸还原酶活性。不同施氮水平是否添加降水对亚硝酸还原酶活性无影响,而增添降水使低氮处理的蛋白酶活性和中、高氮处理水平的硝酸还原酶活性增加、高氮处理的脲酶活性降低。降水在影响蛋白酶和硝酸还原酶活性方面具有主效应,氮沉降在影响亚硝酸还原酶活性方面具有主效应,而降水和施氮处理未表现出明显地交互作用。土壤亚硝酸还原酶活性与土壤碳氮比和NH~+_4-N含量极显著正相关,与NO-3-N含量显著正相关。     

Linking above- and belowground responses to global change at community and ecosystem scales

Cryptic belowground organisms are difficult to observe and their responses to global changes are not well understood. Nevertheless, there is reason to believe that interactions among above- and belowground communities may mediate ecosystem responses to global change. We used grassland mesocosms to manipulate the abundance of one important group of soil organisms, arbuscular mycorrhizal (AM) fungi, and to study community and ecosystem responses to CO₂ and N enrichment. Responses of plants, AM fungi, phospholipid fatty acids and community-level physiological profiles were measured after two growing seasons. Ecosystem responses were examined by measuring net primary production (NPP), evapotranspiration, total soil organic matter (SOM), and extractable mineral N. Structural equation modeling was used to examine the causal relationships among treatments and response variables. We found that while CO₂ and N tended to directly impact ecosystem functions (evapotranspiration and NPP, respectively), AM fungi indirectly impacted ecosystem functions by influencing the community composition of plants and other root fungi, soil fungi and soil bacteria. We found that the mycotrophic status of the dominant plant species in the mesocosms determined whether the presence of AM fungi increased or decreased NPP. Mycotrophic grasses dominated the mesocosm communities during the first growing season, and the mycorrhizal treatments had the highest NPP. In contrast, nonmycotrophic forbs were dominant during the second growing season and the mycorrhizal treatments had the lowest NPP. The composition of the plant community strongly influenced soil N, and the community composition of soil organisms strongly influenced SOM accumulation in the mesocosms. These results show how linkages between above- and belowground communities can determine ecosystem responses to global change.