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Greenhouse gas emissions and land use change - from deforestation, forest degradation and agricultural intensification - are contributing to climate change and biodiversity loss. Important landbased strategies such as planting trees or growing bioenergy crops (with carbon capture and storage) are needed to achieve the goals of the Paris Climate Agreement and to enhance biodiversity. The integrated Land Ecosystems Atmospheric Processes Study (iLEAPS) is an international knowledge-exchange and capacity-building network, specialising in ecosystems and their role in controlling the exchange of water, energy and chemical compounds between the land surface and the atmosphere. We outline priority directions for land-atmosphere interaction research and its contribution to the sustainable development agenda.

Soil respiration rates (Rs) were measured in New Zealand dairy grassland. Both season and soil type significantly affected Rs. Soil temperature and soil type dominated overall Rs. Soil respiration (Rs), the CO2 release from root respiration and microbial metabolism, affects global soil carbon storage and cycling. Only few studies have looked at Rs in the southern hemisphere, especially regarding the interaction between soil type and environmental factors on Rs in dairy grassland. We investigated the relationship between Rs and soil temperature (Ts), soil water content (SWC), soil type, and other environmental factors based on summer and winter measurements at four sites in New Zealand. Across sites, soil respiration rates ranged from 0.29 to 14.58 with a mean of 5.38 +/- 0.13 (mean +/- standard error) mu mol CO2 m-2 s-1. Mean summer R s was 86.5% higher than mean winter Rs, largely driven by organic/gley and pumice soils while ultic soils showed very little seasonal temperature sensitivity. Overall mean Rs in organic/gley soils was 108.0% higher than that in ultic soils. The high Rs rate observed in organic/gley was likely due to high soil organic matter content, while low Rs in ultic and pallic soils resulted from high clay content and low hydraulic conductance. Soil temperature drove overall Rs. Our findings indicate that soil type and soil temperature together best explain Rs. This implies that a mere classification of land use type may be insufficient for global C models and should be supplemented with soil type information, at least locally., Bibliografiskt granskad

Soil respiration rates (Rs) were measured in New Zealand dairy grassland. Both season and soil type significantly affected Rs. Soil temperature and soil type dominated overall Rs.Soil respiration (R s), the CO2 release from root respiration and microbial metabolism, affects global soil carbon storage and cycling. Only few studies have looked at R s in the southern hemisphere, especially regarding the interaction between soil type and environmental factors on R s in dairy grassland. We investigated the relationship between R s and soil temperature (T s), soil water content (SWC), soil type, and other environmental factors based on summer and winter measurements at four sites in New Zealand. Across sites, soil respiration rates ranged from 0.29 to 14.58 with a mean of 5.38 +/- 0.13 (mean +/- standard error) mu mol CO2 m-2 s-1. Mean summer R s was 86.5% higher than mean winter R s, largely driven by organic/gley and pumice soils while ultic soils showed very little seasonal temperature sensitivity. Overall mean R s in organic/gley soils was 108.0% higher than that in ultic soils. The high R s rate observed in organic/gley was likely due to high soil organic matter content, while low R s in ultic and pallic soils resulted from high clay content and low hydraulic conductance. Soil temperature drove overall R s. Our findings indicate that soil type and soil temperature together best explain R s. This implies that a mere classification of land use type may be insufficient for global C models and should be supplemented with soil type information, at least locally.

Optimal stomatal theory predicts that stomata operate to maximise photosynthesis (A(net)) and minimise transpirational water loss to achieve optimal intrinsic water-use efficiency (iWUE). We tested whether this theory can predict stomatal responses to elevated atmospheric CO2 (eCO(2)), and whether it can capture differences in responsiveness among woody plant functional types (PFTs). We conducted a meta-analysis of tree studies of the effect of eCO(2) on iWUE and its components A(net) and stomatal conductance (g(s)). We compared three PFTs, using the unified stomatal optimisation (USO) model to account for confounding effects of leaf-air vapour pressure difference (D). We expected smaller g(s), but greater A(net), responses to eCO(2) in gymnosperms compared with angiosperm PFTs. We found that iWUE increased in proportion to increasing eCO(2) in all PFTs, and that increases in A(net) had stronger effects than reductions in g(s). The USO model correctly captured stomatal behaviour with eCO(2) across most datasets. The chief difference among PFTs was a lower stomatal slope parameter (g(1)) for the gymnosperm, compared with angiosperm, species. Land surface models can use the USO model to describe stomatal behaviour under changing atmospheric CO2 conditions.

