Global Change Factors Research Articles

Plant diversity drives soil microbial biomass carbon in grasslands irrespective of global environmental change factors.

Soil microbial biomass is a key determinant of carbon dynamics in the soil. Several studies have shown that soil microbial biomass significantly increases with plant species diversity, but it remains unclear whether plant species diversity can also stabilize soil microbial biomass in a changing environment. This question is particularly relevant as many global environmental change (GEC) factors, such as drought and nutrient enrichment, have been shown to reduce soil microbial biomass. Experiments with orthogonal manipulations of plant diversity and GEC factors can provide insights whether plant diversity can attenuate such detrimental effects on soil microbial biomass. Here, we present the analysis of 12 different studies with 14 unique orthogonal plant diversity×GEC manipulations in grasslands, where plant diversity and at least one GEC factor (elevated CO2 , nutrient enrichment, drought, earthworm presence, or warming) were manipulated. Our results show that higher plant diversity significantly enhances soil microbial biomass with the strongest effects in long-term field experiments. In contrast, GEC factors had inconsistent effects with only drought having a significant negative effect. Importantly, we report consistent non-significant effects for all 14 interactions between plant diversity and GEC factors, which indicates a limited potential of plant diversity to attenuate the effects of GEC factors on soil microbial biomass. We highlight that plant diversity is a major determinant of soil microbial biomass in experimental grasslands that can influence soil carbon dynamics irrespective of GEC.

Root and leaf traits reflect distinct resource acquisition strategies in tropical lianas and trees.

In Neotropical forests, lianas are increasing in abundance relative to trees. This increased species richness may reflect a positive response to global change factors including increased temperature, atmospheric CO2, habitat fragmentation, and drought severity; however, questions remain as to the specific mechanisms facilitating the response. Previous work suggests that lianas may gain an ecological advantage over trees through leaf functional traits that offer a quick return on investment of resources, although it is unknown whether this pattern extends to root traits and relationships with fungal or bacterial symbionts belowground. We sampled confamilial pairs of liana and tree species and quantified morphological and chemical traits of leaves and fine roots, as well as root symbiont abundance, to determine whether functional traits associated with resource acquisition differed between the two. Compared to trees, lianas possessed higher specific leaf area, specific root length, root branching intensity, and root nitrogen (N) and phosphorus (P) concentrations, and lower leaf and root tissue density, leaf and root carbon (C), root diameter, root C:P and N:P, and mycorrhizal colonization. Our study provides new evidence that liana leaf and root traits are characterized by a rapid resource acquisition strategy relative to trees. These liana functional traits may facilitate their response to global change, raising questions about how increased liana dominance might affect ecosystem processes of Neotropical forests.

Increased rainfall variability and N addition accelerate litter decomposition in a restored prairie.

Anthropogenic nitrogen deposition and projected increases in rainfall variability (the frequency of drought and heavy rainfall events) are expected to strongly influence ecosystem processes such as litter decomposition. However, how these two global change factors interact to influence litter decomposition is largely unknown. I examined how increased rainfall variability and nitrogen addition affected mass and nitrogen loss of litter from two tallgrass prairie species, Schizachyrium scoparium and Solidago canadensis, and isolated the effects of each during plant growth and during litter decomposition. I increased rainfall variability by consolidating ambient rainfall into larger events and simulated chronic nitrogen deposition using a slow-release urea fertilizer. S. scoparium litter decay was more strongly regulated by the treatments applied during plant growth than by those applied during decomposition. During plant growth, increased rainfall variability resulted in S. scoparium litter that subsequently decomposed more slowly and immobilized more nitrogen than litter grown under ambient conditions, whereas nitrogen addition during plant growth accelerated subsequent mass loss of S. scoparium litter. In contrast, S. canadensis litter mass and N losses were enhanced under either N addition or increased rainfall variability both during plant growth and during decomposition. These results suggest that ongoing changes in rainfall variability and nitrogen availability are accelerating nutrient cycling in tallgrass prairies through their combined effects on litter quality, environmental conditions, and plant community composition.

What have we learned from global change manipulative experiments in China? A meta-analysis.

