Plant species richness, elevated CO2, and atmospheric nitrogen deposition alter soil microbial community composition and function

We determined soil microbial community composition and function in a field experiment in which plant communities of increasing species richness were exposed to factorial elevated CO₂ and nitrogen (N) deposition treatments. Because elevated CO₂ and N deposition increased plant productivity to a greater extent in more diverse plant assemblages, it is plausible that heterotrophic microbial communities would experience greater substrate availability, potentially increasing microbial activity, and accelerating soil carbon (C) and N cycling. We, therefore, hypothesized that the response of microbial communities to elevated CO₂ and N deposition is contingent on the species richness of plant communities. Microbial community composition was determined by phospholipid fatty acid analysis, and function was measured using the activity of key extracellular enzymes involved in litter decomposition. Higher plant species richness, as a main effect, fostered greater microbial biomass, cellulolytic and chitinolytic capacity, as well as the abundance of saprophytic and arbuscular mycorrhizal (AM) fungi. Moreover, the effect of plant species richness on microbial communities was significantly modified by elevated CO₂ and N deposition. For instance, microbial biomass and fungal abundance increased with greater species richness, but only under combinations of elevated CO₂ and ambient N, or ambient CO₂ and N deposition. Cellobiohydrolase activity increased with higher plant species richness, and this trend was amplified by elevated CO₂. In most cases, the effect of plant species richness remained significant even after accounting for the influence of plant biomass. Taken together, our results demonstrate that plant species richness can directly regulate microbial activity and community composition, and that plant species richness is a significant determinant of microbial response to elevated CO₂ and N deposition. The strong positive effect of plant species richness on cellulolytic capacity and microbial biomass indicate that the rates of soil C cycling may decline with decreasing plant species richness.     

Fungal root colonization responses in natural grasslands after long-term exposure to elevated atmospheric CO2

Arbuscular mycorrhizae, ubiquitous mutualistic symbioses between plant roots and fungi in the order Glomales, are believed to be important controllers of plant responses to global change, in particular to elevated atmospheric CO₂. In order to test if any effects on the symbiosis can persist after long-term treatment, we examined root colonization by arbuscular mycorrhizal (AM) and other fungi of several plant species from two grassland communities after continuous exposure to elevated atmospheric CO₂ for six growing seasons in the field. For plant species from both a sandstone and a serpentine annual grassland there was evidence for changes in fungal root colonization, with changes occurring as a function of plant host species. We documented decreases in percentage nonmycorrhizal fungal root colonization in elevated CO₂ for several plant species. Total AM root colonization (%) only increased significantly for one out of the five plant species in each grassland. However, when dividing AM fungal hyphae into two groups of hyphae (fine endophyte and coarse endophyte), we could document significant responses of AM fungi that were hidden when only total percentage colonization was measured. We also documented changes in elevated CO₂ in the percentage of root colonized by both AM hyphal types simultaneously. Our results demonstrate that changes in fungal root colonization can occur after long-term CO₂ enrichment, and that the level of resolution of the study of AM fungal responses may have to be increased to uncover significant changes to the CO₂ treatment. This study is also one of the first to document compositional changes in the AM fungi colonizing roots of plants grown in elevated CO₂. Although it is difficult to relate the structural data directly to functional changes, possible implications of the observed changes for plant communities are discussed.     

Species of plants and associated arbuscular mycorrhizal fungi mediate mycorrhizal responses to CO2 enrichment

It has been suggested that enrichment of atmospheric CO₂ should alter mycorrhizal function by simultaneously increasing nutrient‐uptake benefits and decreasing net C costs for host plants. However, this hypothesis has not been sufficiently tested. We conducted three experiments to examine the impacts of CO₂ enrichment on the function of different combinations of plants and arbuscular mycorrhizal (AM) fungi grown under high and low soil nutrient availability. Across the three experiments, AM function was measured in 14 plant species, including forbs, C₃ and C₄ grasses, and plant species that are typically nonmycorrhizal. Five different AM fungal communities were used for inoculum, including mixtures of Glomus spp. and mixtures of Gigasporaceae (i.e. Gigaspora and Scutellospora spp.). Our results do not support the hypothesis that CO₂ enrichment should consistently increase plant growth benefits from AM fungi, but rather, we found CO₂ enrichment frequently reduced AM benefits. Furthermore, we did not find consistent evidence that enrichment of soil nutrients increases plant growth responses to CO₂ enrichment and decreases plant growth responses to AM fungi. Our results show that the strength of AM mutualisms vary significantly among fungal and plant taxa, and that CO₂ levels further mediate AM function. In general, when CO₂ enrichment interacted with AM fungal taxa to affect host plant dry weight, it increased the beneficial effects of Gigasporaceae and reduced the benefits of Glomus spp. Future studies are necessary to assess the importance of temperature, irradiance, and ambient soil fertility in this response. We conclude that the affects of CO₂ enrichment on AM function varies with plant and fungal taxa, and when making predictions about mycorrhizal function, it is unwise to generalize findings based on a narrow range of plant hosts, AM fungi, and environmental conditions.     