Optimal stomatal theory predicts that stomata operate to maximise photosynthesis (A(net)) and minimise transpirational water loss to achieve optimal intrinsic water-use efficiency (iWUE). We tested whether this theory can predict stomatal responses to elevated atmospheric CO2 (eCO(2)), and whether it can capture differences in responsiveness among woody plant functional types (PFTs). We conducted a meta-analysis of tree studies of the effect of eCO(2) on iWUE and its components A(net) and stomatal conductance (g(s)). We compared three PFTs, using the unified stomatal optimisation (USO) model to account for confounding effects of leaf-air vapour pressure difference (D). We expected smaller g(s), but greater A(net), responses to eCO(2) in gymnosperms compared with angiosperm PFTs. We found that iWUE increased in proportion to increasing eCO(2) in all PFTs, and that increases in A(net) had stronger effects than reductions in g(s). The USO model correctly captured stomatal behaviour with eCO(2) across most datasets. The chief difference among PFTs was a lower stomatal slope parameter (g(1)) for the gymnosperm, compared with angiosperm, species. Land surface models can use the USO model to describe stomatal behaviour under changing atmospheric CO2 conditions.

Optimal stomatal theory predicts that stomata operate to maximise photosynthesis (A(net)) and minimise transpirational water loss to achieve optimal intrinsic water-use efficiency (iWUE). We tested whether this theory can predict stomatal responses to elevated atmospheric CO2 (eCO(2)), and whether it can capture differences in responsiveness among woody plant functional types (PFTs). We conducted a meta-analysis of tree studies of the effect of eCO(2) on iWUE and its components A(net) and stomatal conductance (g(s)). We compared three PFTs, using the unified stomatal optimisation (USO) model to account for confounding effects of leaf-air vapour pressure difference (D). We expected smaller g(s), but greater A(net), responses to eCO(2) in gymnosperms compared with angiosperm PFTs. We found that iWUE increased in proportion to increasing eCO(2) in all PFTs, and that increases in A(net) had stronger effects than reductions in g(s). The USO model correctly captured stomatal behaviour with eCO(2) across most datasets. The chief difference among PFTs was a lower stomatal slope parameter (g(1)) for the gymnosperm, compared with angiosperm, species. Land surface models can use the USO model to describe stomatal behaviour under changing atmospheric CO2 conditions.

Optimal stomatal theory predicts that stomata operate to maximise photosynthesis (Anet) and minimise transpirational water loss to achieve optimal intrinsic water-use efficiency (iWUE). We tested whether this theory can predict stomatal responses to elevated atmospheric CO2 (eCO2), and whether it can capture differences in responsiveness among woody plant functional types (PFTs). We conducted a meta-analysis of tree studies of the effect of eCO2 on iWUE and its components Anet and stomatal conductance (gs). We compared three PFTs, using the unified stomatal optimisation (USO) model to account for confounding effects of leaf–air vapour pressure difference (D). We expected smaller gs, but greater Anet, responses to eCO2 in gymnosperms compared with angiosperm PFTs. We found that iWUE increased in proportion to increasing eCO2 in all PFTs, and that increases in Anet had stronger effects than reductions in gs. The USO model correctly captured stomatal behaviour with eCO2 across most datasets. The chief difference among PFTs was a lower stomatal slope parameter (g1) for the gymnosperm, compared with angiosperm, species. Land surface models can use the USO model to describe stomatal behaviour under changing atmospheric CO2 conditions.