Although China has the largest population in the world, a faster rate of warming than the global average, and an active global change research program, results from many of the global change experiments in Chinese terrestrial ecosystems have not been included in global syntheses. Here, we specifically analyze the observed responses of carbon (C) and nitrogen (N) cycling in global change manipulative experiments in China, and compare these responses to those from other regions of the world. Most global change factors, vegetation types, and treatment methods that have been studied or used elsewhere in the world have also been studied and applied in China. The responses of terrestrial ecosystem C and N cycles to N addition and climate warming in China are similar in both direction and intensity to those reported in global syntheses. In Chinese ecosystems as elsewhere, N addition significantly increased aboveground (AGB) and belowground biomass (BGB), litter mass, dissolved organic C, net ecosystem productivity (NEP), and gross ecosystem productivity (GEP). Warming stimulated AGB, BGB and the root-shoot ratio. Increasing precipitation accelerated GEP, NEP, microbial respiration, soil respiration, and ecosystem respiration. Our findings complement and support previous global syntheses and provide insight into regional responses to global change.

The functional quality of decomposing litter outputs from an Arctic plant community is affected by long-term exposure to enhanced UV-B

Plant exposure to enhanced UV-B radiation typically induces changes in leaf secondary metabolite profiles which will be inherited in litter, affecting litter breakdown and the carbon (C) dynamics of sensitive plant communities. A key enzyme in the decomposition process is phenol oxidase which is influenced by litter quality and, hence, a decomposition bioindicator. Here we investigated dwarf shrub litter decomposition following experimental community exposure to enhanced UV-B over two decades in the Swedish sub-Arctic. We examined the hypothesis that foliar UV-B exposure would alter litter quality to elevate phenol oxidase activity. This was tested in the field by measuring phenol oxidase activity in freshly collected mixed-community litter from under our experimental vegetation. A laboratory mesocosm was next used in a decomposition assay to investigate individual species responses over eight weeks, with an emphasis on the quality of leachate outputs from decomposing litter (from Empetrum hermaphroditum, Vaccinium vitis-idaea, Vaccinium uliginosum). In the assay bi-weekly collections of leachate were analysed for phenol oxidase activity, together with total phenolics and dissolved organic C (DOC). At the end of the assay litter mass loss and respired C were also determined. The initial assessment on field mixed-community litter found no enhanced UV-B treatment (henceforth: ‘UV-B treatment’) effect on phenol oxidase activity. However, in the controlled laboratory mesocosm assay, significant species-specific effects of the UV-B treatment were evident, with increased phenol oxidase activity in V. vitis-idaea leachate (P<0.001) and a significant reduction (P=0.05) in respired C. Leachate DOC release from the UV-B treatment was greater in both Vaccinium species and the effect on V. uliginosum was significant (P<0.05). The UV-B treatment had no effect on the total phenolic concentration of litter or leachates for any species, but there were significant differences in leachate total phenolics, both over time and between species. Also the initial phenolic concentration in leachates from the decomposing litter of E. hermaphroditum was greater than both Vaccinum species. Results suggest a species specific role for UV-B in influencing enzyme function and decomposition, dependent on individual traits. This has implications for decomposition dynamics in this system and more widely. Our study highlights the value of using a laboratory assay to gain a mechanistic understanding the species level impacts of a global change factor (UV-B) on decomposition, which are otherwise obscured by community-level responses and difficult to determine under field conditions.

Scale dependence of disease impacts on quaking aspen (Populus tremuloides) mortality in the southwestern United States.

