In recent years, increased awareness of the potential interactions between rising atmospheric
CO2
concentrations (
CO2
) and temperature has illustrated the importance of multifactorial ecosystem ...manipulation experiments for validating Earth System models. To address the urgent need for increased understanding of responses in multifactorial experiments, this article synthesizes how ecosystem productivity and soil processes respond to combined warming and
CO2
manipulation, and compares it with those obtained in single factor
CO2
and temperature manipulation experiments. Across all combined elevated
CO2
and warming experiments, biomass production and soil respiration were typically enhanced. Responses to the combined treatment were more similar to those in the
CO2
‐only treatment than to those in the warming‐only treatment. In contrast to warming‐only experiments, both the combined and the
CO2
‐only treatments elicited larger stimulation of fine root biomass than of aboveground biomass, consistently stimulated soil respiration, and decreased foliar nitrogen (N) concentration. Nonetheless, mineral N availability declined less in the combined treatment than in the
CO2
‐only treatment, possibly due to the warming‐induced acceleration of decomposition, implying that progressive nitrogen limitation (PNL) may not occur as commonly as anticipated from single factor
CO2
treatment studies. Responses of total plant biomass, especially of aboveground biomass, revealed antagonistic interactions between elevated
CO2
and warming, i.e. the response to the combined treatment was usually less‐than‐additive. This implies that productivity projections might be overestimated when models are parameterized based on single factor responses. Our results highlight the need for more (and especially more long‐term) multifactor manipulation experiments. Because single factor
CO2
responses often dominated over warming responses in the combined treatments, our results also suggest that projected responses to future global warming in Earth System models should not be parameterized using single factor warming experiments.
Changes in rainfall amounts and patterns have been observed and are expected to continue in the near future with potentially significant ecological and societal consequences. Modelling vegetation ...responses to changes in rainfall is thus crucial to project water and carbon cycles in the future. In this study, we present the results of a new model‐data intercomparison project, where we tested the ability of 10 terrestrial biosphere models to reproduce the observed sensitivity of ecosystem productivity to rainfall changes at 10 sites across the globe, in nine of which, rainfall exclusion and/or irrigation experiments had been performed. The key results are as follows: (a) Inter‐model variation is generally large and model agreement varies with timescales. In severely water‐limited sites, models only agree on the interannual variability of evapotranspiration and to a smaller extent on gross primary productivity. In more mesic sites, model agreement for both water and carbon fluxes is typically higher on fine (daily–monthly) timescales and reduces on longer (seasonal–annual) scales. (b) Models on average overestimate the relationship between ecosystem productivity and mean rainfall amounts across sites (in space) and have a low capacity in reproducing the temporal (interannual) sensitivity of vegetation productivity to annual rainfall at a given site, even though observation uncertainty is comparable to inter‐model variability. (c) Most models reproduced the sign of the observed patterns in productivity changes in rainfall manipulation experiments but had a low capacity in reproducing the observed magnitude of productivity changes. Models better reproduced the observed productivity responses due to rainfall exclusion than addition. (d) All models attribute ecosystem productivity changes to the intensity of vegetation stress and peak leaf area, whereas the impact of the change in growing season length is negligible. The relative contribution of the peak leaf area and vegetation stress intensity was highly variable among models.