Responses of the terrestrial biosphere to rapidly changing environmental conditions are a major source of uncertainty in climate projections. In an effort to reduce this uncertainty, a wide range of global change experiments have been conducted that mimic future conditions in terrestrial ecosystems, manipulating CO2, temperature, and nutrient and water availability. Syntheses of results across experiments provide a more general sense of ecosystem responses to global change, and help to discern the influence of background conditions such as climate and vegetation type in determining global change responses. Several independent syntheses of published data have yielded distinct databases for specific objectives. Such parallel, uncoordinated initiatives carry the risk of producing redundant data collection efforts and have led to contrasting outcomes without clarifying the underlying reason for divergence. These problems could be avoided by creating a publicly available, updatable, curated database. Here, we report on a global effort to collect and curate 57,089 treatment responses across 3644 manipulation experiments at 1145 sites, simulating elevated CO2, warming, nutrient addition, and precipitation changes. In the resulting Manipulation Experiments Synthesis Initiative (MESI) database, effects of experimental global change drivers on carbon and nutrient cycles are included, as well as ancillary data such as background climate, vegetation type, treatment magnitude, duration, and, unique to our database, measured soil properties. Our analysis of the database indicates that most experiments are short term (one or few growing seasons), conducted in the USA, Europe, or China, and that the most abundantly reported variable is aboveground biomass. We provide the most comprehensive multifactor global change database to date, enabling the research community to tackle open research questions, vital to global policymaking. The MESI database, freely accessible at doi.org/10.528

Optimal stomatal theory predicts that stomata operate to maximise photosynthesis (A(net)) and minimise transpirational water loss to achieve optimal intrinsic water-use efficiency (iWUE). We tested whether this theory can predict stomatal responses to elevated atmospheric CO2 (eCO(2)), and whether it can capture differences in responsiveness among woody plant functional types (PFTs). We conducted a meta-analysis of tree studies of the effect of eCO(2) on iWUE and its components A(net) and stomatal conductance (g(s)). We compared three PFTs, using the unified stomatal optimisation (USO) model to account for confounding effects of leaf-air vapour pressure difference (D). We expected smaller g(s), but greater A(net), responses to eCO(2) in gymnosperms compared with angiosperm PFTs. We found that iWUE increased in proportion to increasing eCO(2) in all PFTs, and that increases in A(net) had stronger effects than reductions in g(s). The USO model correctly captured stomatal behaviour with eCO(2) across most datasets. The chief difference among PFTs was a lower stomatal slope parameter (g(1)) for the gymnosperm, compared with angiosperm, species. Land surface models can use the USO model to describe stomatal behaviour under changing atmospheric CO2 conditions.

Optimal stomatal theory predicts that stomata operate to maximise photosynthesis (A(net)) and minimise transpirational water loss to achieve optimal intrinsic water-use efficiency (iWUE). We tested whether this theory can predict stomatal responses to elevated atmospheric CO2 (eCO(2)), and whether it can capture differences in responsiveness among woody plant functional types (PFTs). We conducted a meta-analysis of tree studies of the effect of eCO(2) on iWUE and its components A(net) and stomatal conductance (g(s)). We compared three PFTs, using the unified stomatal optimisation (USO) model to account for confounding effects of leaf-air vapour pressure difference (D). We expected smaller g(s), but greater A(net), responses to eCO(2) in gymnosperms compared with angiosperm PFTs. We found that iWUE increased in proportion to increasing eCO(2) in all PFTs, and that increases in A(net) had stronger effects than reductions in g(s). The USO model correctly captured stomatal behaviour with eCO(2) across most datasets. The chief difference among PFTs was a lower stomatal slope parameter (g(1)) for the gymnosperm, compared with angiosperm, species. Land surface models can use the USO model to describe stomatal behaviour under changing atmospheric CO2 conditions.

Despite a large number of studies examining the effects of abiotic stress factors on plants, the mechanistic explanations of drought-induced tree mortality remain inconclusive and even less is known about how multiple stressors interact. The role of non-structural carbohydrates (NSCs) in preventing or postponing drought mortality is gaining attention. Here, we tested the role of NSCs in mitigating the effects of drought and salinity in New Zealand mangroves, Avicennia marina subsp. australasica. We experimentally manipulated plant NSC levels, prior to subjecting them to combinations of drought and salinity. Plant growth and survival rates were 2- and 3-fold higher in the high-NSC (H-NSC) group than in the low-NSC (L-NSC) group under high salinity and drought conditions, respectively. After 12 weeks under high salinity-high drought conditions, the H-NSC group showed higher stem hydraulic conductivity (281 +/- 50 mmol cm(-1) s(-1) MPa-1) compared with the L-NSC group (134 +/- 40 mmol cm(-1) s(-1) MPa-1). Although starch levels remained relatively constant, we found a 20% increase in soluble sugars in the stems of H-NSC group under high drought and high salinity in week 8 compared with week 12. Our results suggest (i) an important role of NSCs in mitigating the effects of low soil water potential caused by drought and salinity, and (ii) sink-limited growth under conditions of combined salinity and drought.