Depending on how disease impacts tree exposure to risk, both the prevalence of disease and disease effects on survival may contribute to patterns of mortality risk across a species' range. Disease may accelerate tree species' declines in response to global change factors, such as drought, biotic interactions, such as competition, or functional traits, such as allometry. To assess the role of disease in mediating mortality risk in quaking aspen (Populus tremuloides), we developed hierarchical Bayesian models for both disease prevalence in live aspen stems and the resulting survival rates of healthy and diseased aspen near the species' southern range limit using 5088 individual trees on 281 United States Forest Service Forest Inventory and Analysis plots in the southwestern United States. We found that disease prevalence depended primarily on tree size, tree allometry, and spatial variation in precipitation, while mortality depended on tree size, allometry, competition, spatial variation in summer temperature, and both temporal and spatial variation in summer precipitation. Disease prevalence was highest in large trees with low slenderness found on dry sites. For healthy trees, mortality decreased with diameter, slenderness, and temporal variation in summer precipitation, but increased with competition and spatial variation in summer temperature. Mortality of diseased trees decreased with diameter and aspen relative basal area and increased with mean summer temperature and precipitation. Disease infection increased aspen mortality, especially in trees of intermediate size and trees on plots at climatic extremes (i.e., cool, wet and warm, dry climates). By examining variation in disease prevalence, mortality of healthy trees, and mortality of diseased trees, we showed that the role of disease in aspen tree mortality depended on the scale of inference. For variation among individuals in diameter, disease tended to expose intermediate-size trees experiencing moderate risk to greater risk. For spatial variation in summer temperature, disease exposed lower risk populations to greater mortality probabilities, but the magnitude of this exposure depended on summer precipitation. Furthermore, the importance of diameter and slenderness in mediating responses to climate supports the increasing emphasis on trait variation in studies of ecological responses to global change.

Impacts of urban sprawl on species richness of plants, butterflies, gastropods and birds: not only built-up area matters

Urban growth is a major factor of global environmental change and has important impacts on biodiversity, such as changes in species composition and biotic homogenization. Most previous studies have focused on effects of urban area as a general measure of urbanization, and on few or single taxa. Here, we analyzed the impacts of the different components of urban sprawl (i.e., scattered and widespread urban growth) on species richness of a variety of taxonomic groups covering mosses, vascular plants, gastropods, butterflies, and birds at the habitat and landscape scales. Besides urban area, we considered the average age, imperviousness, and dispersion degree of urban area, along with human population density, to disentangle the effects of the different components of urban sprawl on biodiversity. The study was carried out in the Swiss Plateau that has undergone substantial urban sprawl in recent decades. Vascular plants and birds showed the strongest responses to urban sprawl, especially at the landscape scale, with non-native and ruderal plants proliferating and common generalist birds increasing at the expense of specialist birds as urban sprawl grew. Overall, urban area had the greatest contribution on such impacts, but additional effects of urban dispersion (i.e., increase of non-native plants) and human population density (i.e., increases of ruderal plants and common generalist birds) were found. Our findings support the hypothesis that negative impacts of urban sprawl on biodiversity can be reduced by compacting urban growth while still avoiding the formation of very densely populated areas.

Biotic interactions mediate soil microbial feedbacks to climate change.

Decomposition of organic material by soil microbes generates an annual global release of 50-75 Pg carbon to the atmosphere, ∼7.5-9 times that of anthropogenic emissions worldwide. This process is sensitive to global change factors, which can drive carbon cycle-climate feedbacks with the potential to enhance atmospheric warming. Although the effects of interacting global change factors on soil microbial activity have been a widespread ecological focus, the regulatory effects of interspecific interactions are rarely considered in climate feedback studies. We explore the potential of soil animals to mediate microbial responses to warming and nitrogen enrichment within a long-term, field-based global change study. The combination of global change factors alleviated the bottom-up limitations on fungal growth, stimulating enzyme production and decomposition rates in the absence of soil animals. However, increased fungal biomass also stimulated consumption rates by soil invertebrates, restoring microbial process rates to levels observed under ambient conditions. Our results support the contemporary theory that top-down control in soil food webs is apparent only in the absence of bottom-up limitation. As such, when global change factors alleviate the bottom-up limitations on microbial activity, top-down control becomes an increasingly important regulatory force with the capacity to dampen the strength of positive carbon cycle-climate feedbacks.

Microbial lipid and amino sugar responses to long-term simulated global environmental changes in a California annual grassland.