Short-term decomposition of litter produced by plants grown in ambient and elevated atmospheric CO2 concentrations

The effects of elevated atmospheric CO₂ (ambient + 340 μmol mol^–1) on above-ground litter decomposition were investigated over a 6-week period using a field-based mesocosm system. Soil respiratory activity in mesocosms incubated in ambient and elevated atmospheric CO₂ concentrations were not significantly different (t-test, P > 0.05) indicating that there were no direct effects of elevated atmospheric CO₂ on litter decomposition. A study of the indirect effects of CO₂ on soil respiration showed that soil mesocosms to which naturally senescent plant litter had been added (0.5% w/w) from the C₃ sedge Scirpus olneyi grown in elevated atmospheric CO₂ was reduced by an average of 17% throughout the study when compared to soil mesocosms to which litter from Scirpus olneyi grown in ambient conditions had been added. In contrast, similar experiments using senescent material from the C₄ grass Spartina patens showed no difference in soil respiration rates between mesocosms to which litter from plants grown in elevated or ambient CO₂ conditions had been added. Analysis of the C:N ratio and lignin content of the senescent material showed that, while the C:N ratio and lignin content of the Spartina patens litter did not vary with atmospheric CO₂ conditions, the C:N ratio (but not the lignin content) of the litter from Scirpus olneyi was significantly greater (t-test;P < 0.05) when derived from plants grown under elevated CO₂ (105:1 compared to 86:1 for litter derived from Scirpus olneyi grown under ambient conditions). The results suggest that the increased C:N ratio of the litter from the C₃ plant Scirpus olneyi grown under elevated CO₂ led to the lower rates of biodegradation observed as reduced soil respiration in the mesocosms. Further long-term experiments are now required to determine the effects of elevated CO₂ on C partitioning in terrestrial ecosystems.     

Effects of elevated [CO2] at the community level mediated by root symbionts

This review examines the effects of elevated [CO₂] on plant symbioses with mycorrhizal fungi and root nodule bacteria, with emphasis on community and ecosystem processes. The effects of elevated [CO₂] on the relationships between single plant species and root symbionts are considered first. There is some evidence that plant infection by and/or biomass of root symbionts are stimulated by elevated [CO₂], but growth enhancement of the host seemingly depends on its degree of dependence on symbiosis and on soil nutrient availability. Second, the effects of elevated [CO₂] on the relationships between plant multispecies assemblages and soil, and likely impacts on above-ground and belowground diversity, are analysed. Experimental and modelling work have suggested the existence of complex feedbacks in the responses of plants and the rhizosphere to CO₂ enrichment. By modifying C inputs from plants to soil, elevated [CO₂] may affect the biomass, the infectivity, and the species/isolate composition of root symbionts. This has the potential to alter community structure and ecosystem functioning. Finally, the incorporation of type and degree of symbiotic dependence into the definition of plant functional types, and into experimental work within the context of global change research, are discussed. More experimental work on the effects of elevated [CO₂] at the community/ecosystem level, explicitly considering the role of root symbioses, is urgently needed.     