Atmospheric carbon dioxide concentration ([CO2]) is increasing, which increases leaf‐scale photosynthesis and intrinsic water‐use efficiency. These direct responses have the potential to increase plant growth, vegetation biomass, and soil organic matter; transferring carbon from the atmosphere into terrestrial ecosystems (a carbon sink). A substantial global terrestrial carbon sink would slow the rate of [CO2] increase and thus climate change. However, ecosystem CO2 responses are complex or confounded by concurrent changes in multiple agents of global change and evidence for a [CO2]‐driven terrestrial carbon sink can appear contradictory. Here we synthesize theory and broad, multidisciplinary evidence for the effects of increasing [CO2] (iCO2) on the global terrestrial carbon sink. Evidence suggests a substantial increase in global photosynthesis since pre‐industrial times. Established theory, supported by experiments, indicates that iCO2 is likely responsible for about half of the increase. Global carbon budgeting, atmospheric data, and forest inventories indicate a historical carbon sink, and these apparent iCO2 responses are high in comparison to experiments and predictions from theory. Plant mortality and soil carbon iCO2 responses are highly uncertain. In conclusion, a range of evidence supports a positive terrestrial carbon sink in response to iCO2, albeit with uncertain magnitude and strong suggestion of a role for additional agents of global change.

Atmospheric carbon dioxide concentration ([CO2]) is increasing, which increases leaf-scale photosynthesis and intrinsic water-use efficiency. These direct responses have the potential to increase plant growth, vegetation biomass, and soil organic matter; transferring carbon from the atmosphere into terrestrial ecosystems (a carbon sink). A substantial global terrestrial carbon sink would slow the rate of [CO2] increase and thus climate change. However, ecosystem CO2 responses are complex or confounded by concurrent changes in multiple agents of global change and evidence for a [CO2]-driven terrestrial carbon sink can appear contradictory. Here we synthesize theory and broad, multidisciplinary evidence for the effects of increasing [CO2] (iCO2) on the global terrestrial carbon sink. Evidence suggests a substantial increase in global photosynthesis since pre-industrial times. Established theory, supported by experiments, indicates that iCO2 is likely responsible for about half of the increase. Global carbon budgeting, atmospheric data, and forest inventories indicate a historical carbon sink, and these apparent iCO2 responses are high in comparison to experiments and predictions from theory. Plant mortality and soil carbon iCO2 responses are highly uncertain. In conclusion, a range of evidence supports a positive terrestrial carbon sink in response to iCO2, albeit with uncertain magnitude and strong suggestion of a role for additional agents of global change.

Anthropogenic environmental changes, or 'stressors', increasingly threaten biodiversity and ecosystem functioning worldwide. Multiple-stressor research is a rapidly expanding field of science that seeks to understand and ultimately predict the interactions between stressors. Reviews and meta-analyses of the primary scientific literature have largely been specific to either freshwater, marine or terrestrial ecology, or ecotoxicology. In this cross-disciplinary study, we review the state of knowledge within and among these disciplines to highlight commonality and division in multiple-stressor research. Our review goes beyond a description of previous research by using quantitative bibliometric analysis to identify the division between disciplines and link previously disconnected research communities. Towards a unified research framework, we discuss the shared goal of increased realism through both ecological and temporal complexity, with the overarching aim of improving predictive power. In a rapidly changing world, advancing our understanding of the cumulative ecological impacts of multiple stressors is critical for biodiversity conservation and ecosystem management. Identifying and overcoming the barriers to interdisciplinary knowledge exchange is necessary in rising to this challenge. Division between ecosystem types and disciplines is largely a human creation. Species and stressors cross these borders and so should the scientists who study them.