Global environmental change is predicted to have major consequences for carbon cycling and the functioning of soil ecosystems. However, we have limited knowledge about its impacts on the microorganisms, which act as a “valve” between carbon sequestered in soils versus released into the atmosphere. In this study we examined microbial response to continuous 9-years manipulation of three global change factors (elevated CO2, warming, and nitrogen deposition), singly and in combination using two methods: lipid and amino sugar biomarkers at the Jasper Ridge Global Change Experiment (JRGCE). The two methods yielded important distinctions. There were limited microbial lipid differences, but many significant effects for microbial amino sugars. We found that CO2 was not a direct factor influencing soil carbon and major amino sugar pools, but had a positive impact on bacterial-derived muramic acid. Likewise, warming and nitrogen deposition appeared to enrich residues specific to bacteria despite an overall depletion in total amino sugars. The results indicate that elevated CO2, warming, and nitrogen deposition all appeared to increase bacterial-derived residues, but this accumulation effect was far offset by a corresponding decline in fungal residues. The sensitivity of microbial residue biomarker amino sugars to warming and nitrogen deposition may have implications for our predictions of global change impacts on soil stored carbon.

Anthropogenic nitrogen deposition enhances carbon sequestration in boreal soils.

It is proposed that carbon (C) sequestration in response to reactive nitrogen (Nr ) deposition in boreal forests accounts for a large portion of the terrestrial sink for anthropogenic CO2 emissions. While studies have helped clarify the magnitude by which Nr deposition enhances C sequestration by forest vegetation, there remains a paucity of long-term experimental studies evaluating how soil C pools respond. We conducted a long-term experiment, maintained since 1996, consisting of three N addition levels (0, 12.5, and 50 kg N ha(-1) yr(-1) ) in the boreal zone of northern Sweden to understand how atmospheric Nr deposition affects soil C accumulation, soil microbial communities, and soil respiration. We hypothesized that soil C sequestration will increase, and soil microbial biomass and soil respiration will decrease, with disproportionately large changes expected compared to low levels of N addition. Our data showed that the low N addition treatment caused a non-significant increase in the organic horizon C pool of ~15% and a significant increase of ~30% in response to the high N treatment relative to the control. The relationship between C sequestration and N addition in the organic horizon was linear, with a slope of 10 kg C kg(-1) N. We also found a concomitant decrease in total microbial and fungal biomasses and a ~11% reduction in soil respiration in response to the high N treatment. Our data complement previous data from the same study system describing aboveground C sequestration, indicating a total ecosystem sequestration rate of 26 kg C kg(-1) N. These estimates are far lower than suggested by some previous modeling studies, and thus will help improve and validate current modeling efforts aimed at separating the effect of multiple global change factors on the C balance of the boreal region.

Fungal communities respond to long-term CO2 elevation by community reassembly.

Fungal communities play a major role as decomposers in the Earth's ecosystems. Their community-level responses to elevated CO2 (eCO2), one of the major global change factors impacting ecosystems, are not well understood. Using 28S rRNA gene amplicon sequencing and co-occurrence ecological network approaches, we analyzed the response of soil fungal communities in the BioCON (biodiversity, CO2, and N deposition) experimental site in Minnesota, USA, in which a grassland ecosystem has been exposed to eCO2 for 12 years. Long-term eCO2 did not significantly change the overall fungal community structure and species richness, but significantly increased community evenness and diversity. The relative abundances of 119 operational taxonomic units (OTU; ∼27% of the total captured sequences) were changed significantly. Significantly changed OTU under eCO2 were associated with decreased overall relative abundance of Ascomycota, but increased relative abundance of Basidiomycota. Co-occurrence ecological network analysis indicated that eCO2 increased fungal community network complexity, as evidenced by higher intermodular and intramodular connectivity and shorter geodesic distance. In contrast, decreased connections for dominant fungal species were observed in the eCO2 network. Community reassembly of unrelated fungal species into highly connected dense modules was observed. Such changes in the co-occurrence network topology were significantly associated with altered soil and plant properties under eCO2, especially with increased plant biomass and NH4 (+) availability. This study provided novel insights into how eCO2 shapes soil fungal communities in grassland ecosystems.