In this research we evaluated the skill of 10 terrestrial ecosystem models in reproducing aboveground net primary productivity responses as measured in 10 rainfall manipulation experiments.See also the Commentary on this article by Xue Feng, 26, 3190–3192
Nitrogen (N) is a critical resource for plant growth in tundra ecosystems, and species differences in the timing of N uptake may be an important feature regulating community composition and ecosystem ...productivity. We added 15 N-labelled glycine to a subarctic heath tundra dominated by dwarf shrubs, mosses and graminoids in fall, and investigated its partitioning among ecosystem components at several time points (October, November, April, May, June) through to the following spring/early summer. Soil microbes had acquired 65 ± 7% of the 15 N tracer by October, but this pool decreased through winter to 37 ± 7% by April indicating significant microbial N turnover prior to spring thaw. Only the evergreen dwarf shrubs showed active 15 N acquisition before early May indicating that they had the highest potential of all functional groups for acquiring nutrients that became available in early spring. The faster-growing deciduous shrubs did not resume 15 N acquisition until after early May indicating that they relied more on nitrogen made available later during the spring/early summer. The graminoids and mosses had no significant increases in 15 N tracer recovery or tissue 15 N tracer concentrations after the first harvest in October. However, the graminoids had the highest root 15 N, tracer concentrations of all functional groups in October indicating that they primarily relied on N made available during summer and fall. Our results suggest a temporal differentiation among plant functional groups in the post-winter resumption of N uptake with evergreen dwarf shrubs having the highest potential for early N uptake, followed by deciduous dwarf shrubs and graminoids.
Field-scale experiments simulating realistic future climate scenarios are important tools for investigating the effects of current and future climate changes on ecosystem functioning and ...biogeochemical cycling. We exposed a seminatural Danish heathland ecosystem to elevated atmospheric carbon dioxide (CO₂), warming, and extended summer drought in all combinations. Here, we report on the short-term responses of the nitrogen (N) cycle after 2 years of treatments. Elevated CO₂ significantly affected aboveground stoichiometry by increasing the carbon to nitrogen (C/N) ratios in the leaves of both co-dominant species (Calluna vulgaris and Deschampsia flexuosa), as well as the C/N ratios of Calluna flowers and by reducing the N concentration of Deschampsia litter. Belowground, elevated CO₂ had only minor effects, whereas warming increased N turnover, as indicated by increased rates of microbial NH₄ ⁺ consumption, gross mineralization, potential nitrification, denitrification and N₂O emissions. Drought reduced belowground gross N mineralization and decreased fauna N mass and fauna N mineralization. Leaching was unaffected by treatments but was significantly higher across all treatments in the second year than in the much drier first year indicating that ecosystem N loss is highly sensitive to changes and variability in amount and timing of precipitation. Interactions between treatments were common and although some synergistic effects were observed, antagonism dominated the interactive responses in treatment combinations, i.e. responses were smaller in combinations than in single treatments. Nonetheless, increased C/N ratios of photosynthetic tissue in response to elevated CO₂, as well as drought-induced decreases in litter N production and fauna N mineralization prevailed in the full treatment combination. Overall, the simulated future climate scenario therefore lead to reduced N turnover, which could act to reduce the potential growth response of plants to elevated atmospheric CO₂ concentration.
Climatic extreme events are expected to occur more frequently in the future, increasing the likelihood of unprecedented climate extremes (UCEs) or record-breaking events. UCEs, such as extreme ...heatwaves and droughts, substantially affect ecosystem stability and carbon cycling by increasing plant mortality and delaying ecosystem recovery. Quantitative knowledge of such effects is limited due to the paucity of experiments focusing on extreme climatic events beyond the range of historical experience. Here, we present a road map of how dynamic vegetation demographic models (VDMs) can be used to investigate hypotheses surrounding ecosystem responses to one type of UCE: unprecedented droughts. As a result of nonlinear ecosystem responses to UCEs that are qualitatively different from responses to milder extremes, we consider both biomass loss and recovery rates over time by reporting a time-integrated carbon loss as a result of UCE, relative to the absence of drought. Additionally, we explore how unprecedented droughts in combination with increasing atmospheric CO2 and/or temperature may affect ecosystem stability and carbon cycling. We explored these questions using simulations of pre-drought and post-drought conditions at well-studied forest sites using well-tested models (ED2 and LPJ-GUESS). The severity and patterns of biomass losses differed substantially between models. For example, biomass loss could be sensitive to either drought duration or drought intensity depending on the model approach. This is due to the models having different, but also plausible, representations of processes and interactions, highlighting the complicated variability of UCE impacts that still need to be narrowed down in models. Elevated atmospheric CO2 concentrations (eCO2) alone did not completely buffer the ecosystems from carbon losses during UCEs in the majority of our simulations. Our findings highlight the consequences of differences in process formulations and uncertainties in models, most notably related to availability in plant carbohydrate storage and the diversity of plant hydraulic schemes, in projecting potential ecosystem responses to UCEs. We provide a summary of the current state and role of many model processes that give way to different underlying hypotheses of plant responses to UCEs, reflecting knowledge gaps which in future studies could be tested with targeted field experiments and an iterative modeling–experimental conceptual framework.