Impacts of long‐term elevated atmospheric CO2 concentrations on communities of arbuscular mycorrhizal fungi

The ecological impacts of long‐term elevated atmospheric CO₂ (eCO₂) levels on soil microbiota remain largely unknown. This is particularly true for the arbuscular mycorrhizal (AM) fungi, which form mutualistic associations with over two‐thirds of terrestrial plant species and are entirely dependent on their plant hosts for carbon. Here, we use high‐resolution amplicon sequencing (Illumina, HiSeq) to quantify the response of AM fungal communities to the longest running (>15 years) free‐air carbon dioxide enrichment (FACE) experiment in the Northern Hemisphere (GiFACE); providing the first evaluation of these responses from old‐growth (>100 years) semi‐natural grasslands subjected to a 20% increase in atmospheric CO₂. eCO₂ significantly increased AM fungal richness but had a less‐pronounced impact on the composition of their communities. However, while broader changes in community composition were not observed, more subtle responses of specific AM fungal taxa were with populations both increasing and decreasing in abundance in response to eCO₂. Most population‐level responses to eCO₂ were not consistent through time, with a significant interaction between sampling time and eCO₂ treatment being observed. This suggests that the temporal dynamics of AM fungal populations may be disturbed by anthropogenic stressors. As AM fungi are functionally differentiated, with different taxa providing different benefits to host plants, changes in population densities in response to eCO₂ may significantly impact terrestrial plant communities and their productivity. Thus, predictions regarding future terrestrial ecosystems must consider changes both aboveground and belowground, but avoid relying on broad‐scale community‐level responses of soil microbes observed on single occasions.     

Alteration of Soil Carbon Pools and Communities of Mycorrhizal Fungi in Chaparral Exposed to Elevated Carbon Dioxide

We examined the effects of atmospheric carbon dioxide (CO₂) enrichment on belowground carbon (C) pools and arbuscular mycorrhizal (AM) fungi in a chaparral community in southern California. Chambers enclosing intact mesocosms dominated by Adenostoma fasciculatum were exposed for 3.5 years to CO₂ levels ranging from 250 to 750 ppm. Pools of total C in bulk soil and in water-stable aggregates (WSA) increased 1.5- and threefold, respectively, between the 250- and 650-ppm treatments. In addition, the abundance of live AM hyphae and spores rose markedly over the same range of CO₂, and the community composition shifted toward dominance by the AM genera Scutellospora and Acaulospora. Net ecosystem exchange of C with the atmosphere declined with CO₂ treatment. It appears that under CO₂ enrichment, extra C was added to the soil via AM fungi. Moreover, AM fungi were predominant in WSA and may shunt C into these aggregates versus bulk soil. Alternatively, C may be retained longer within WSA than within bulk soil. We note that differences between the soil fractions may act as a potential feedback on C cycling between the soil and atmosphere.     

Seven years of carbon dioxide enrichment, nitrogen fertilization and plant diversity influence arbuscular mycorrhizal fungi in a grassland ecosystem

? We tested the prediction that the abundance and diversity of arbuscular mycorrhizal (AM) fungi are influenced by resource availability and plant community composition by examining the joint effects of carbon dioxide (CO(2) ) enrichment, nitrogen (N) fertilization and plant diversity on AM fungi. ? We quantified AM fungal spores and extramatrical hyphae in 176 plots after 7 yr of treatment with all combinations of ambient or elevated CO(2) (368 or 560 ppm), with or without N fertilization (0 or 4 g Nm(-2) ), and one (monoculture) or 16 host plant species (polyculture) in the BioCON field experiment at Cedar Creek Ecosystem Science Reserve, Minnesota, USA. ? Extramatrical hyphal lengths were increased by CO(2) enrichment, whereas AM spore abundance decreased with N fertilization. Spore abundance, morphotype richness and extramatrical hyphal lengths were all greater in monoculture plots. A structural equation model showed AM fungal biovolume was most influenced by CO(2) enrichment, plant community composition and plant richness, whereas spore richness was most influenced by fungal biovolume, plant community composition and plant richness. ? Arbuscular mycorrhizal fungi responded to differences in host community and resource availability, suggesting that mycorrhizal functions, such as carbon sequestration and soil stability, will be affected by global change.     