Erosion of beta diversity under interacting global change impacts in a semi‐arid grassland

Summary Human impacts are often thought of as homogenizing natural communities, but it is unclear how these impacts interact to alter the beta diversity (spatial variability) of plant communities. In a grassland with high beta diversity along a soil fertility gradient, we asked which combinations of nutrient enrichment, precipitation enhancement and disturbance would homogenize communities along the gradient by allowing dominant species from one part of the gradient to increase their abundances in other parts of the gradient. In particular, joint addition of nutrients and water might especially benefit resource‐acquisitive and/or exotic species, which have been observed to compete well in resource‐rich conditions; this would tend to reduce variation in species composition across the gradient. To alleviate dispersal limitation as a constraint on treatment responses, we added seeds of 15 species abundant in different parts of the soil fertility gradient to our 132 plots. Fertilization and water addition allowed species from fertile soils to increase their abundance in infertile soils, an effect that strengthened with time. Disturbance permitted species from infertile soils to increase in abundance in infertile soils, but only transiently. Plant height, specific leaf area and exotic status best predicted the abilities of species to attain greater cover in harsh habitats under ameliorated conditions. Among‐habitat beta diversity of seeded species was reduced by the interactive effect of fertilization and watering, although not by watering alone. Synthesis. Our findings are a novel demonstration of how interacting global change factors reduce beta diversity by eroding the biotic and abiotic resistances that control community structure along soil fertility gradients.

The role and importance of international organizations in maintaining ecological and human security

This paper presents key reflections on compromised ecological and human security. Man's growing insecurity has been affected by global ecological issues in two ways, directly and indirectly. The former involves health implications of exposure to ultraviolet radiation, while the latter is linked to potential climate changes and malnutrition. Not all ecological hazards are global, but it is clear that they exacerbate local pollution-related concerns and that individual countries are not capable of addressing them on their own. How the perspective of human security can be incorporated into the analysis of environmental challenges and how the ecological dimension can be bolstered in the analysis of human security are some of the other issues examined in this paper. The paper aims to recognize the role and importance of international organizations based on conceptual, analytical and operational considerations of ecological and human security issues. Namely, through mutual cooperation and joint activities these organizations are making great efforts to produce results by analyzing the interaction between factors of global change and degradation of the environment and factors affecting human security. Among the more important organizations mentioned are the UN and several of its agencies and suborganizations that are part of the 'UN family' (UNDP, UNESCO, UNU, UNU-EHS, UNEP, FAO) as well as a number of regional organizations such as OSCE and EU.

Introduction to the Special Issue: Across the horizon: scale effects in global change research.

As a result of the increasing speed and magnitude in which habitats worldwide are experiencing environmental change, making accurate predictions of the effects of global change on ecosystems and the organisms that inhabit them have become an important goal for ecologists. Experimental and modelling approaches aimed at understanding the linkages between factors of global change and biotic responses have become numerous and increasingly complex in order to adequately capture the multifarious dynamics associated with these relationships. However, constrained by resources, experiments are often conducted at small spatiotemporal scales (e.g. looking at a plot of a few square metres over a few years) and at low organizational levels (looking at organisms rather than ecosystems) in spite of both theoretical and experimental work that suggests ecological dynamics across scales can be dissimilar. This phenomenon has been hypothesized to occur because the mechanisms that drive dynamics across scales differ. A good example is the effect of elevated CO2 on transpiration. While at the leaf level, transpiration can be reduced, at the stand level, transpiration can increase because leaf area per unit ground area increases. The reported net effect is then highly dependent on the spatiotemporal scale. This special issue considers the biological relevancy inherent in the patterns associated with the magnitude and type of response to changing environmental conditions, across scales. This collection of papers attempts to provide a comprehensive treatment of this phenomenon in order to help develop an understanding of the extent of, and mechanisms involved with, ecological response to global change.

Linking Azolla filiculoides invasion to increased winter temperatures in the Doñana marshland (SW Spain)

Unravelling how a multiplicity of global change factors might influence the expansion of alien plants is a major goal of invasion biology. We explored the association of climatic conditions (i.e. annual rainfall and average minimum temperature during the coldest months) and water quality (orthophosphate, nitrate concentrations and electrical conductivity), with blooms of the invasive fern Azolla filiculoides Lam. in the Donana marshland (SW Spain), one of the most extensive wetlands in Europe. Azolla was first detected in Donana in 2001. Its annual cover has been quantified since then through remote sensing. In the last decade there has been a considerable increase of orthophosphate and yearly fluctuations of nitrate. The first orthophosphate increase coincided with the presence of Azolla in Donana. Since then, there has been a positive relationship between inter-annual variation in Azolla cover and minimum temperature during the coldest months. Our study shows that the Donana marshland is facing rapid eutrophication and invasion by Azolla, most notably in years with high temperatures during the winter.