Direct quantification of terrestrial biosphere responses to global change is crucial for projections of future climate change in Earth system models. Here, we synthesized ecosystem carbon-cycling data from 1,119 experiments performed over the past four decades concerning changes in temperature, precipitation, CO2 and nitrogen across major terrestrial vegetation types of the world. Most experiments manipulated single rather than multiple global change drivers in temperate ecosystems of the USA, Europe and China. The magnitudes of warming and elevated CO2 treatments were consistent with the ranges of future projections, whereas those of precipitation changes and nitrogen inputs often exceeded the projected ranges. Increases in global change drivers consistently accelerated, but decreased precipitation slowed down carbon-cycle processes. Nonlinear (including synergistic and antagonistic) effects among global change drivers were rare. Belowground carbon allocation responded negatively to increased precipitation and nitrogen addition and positively to decreased precipitation and elevated CO2. The sensitivities of carbon variables to multiple global change drivers depended on the background climate and ecosystem condition, suggesting that Earth system models should be evaluated using site-specific conditions for best uses of this large dataset. Together, this synthesis underscores an urgent need to explore the interactions among multiple global change drivers in under-represented regions such as semi-arid ecosystems, forests in the tropics and subtropics, and Arctic tundra when forecasting future terrestrial carbon-climate feedback.

Direct quantification of terrestrial biosphere responses to global change is crucial for projections of future climate change in Earth system models. Here, we synthesized ecosystem carbon-cycling data from 1,119 experiments performed over the past four decades concerning changes in temperature, precipitation, CO2 and nitrogen across major terrestrial vegetation types of the world. Most experiments manipulated single rather than multiple global change drivers in temperate ecosystems of the USA, Europe and China. The magnitudes of warming and elevated CO2 treatments were consistent with the ranges of future projections, whereas those of precipitation changes and nitrogen inputs often exceeded the projected ranges. Increases in global change drivers consistently accelerated, but decreased precipitation slowed down carbon-cycle processes. Nonlinear (including synergistic and antagonistic) effects among global change drivers were rare. Belowground carbon allocation responded negatively to increased precipitation and nitrogen addition and positively to decreased precipitation and elevated CO2. The sensitivities of carbon variables to multiple global change drivers depended on the background climate and ecosystem condition, suggesting that Earth system models should be evaluated using site-specific conditions for best uses of this large dataset. Together, this synthesis underscores an urgent need to explore the interactions among multiple global change drivers in under-represented regions such as semi-arid ecosystems, forests in the tropics and subtropics, and Arctic tundra when forecasting future terrestrial carbon-climate feedback.

There is evidence of continued stimulation of foliage photosynthesis in trees exposed to elevated atmospheric CO2 concentrations; however, this is mostly without a proportional growth response. Consequently, we lack information on the fate of this extra carbon (C) acquired. By a steady application of a 13CO2 label in a free air CO2 enrichment (FACE) experiment, we traced the fate of C in 37-m-tall, ca. 110-year-old Picea abies trees in a natural forest in Switzerland. Hence, we are not reporting tree responses to elevated CO2 (which would require equally 13C labeled controls), but are providing insights into assimilate processing in such trees. Sunlit needles and branchlets grow almost exclusively from current assimilates, whereas shaded parts of the crowns also rely on stored C. Only 2.5years after FACE initiation, tree rings contained 100% new C. Stem-respiratory CO2 averaged 50% of new C over the entire FACE period. Fine roots and mycorrhizal fungi contained 49-56 and 26-43% new C, respectively, after 2.5years. The isotopic signals in soil CO2 arrived 12days after the onset of FACE, yet it contained only ca. 15% new C thereafter. We conclude that new C first feeds into fast turnover C pools in the canopy and becomes increasingly mixed with older C sources as one moves away (downward) from the crown. We speculate that enhanced C turnover (its metabolic cost) along the phloem path, as evidenced by basipetal isotope signal depletion, explains part of the ‘missing carbon' in trees that assimilated more C under elevated CO2.

Over the past 25years, countless experiments have been conducted on the impact of increased atmospheric CO2 concentration on various plants and ecosystems. While this research was motivated to better understand and predict how rising CO2 will affect the structure and function of ecosystems in the future, it also shed light on some general, CO2-research independent, aspects in ecological research. Interestingly, it is these general aspects that continue to create confusion and lead to misinterpretation. Here, we focus on seven interrelated key issues including (1) the confusion between fluxes and pools, (2) the stoichiometric aspects of growth and biomass production, (3) resource allocation within organisms, (4) data scaling and the choice of a reference metric, (5) the consideration of time and timing (experimental duration, ontogenetic shifts), (6) confounding and second-order (indirect or feedback) effects, and (7) the key role of biodiversity. The principles deriving from addressing these issues relate strongly to each other. Their concurrent consideration requires experimenters and modellers to likewise maintain a broad, holistic perspective. In this synthesis, we attempt to show how appropriate consideration of these principles can greatly enhance the assessment of the validity, plausibility and generality of experimental and modelling results. We conclude that neglecting to adequately address these key issues in ecological research may lead to overestimations of measured responses and/or simplistic interpretations. Our examples mostly originate from research on plant responses to elevated atmospheric CO2, but are also applicable to other areas of ecological research. We provide a checklist for the planning of ecological field experiments and the interpretation of their results that may help in avoiding common pitfalls