This study investigated the impact of predicted future climatic and atmospheric conditions on soil respiration (RS) in a Danish Calluna‐Deschampsia‐heathland. A fully factorial in situ experiment ...with treatments of elevated atmospheric CO2 (+130 ppm), raised soil temperature (+0.4 °C) and extended summer drought (5–8% precipitation exclusion) was established in 2005. The average RS, observed in the control over 3 years of measurements (1.7 μmol CO2 m−2 sec−1), increased 38% under elevated CO2, irrespective of combination with the drought or temperature treatments. In contrast, extended summer drought decreased RS by 14%, while elevated soil temperature did not affect RS overall. A significant interaction between elevated temperature and drought resulted in further reduction of RS when these treatments were combined. A detailed analysis of short‐term RS dynamics associated with drought periods showed that RS was reduced by ~50% and was strongly correlated with soil moisture during these events. Recovery of RS to pre‐drought levels occurred within 2 weeks of rewetting; however, unexpected drought effects were observed several months after summer drought treatment in 2 of the 3 years, possibly due to reduced plant growth or changes in soil water holding capacity. An empirical model that predicts RS from soil temperature, soil moisture and plant biomass was developed and accounted for 55% of the observed variability in RS. The model predicted annual sums of RS in 2006 and 2007, in the control, were 672 and 719 g C m−2 y−1, respectively. For the full treatment combination, i.e. the future climate scenario, the model predicted that soil respiratory C losses would increase by ~21% (140–150 g C m−2 y−1). Therefore, in the future climate, stimulation of C storage in plant biomass and litter must be in excess of 21% for this ecosystem to not suffer a reduction in net ecosystem exchange.
Whether species can respond evolutionarily to current climate change is crucial for the persistence of many species. Yet, very few studies have examined genetic responses to climate change in ...manipulated experiments carried out in natural field conditions. We examined the evolutionary response to climate change in a common annelid worm using a controlled replicated experiment where climatic conditions were manipulated in a natural setting. Analyzing the transcribed genome of 15 local populations, we found that about 12% of the genetic polymorphisms exhibit differences in allele frequencies associated to changes in soil temperature and soil moisture. This shows an evolutionary response to realistic climate change happening over short‐time scale, and calls for incorporating evolution into models predicting future response of species to climate change. It also shows that designed climate change experiments coupled with genome sequencing offer great potential to test for the occurrence (or lack) of an evolutionary response.
Plant community biomass production is co‐dependent on climatic and edaphic factors that are often covarying and non‐independent. Disentangling how these factors act in isolation is challenging, ...especially along large climatic gradients that can mask soil effects. As anthropogenic pressure increasingly alters local climate and soil resource supply unevenly across landscapes, our ability to predict concurrent changes in plant community processes requires clearer understandings of independent and interactive effects of climate and soil. To address this, we developed a multispecies phytometer (i.e., standardized plant community) for separating key drivers underlying plant productivity across gradients. Phytometers were composed of three globally cosmopolitan herbaceous perennials, Dactylis glomerata, Plantago lanceolata, and Trifolium pratense. In 2017, we grew phytometer communities in 18 sites across a pan‐European aridity gradient in local site soils and a standardized substrate and compared biomass production. Standard substrate phytometers succeeded in providing a standardized climate biomass response independent of local soil effects. This allowed us to factor out climate effects in local soil phytometers, establishing that nitrogen availability did not predict biomass production, while phosphorus availability exerted a strong, positive effect independent of climate. Additionally, we identified a negative relationship between biomass production and potassium and magnesium availability. Species‐specific biomass responses to the environment in the climate‐corrected biomass were asynchronous, demonstrating the importance of species interactions in vegetation responses to global change. Biomass production was co‐limited by climatic and soil drivers, with each species experiencing its own unique set of co‐limitations. Our study demonstrates the potential of phytometers for disentangling effects of climate and soil on plant biomass production and suggests an increasing role of P limitation in the temperate regions of Europe.