Nematode functional guilds, not trophic groups, reflect shifts in soil food webs and processes in response to interacting global change factors

a b s t r a c t Soils store ∼80% of global terrestrial organic carbon and alterations in fluxes into and out of this pool may interact with ongoing climate change. Belowground food webs drive soil C dynamics, but little is known about their responses to co-occurring global change agents. We investigated open-air experi- mental grassland communities at ambient and elevated atmospheric CO2 concentration, ambient and enriched nitrogen input, and ambient and reduced summer precipitation to evaluate how these agents interactively affect soil nematodes, which are often used as an indicator group for soil food web structure and soil health. The aim of the study was to elucidate the response of the functional diversity of soil nematodes to changing environmental conditions by using nematode functional guilds and indices as indicators. The results suggest that nematode functional guilds surpass nematode trophic groups as soil indicators, suggesting that more detailed data on nematode community structure is essential to capture functional changes in response to environmental change. For instance, the density of opportunistic fungal feeders increased due to N addition with the response being more pronounced at elevated CO2, whereas densities of sensitive fungal-feeders were increased at ambient N and elevated CO2, illustrating opposing responses within one trophic group. Opportunistic bacterial feeders increased at elevated N, but did not respond to other environmental factors studied. Root-feeding Longidoridae were significantly reduced at elevated CO2 and elevated N compared to ambient conditions, whereas other plant feeders were little affected by the manipulations. Predacious nematodes were less abundant at elevated N, and the Structure Index (which indicates food web structure) suggested reduced top-down forces and simplified soil food webs, although omnivores did not vary significantly. Elevated CO2 buffered the effect of reduced precipitation on the Enrichment Index (which indicates increased resource availability) and the Channel Index (which indicates changes in decomposition channel) probably due to reduced stomatal conductance at elevated CO2. Further, the results suggest that the decomposer community switched from a bacterial-dominated to a fungal-dominated system at elevated N, indicating shifts in the microbial community as well as in the functioning of belowground food webs. Overall, the studied global change agents interactively and differentially affected functional guilds of soil nematodes, suggesting complex changes in soil processes. We highlight that detailed information on the functional guilds of nematodes is likely necessary to fully understand alterations in soil food webs and related processes due to global environmental change.

Direct and indirect drivers of plant diversity responses to climate and clipping across northern temperate grassland

It is well known that climate can influence plant community assembly via a multitude of indirect and direct pathways. However, interpretations of plant diversity responses to simulated climate change experiments, and subsequent predictions of plant communities under future climate scenarios, rarely address the importance of indirect effects. Networks of direct and indirect effects are also critical in understanding linkages between climate and grazing, a common land use of grasslands, and implications for plant diversity. We characterized the roles of indirect vs. direct effects in determining plant diversity responses to climate and grazing using data from three northern temperate grasslands in which we conducted factorial experiments manipulating precipitation, air temperature, and clipping intensity. Utilizing a structural equation modeling framework to address the multivariate networks, we found warming operated directly, causing species loss at all sites. We identified shoot biomass as the key indirect driver of diversity loss in response to both precipitation and clipping, regardless of site. However, site‐specific contingencies in the network of interactions were important for understanding varied precipitation effects. At the driest site only, shoot biomass was resistant to reduced precipitation, and diversity was consequently unaltered. Similarly, disconnect between primary drivers and responses explained relatively idiosyncratic responses of evenness compared to richness. Importantly, the finding of widespread, directly controlled plant diversity loss with warming aligns with concerns about declining biodiversity under climate change. However, using a framework of network interactions also allowed us to pinpoint the source of variability in response across systems. Looking forward, we can use the identification of this key indirect pathway to guide an understanding of community assembly under factors likely to control shoot biomass. Viewing a multifactorial, multisite experimental approach through a framework of network interactions allowed us to both identify generalized responses and distill the complexity of contingent responses. This, along with the practical need to identify diversity responses to climate change and grazing, underscores the importance of understanding both indirect and direct drivers of ecosystem responses to global change factors.