The rate of progression and stability of progressive nitrogen limitation at elevated atmospheric CO2 in a grazed grassland over 11 years of Free Air CO2 enrichment

A decline in the availability of nitrogen (N) for plant growth (progressive nitrogen limitation or PNL) is a feedback that could constrain terrestrial ecosystem responses to elevated atmospheric CO₂. Several long-term CO₂ enrichment experiments have measured changes in plant and soil pools and fluxes consistent with PNL but evidence for PNL in grasslands is limited. In an 11 year Free Air CO₂ Enrichment (FACE) experiment on grazed grassland we found the amount of N harvested in aboveground plant biomass was greater at elevated CO₂ but declined over time to be indistinguishable from ambient after 5 years. Re-wetting after a major drought resulted in a large input of N from mineralisation and a return to a higher N harvested under elevated CO₂ followed by a further decline. Over these two periods the amount of N in soil significantly increased at elevated CO₂. Data from mesocosms introduced into the rings at intervals, and therefore having different lengths of exposure to CO₂, showed plant N availability declined at elevated CO₂ reaching a new equilibrium after 6 years of exposure. We conclude that the availability of N for plants in this grassland is dynamic but the underlying trend at elevated CO₂ is for PNL.     

Soil microbial response in tallgrass prairie to elevated CO2

Terrestrial responses to increasing atmospheric CO₂ are important to the global carbon budget. Increased plant production under elevated CO₂ is expected to increase soil C which may induce N limitations. The objectives of this study were to determine the effects of increased CO₂ on 1) the amount of carbon and nitrogen stored in soil organic matter and microbial biomass and 2) soil microbial activity. A tallgrass prairie ecosystem was exposed to ambient and twice-ambient CO₂ concentrations in open-top chambers in the field from 1989 to 1992 and compared to unchambered ambient CO₂ during the entire growing season. During 1990 and 1991, N fertilizer was included as a treatment. The soil microbial response to CO₂ was measured during 1991 and 1992. Soil organic C and N were not significantly affected by enriched atmospheric CO₂. The response of microbial biomass to CO₂ enrichment was dependent upon soil water conditions. In 1991, a dry year, CO₂ enrichment significantly increased microbial biomass C and N. In 1992, a wet year, microbial biomass C and N were unaffected by the CO₂ treatments. Added N increased microbial C and N under CO₂ enrichment. Microbial activity was consistently greater under CO₂ enrichment because of better soil water conditions. Added N stimulated microbial activity under CO₂ enrichment. Increased microbial N with CO₂ enrichment may indicate plant production could be limited by N availability. The soil system also could compensate for the limited N by increasing the labile pool to support increased plant production with elevated atmospheric CO₂. Longer-term studies are needed to determine how tallgrass prairie will respond to increased C input.     

Plant nitrogen acquisition and interactions under elevated carbon dioxide: impact of endophytes and mycorrhizae

Both endophytic and mycorrhizal fungi interact with plants to form symbiosis in which the fungal partners rely on, and sometimes compete for, carbon (C) sources from their hosts. Changes in photosynthesis in host plants caused by atmospheric carbon dioxide (CO₂) enrichment may, therefore, influence those mutualistic interactions, potentially modifying plant nutrient acquisition and interactions with other coexisting plant species. However, few studies have so far examined the interactive controls of endophytes and mycorrhizae over plant responses to atmospheric CO₂ enrichment. Using Festuca arundinacea Schreb and Plantago lanceolata L. as model plants, we examined the effects of elevated CO₂ on mycorrhizae and endophyte (Neotyphodium coenophialum) and plant nitrogen (N) acquisition in two microcosm experiments, and determined whether and how mycorrhizae and endophytes mediate interactions between their host plant species. Endophyte‐free and endophyte‐infected F. arundinacea varieties, P. lanceolata L., and their combination with or without mycorrhizal inocula were grown under ambient (400 μmol mol⁻¹) and elevated CO₂ (ambient + 330 μmol mol⁻¹). A ¹⁵N isotope tracer was used to quantify the mycorrhiza‐mediated plant acquisition of N from soil. Elevated CO₂ stimulated the growth of P. lanceolata greater than F. arundinacea, increasing the shoot biomass ratio of P. lanceolata to F. arundinacea in all the mixtures. Elevated CO₂ also increased mycorrhizal root colonization of P. lanceolata, but had no impact on that of F. arundinacea. Mycorrhizae increased the shoot biomass ratio of P. lanceolata to F. arundinacea under elevated CO₂. In the absence of endophytes, both elevated CO₂ and mycorrhizae enhanced ¹⁵N and total N uptake of P. lanceolata but had either no or even negative effects on N acquisition of F. arundinacea, altering N distribution between these two species in the mixture. The presence of endophytes in F. arundinacea, however, reduced the CO₂ effect on N acquisition in P. lanceolata, although it did not affect growth responses of their host plants to elevated CO₂. These results suggest that mycorrhizal fungi and endophytes might interactively affect the responses of their host plants and their coexisting species to elevated CO₂.     