In temperate regions, climate change is predicted to increase annual mean temperature and intensify the duration and frequency of summer droughts, which together with elevated atmospheric carbon ...dioxide (CO₂) concentrations, may affect the exchange of nitrous oxide (N₂O) and methane (CH₄) between terrestrial ecosystems and the atmosphere. We report results from the CLIMAITE experiment, where the effects of these three climate change parameters were investigated solely and in all combinations in a temperate heathland. Field measurements of N₂O and CH₄ fluxes took place 1–2 years after the climate change manipulations were initiated. The soil was generally a net sink for atmospheric CH₄. Elevated temperature (T) increased the CH₄ uptake by on average 10 μg C m⁻² h⁻¹, corresponding to a rise in the uptake rate of about 20%. However, during winter elevated CO₂ (CO₂) reduced the CH₄ uptake, which outweighed the positive effect of warming when analyzed across the study period. Emissions of N₂O were generally low (<10 μg N m⁻² h⁻¹). As single experimental factors, elevated CO₂, temperature and summer drought (D) had no major effect on the N₂O fluxes, but the combination of CO₂ and warming (TCO₂) stimulated N₂O emission, whereas the N₂O emission ceased when CO₂ was combined with drought (DCO₂). We suggest that these N₂O responses are related to increased rhizodeposition under elevated CO₂ combined with increased and reduced nitrogen turnover rates caused by warming and drought, respectively. The N₂O flux in the multifactor treatment TDCO₂ was not different from the ambient control treatment. Overall, our study suggests that in the future, CH₄ uptake may increase slightly, while N₂O emission will remain unchanged in temperate ecosystems on well-aerated soils. However, we propose that continued exposure to altered climate could potentially change the greenhouse gas flux pattern in the investigated heathland.
Recent evidence suggests that biogeochemical processes in the Arctic during late winter and spring-thaw strongly affect the annual cycling of carbon and nutrients, indicating high susceptibility to ...climate change. We therefore examined the carbon and nutrient dynamics in a sub-arctic heath and a birch forest with high temporal resolution from March until snowmelt at both ambient and experimentally increased snow depths. Ecosystem respiration (ER) from mid-March to snowmelt at ambient snow was high, reaching 99 ± 19 (birch) and 67 ± 1.4 g C m−2 (heath). Enhanced snow depth by about 20–30 cm increased ER by 77–157% during late winter but had no effects during spring-thaw. ER rates at the birch site were poorly described by classic first-order exponential models (R2 = 0.06–0.10) with temperature as a single variable, but model fit improved considerably by including the supply of dissolved organic carbon (DOC) or nitrogen (DON) in the model (R2 = 0.40–0.47). At the heath, model fit with temperature as the single variable was better (R2 = 0.38–0.52), yet it improved when the supply of DOC or DON was included (R2 = 0.65–0.72). Microbial carbon decreased by 43% within a few days after the first soil freeze-thaw event, while microbial nitrogen and phosphorus decreased more slowly. Because soil inorganic nitrogen and phosphorus concentrations were low, nutrients released from lysed microbial cells may have been sequestered by surviving microbes or by plants resuming growth. The fast change in microbial biomass and the dependence of ER on substrate availability stress the need for high temporal resolution in future research on ecosystem carbon and nutrient dynamics at snowmelt in order to make robust models of their turnover.