Successional processes and global change: tree seedling establishment in response to warming and N addition in a temperate old field

Climate change and increased atmospheric nitrogen (N) deposition can alter successional processes, either by directly altering the establishment and growth of later successional species or by altering their interactions with earlier successional species. We examined the effects of these global change factors on post-germination tree establishment in a grass-dominated old field in London, Ontario, Canada, by transplanting tree seedlings into the plots of a warming and N addition field experiment. Transplantation was duplicated over two growing seasons. We also administered warming and N addition treatments to tree seedlings in the absence of grasses in a greenhouse in order to examine the direct seedling responses. In the field, seedling survival was generally low over the first growing season, which was very dry. The only significant treatment effects were those of warming decreasing the survival and biomass of Malus coronaria, and N addition increasing the survival of Elaeagnus umbellata. Grass biomass increased in response to N addition, and seedling condition and survival were positively correlated with increased grass biomass for most species. Seedling survival was much higher in the second growing season, although warming decreased the survival of Rhamnus cathartica and Rhus typhina. N addition increased the biomass and leaf area of M. coronaria, but decreased these responses for R. typhina, and warming increased the leaf area of Fraxinus americana. Grass biomass again increased in response to N addition, but unlike the previous year, seedling condition and survival were generally negatively correlated with increased grass biomass. In the greenhouse (grasses absent), the seedlings generally increased in biomass in response to N addition and were unresponsive to warming. Our results indicate that both warming and N addition can affect the processes determining post-germination tree seedling establishment, and that depending on the environmental conditions, and in particular water availability, the responses of grasses appear to either facilitate or suppress seedling condition and survival.

Litters of photosynthetically divergent grasses exhibit differential metabolic responses to warming and elevated CO2

Climatic stress such as warming would alter physiological pathways in plants leading to changes in tissue chemistry. Elevated CO2 could partly mitigate warming induced moisture stress, and the degree of this mitigation may vary with plant functional types. We studied the composition of structural and non‐structural metabolites in senesced tissues of Bouteloua gracilis (C4) and Pascopyrum smithii (C3) at the Prairie Heating and CO2 Enrichment experiment, Wyoming, USA. We hypothesized that P. smithii and B. gracilis would respond to unfavorable global change factors by producing structural metabolites and osmoregulatory compounds that are necessary to combat stress. However, due to the inherent variation in the tolerance of their photosynthetic pathways to warming and CO2, we hypothesized that these species will exhibit differential response under different combinations of warming and CO2 conditions. Due to a lower thermo‐tolerance of the C4 photosynthesis we expected B. gracilis to exhibit a greater metabolic response under warming with ambient CO2 (cT) and P. smithii to exhibit a similar response under warming combined with elevated CO2 (CT). Our hypothesis was supported by the differential response of structural compounds in these two species, where cT increased the content of lignin and cuticular‐matrix in B. gracilis. In P. smithii a similar response was observed in plants exposed to CT, possibly due to the partial alleviation of moisture stress. With warming, the total cell‐wall bound phenolic acids that cross link polysaccharides to lignins increased in B. gracilis and decreased in P. smithii, indicating a potentially adaptive response of C4 pathway to warming alone. Similarly, in B. gracilis, extractable polar metabolites such as sugars and phenolic acids increased with the main effect of warming. Conversely, in P. smithii, only sugars showed a higher abundance in plants exposed to warming treatments indicating that warming alone might be metabolically too disruptive for the C3 photosynthetic pathway. Here we show for the first time, that along with traditionally probed extractable metabolites, warming and elevated CO2 differentially influence the structural metabolites in litters of photosynthetically divergent grass species. If these unique metabolite responses occur in other species of similar functional types, this could potentially alter carbon cycling in grasslands due to the varying degradability of these litters.

Using root form to improve our understanding of root function.

Using root form to improve our understanding of root function.