Plant growth and reproduction along CO2 gradients: non-linear responses and implications for community change

The effects of rising atmospheric CO₂ concentrations on natural plant communities will depend upon the cumulative responses of plant growth and reproduction to gradual, incremental changes in climatic conditions. We analysed published studies of plant responses to elevated CO₂ to address whether reproductive and total biomass exhibit similar enhancement to elevated vs. ambient CO₂ concentrations, and to assess the patterns of plant response along gradients of CO₂ concentrations. In six annual plant species, mean enhancement at double ambient vs. ambient CO₂ was 1.13 for total biomass and 1.30 for reproductive biomass. The two measures were significantly correlated, but there was considerable scatter in the relationship, indicating that reproductive responses cannot be consistently predicted from enhancement of total biomass. Along experimental CO₂ gradients utilizing three concentrations, there was a great diversity of response patterns, including positive, negative, non-monotonic and non-significant (flat) responses. The distribution of response patterns differed for plants grown in stands compared to those grown individually. Positive responses were less frequent in competitive environments, and non-monotonic responses were more frequent. These results emphasize that interpolation of plant response based on enhancement ratios measured at elevated vs. ambient CO₂ concentrations is not sufficient to predict community responses to incremental changes in atmospheric conditions. The consequences of differential response patterns were assessed in a simulation of community dynamics for four species of annual plants. The model illustrates that the final community composition at a future point in time depends critically on both the magnitude and the rate of increase of atmospheric CO₂.     

The influence of plant species,fertilization and elevated CO2 on soil aggregate stability

We tested the effects of plant species, fertilization and elevated CO₂ on water-stable soil aggregation. Five annual grassland species and a plant community were grown in outdoor mesocosms for 4 years, with and without NPK fertilization, at ambient or elevated atmospheric CO₂ concentrations. Aggregate stability (resistance of aggregates to slaking) in the top 0.15 m of soil differed among plant species. However, the more diverse plant community did not enhance aggregate stability relative to most monocultures. Species differences in aggregate stability were positively correlated with soil active bacterial biomass, but did not correlate with root biomass or fungal length. Plant species did not affect aggregate stability lower in the soil profile (0.15–0.45 m), where soil biological activity is generally decreased. Elevated CO₂ and NPK fertilization altered many of the factors known to influence aggregation, but did not affect water-stable aggregation at either depth, in any of the plant treatments. These results suggest that global changes will alter soil structure primarily due to shifts in vegetation composition.     

Responses of an old-field plant community to interacting factors of elevated [CO2], warming, and soil moisture

Aims: The direct effects of atmospheric and climatic change factors—atmospheric[CO₂], air temperature and changes in precipitation—canshape plant community composition and alter ecosystem function.It is essential to understand how these factors interact tomake better predictions about how ecosystems may respond tochange. We investigated the direct and interactive effects of[CO₂], warming and altered soil moisture in open-top chambers(OTCs) enclosing a constructed old-field community to test howthese factors shape plant communities. Materials and methods: The experimental facility in Oak Ridge, TN, USA, made use of4-m diameter OTCs and rain shelters to manipulate [CO₂] (ambient,ambient + 300 ppm), air temperature (ambient, ambient + 3.5°C)and soil moisture (wet, dry). The plant communities within thechambers comprised seven common old-field species, includinggrasses, forbs and legumes. We tracked foliar cover for eachspecies and calculated community richness, evenness and diversityfrom 2003 to 2005. Important findings: This work resulted in three main findings: (1) warming had species-specificeffects on foliar cover that varied through time and were alteredby soil moisture treatments; (2) [CO₂] had little effect onindividual species or the community; (3) diversity, evennessand richness were influenced most by soil moisture, primarilyreflecting the response of one dominant species. We concludethat individualistic species responses to atmospheric and climaticchange can alter community composition and that plant populationsand communities should be considered as part of analyses ofterrestrial ecosystem response to climate change. However, predictionof plant community responses may be difficult given interactionsbetween factors and changes in response through time.     

Global change belowground: impacts of elevated CO2, nitrogen,and summer drought on soil food webs and biodiversity

The world's ecosystems are subjected to various anthropogenic global change agents, such as enrichment of atmospheric CO₂ concentrations, nitrogen (N) deposition, and changes in precipitation regimes. Despite the increasing appreciation that the consequences of impending global change can be better understood if varying agents are studied in concert, there is a paucity of multi‐factor long‐term studies, particularly on belowground processes. Herein, we address this gap by examining the responses of soil food webs and biodiversity to enrichment of CO₂, elevated N, and summer drought in a long‐term grassland study at Cedar Creek, Minnesota, USA (BioCON experiment). We use structural equation modeling (SEM), various abiotic and biotic explanatory variables, and data on soil microorganisms, protozoa, nematodes, and soil microarthropods to identify the impacts of multiple global change effects on drivers belowground. We found that long‐term (13‐year) changes in CO₂ and N availability resulted in modest alterations of soil biotic food webs and biodiversity via several mechanisms, encompassing soil water availability, plant productivity, and – most importantly – changes in rhizodeposition. Four years of manipulation of summer drought exerted surprisingly minor effects, only detrimentally affecting belowground herbivores and ciliate protists at elevated N. Elevated CO₂ increased microbial biomass and the density of ciliates, microarthropod detritivores, and gamasid mites, most likely by fueling soil food webs with labile C. Moreover, beneficial bottom‐up effects of elevated CO₂ compensated for detrimental elevated N effects on soil microarthropod taxa richness. In contrast, nematode taxa richness was lowest at elevated CO₂ and elevated N. Thus, enrichment of atmospheric CO₂ concentrations and N deposition may result in taxonomically and functionally altered, potentially simplified, soil communities. Detrimental effects of N deposition on soil biodiversity underscore recent reports on plant community simplification. This is of particular concern, as soils house a considerable fraction of global biodiversity and ecosystem functions.     

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.     

Mycorrhizal dynamics under elevated CO2 and nitrogen fertilization in a warm temperate forest

We examined the response of mycorrhizal fungi to free-air CO₂ enrichment (FACE) and nitrogen (N) fertilization in a warm temperate forest to better understand potential influences over plant nutrient uptake and soil carbon (C) storage. In particular, we hypothesized that mycorrhizal fungi and glomalin would become more prevalent under elevated CO₂ but decrease under N fertilization. In addition, we predicted that N fertilization would mitigate any positive effects of elevated CO₂ on mycorrhizal abundance. Overall, we observed a 14% increase in ectomycorrhizal (ECM) root colonization under CO₂ enrichment, which implies that elevated CO₂ results in greater C investments in these fungi. Arbuscular mycorrhizal (AM) hyphal length and glomalin stocks did not respond substantially to CO₂ enrichment, and effects of CO₂ on AM root colonization varied by date. Nitrogen effects on AM fungi were not consistent with our hypothesis, as we found an increase in AM colonization under N fertilization. Lastly, neither glomalin concentrations nor ECM colonization responded significantly to N fertilization or to an N-by-CO₂ interaction. A longer duration of N fertilization may be required to detect effects on these parameters.     

Plant and microbial N acquisition under elevated atmospheric CO2 in two mesocosm experiments with annual grasses

The impact of elevated CO₂ on terrestrial ecosystem C balance, both in sign or magnitude, is not clear because the resulting alterations in C input, plant nutrient demand and water use efficiency often have contrasting impacts on microbial decomposition processes. One major source of uncertainty stems from the impact of elevated CO₂ on N availability to plants and microbes. We examined the effects of atmospheric CO₂ enrichment (ambient+370 μmol mol^?1) on plant and microbial N acquisition in two different mesocosm experiments, using model plant species of annual grasses of Avena barbata and A. fatua, respectively. The A. barbata experiment was conducted in a N‐poor sandy loam and the A. fatua experiment was on a N‐rich clayey loam. Plant–microbial N partitioning was examined through determining the distribution of a ¹⁵N tracer. In the A. barbata experiment, ¹⁵N tracer was introduced to a field labeling experiment in the previous year so that ¹⁵N predominantly existed in nonextractable soil pools. In the A. fatua experiment, ¹⁵N was introduced in a mineral solution [(¹⁵NH₄)₂SO₄ solution] during the growing season of A. fatua. Results of both N budget and ¹⁵N tracer analyses indicated that elevated CO₂ increased plant N acquisition from the soil. In the A. barbata experiment, elevated CO₂ increased plant biomass N by ca. 10% but there was no corresponding decrease in soil extractable N, suggesting that plants might have obtained N from the nonextractable organic N pool because of enhanced microbial activity. In the A. fatua experiment, however, the CO₂‐led increase in plant biomass N was statistically equal to the reduction in soil extractable N. Although atmospheric CO₂ enrichment enhanced microbial biomass C under A. barbata or microbial activity (respiration) under A. fatua, it had no significant effect on microbial biomass N in either experiment. Elevated CO₂ increased the colonization of A. fatua roots by arbuscular mycorrhizal fungi, which coincided with the enhancement of plant competitiveness for soluble soil N. Together, these results suggest that elevated CO₂ may tighten N cycling through facilitating plant N acquisition. However, it is unknown to what degree results from these short‐term microcosm experiments can be extrapolated to field conditions. Long‐term studies in less‐disturbed soils are needed to determine whether CO₂‐enhancement of plant N acquisition can significantly relieve N limitation over plant growth in an elevated CO₂ environment.     

Plant species-specific changes in root-inhabiting fungi in a California annual grassland: responses to elevated CO2 and nutrients

Five co-occurring plant species from an annual mediterranean grassland were grown in monoculture for 4 months in pots inside open-top chambers at the Jasper Ridge Biological Preserve (San Mateo County, California). The plants were exposed to elevated atmospheric CO₂ and soil nutrient enrichment in a complete factorial experiment. The response of root-inhabiting non-mycorrhizal and arbuscular mycorrhizal fungi to the altered resource base depended strongly on the plant species. Elevated CO₂ and fertilization altered the ratio of non-mycorrhizal to mycorrhizal fungal colonization for some plant species, but not for others. Percent root infection by non-mycorrhizal fungi increased by over 500% for Linanthus parviflorus in elevated CO₂, but decreased by over 80% for Bromus hordeaceus. By contrast, the mean percent infection by mycorrhizal fungi increased in response to elevated CO₂ for all species, but significantly only for Avena barbata and B. hordeaceus. Percent infection by mycorrhizal fungi increased, decreased, or remained unchanged for different plant hosts in response to fertilization. There was evidence of a strong interaction between the two treatments for some plant species and non-mycorrhizal and mycorrhizal fungi. This study demonstrated plant species- and soil fertility-dependent shifts in below-ground plant resource allocation to different morpho-groups of fungal symbionts. This may have consequences for plant community responses to elevated CO₂ in this California grassland ecosystem. Received: 2 June 1997 / Accepted: 22 August 1997     

Microbial assimilation of new photosynthate is altered by plant species richness and nitrogen deposition

To determine how plant species richness impacts microbial assimilation of new photosynthate, and how this may be modified by atmospheric N deposition, we analyzed the microbial assimilation of recent photosynthate in a 6-year-long field experiment in which plant species richness, atmospheric N deposition, and atmospheric CO₂ concentration were manipulated in concert. The depleted δ¹³C of fumigation CO₂ enabled us to investigate the effect of plant species richness and atmospheric N deposition on the metabolism of soil microbial communities in the elevated CO₂ treatment. To accomplish this, we determined the δ¹³C of bacterial, actinobacterial, and fungal phospholipid fatty acids (PLFAs). In the elevated CO₂ conditions of this study, the δ¹³C of bacterial PLFAs (i15:0, i16:0, 16:1ω7c, 16:1ω9c, 10Me16:0, and 10Me18:0) and the fungal PLFA 18:1ω9c was significantly lower in species-rich plant communities than in species-poor plant communities, indicating that microbial incorporation of new C increased with plant species richness. Despite an increase in plant production, total PLFA decreased under N deposition. Moreover, N deposition also decreased fungal relative abundance in species-rich plant communities. In our study, plant species richness directly increased microbial incorporation of new photosynthate, providing a mechanistic link between greater plant detritus production in species-rich plant communities and larger and more active soil microbial community.