An organism's pattern of resource allocation to reproduction and growth over time critically impacts on its lifetime reproductive success. During times of low resource availability, there are two fundamental, mutually exclusive strategies of energy investment: maintenance of somatic tissues to support survival and later reproduction or investment into an immediate reproductive event at the risk of subsequent death. Here, we examine energy allocation patterns in the coral Montipora digitata to determine whether energy investment during periods of resource shortage favours growth or reproduction in a sessile, modular marine species. We manipulated light regimes (two levels of shading) on plots within a shallow reef flat habitat (Orpheus Island, Great Barrier Reef, Australia) and quantified energy uptake (rates of net photosynthesis), energy investment into reproduction (E R ), tissue growth per unit surface area (E T ) and energy channelled into calcification (E C ). With declining resource availability (i.e. reduced photosynthesis), relative energy investment shifted from high (~80%) allocation to tissue growth (E R :E T :E C =11:81:8%) to an increasing proportion channelled into reproduction and skeletal growth (20:31:49%). At the lowest light regime, calcification was maintained but reproduction was halted and thus energy content per unit surface area of tissue declined, although no mortality was observed. The changing hierarchy in energy allocation among life functions with increasing resource limitation found here for an autotrophic coral, culminating in cessation of reproduction when limitations are severe, stands in contrast to observations from annual plants. However, the strategy may be optimal for maximising fitness components (growth, reproduction and survival) through time in marine modular animals

Stomatal conductance (g s) of mature trees exposed to elevated CO2 concentrations was examined in a diverse deciduous forest stand in NW Switzerland. Measurements of g s were carried out on upper canopy foliage before noon, over four growing seasons, including an exceptionally dry summer (2003). Across all species reductions in stomatal conductance were smaller than 25% most likely around 10%, with much variation among species and trees. Given the large heterogeneity in light conditions within a tree crown, this signal was not statistically significant, but the responses within species were surprisingly consistent throughout the study period. Except during a severe drought, stomatal conductance was always lower in trees of Carpinus betulus exposed to elevated CO2 compared to Carpinus trees in ambient air, but the difference was only statistically significant on 2 out of 15 days. In contrast, stomatal responses in Fagus sylvatica and Quercus petraea varied around zero with no consistent trend in relation to CO2 treatment. During the 2003 drought in the third treatment year, the CO2 effect became reversed in Carpinus, resulting in higher g s in trees exposed to elevated CO2 compared to control trees, most likely due to better water supply because of the previous soil water savings. This was supported by less negative predawn leaf water potential in CO2 enriched Carpinus trees, indicating an improved water status. These findings illustrate (1) smaller than expected CO2-effects on stomata of mature deciduous forest trees, and (2) the possibility of soil moisture feedback on canopy water relations under elevated CO2

The global latitudinal gradient in biodiversity weakens in the high polar biome and so an alternative explanation for distribution of Arctic and Antarctic photoautotrophs is required. Here we identify how temporal, microclimate and evolutionary drivers of biogeography are important, rather than the macroclimate features that drive plant diversity patterns elsewhere. High polar ecosystems are biologically unique, with a more central role for bryophytes, lichens and microbial photoautotrophs over that of vascular plants. Constraints on vascular plants arise mainly due to stature and ontogenetic barriers. Conversely non-vascular plant and microbial photoautotroph distribution is correlated with favourable microclimates and the capacity for poikilohydric dormancy. Contemporary distribution also depends on evolutionary history, with adaptive and dispersal traits as well as legacy influencing biogeography. We highlight the relevance of these findings to predicting future impacts on polar plant diversity and to the current status of plants in Arctic and Antarctic conservation policy frameworks.

The global latitudinal gradient in biodiversity weakens in the high polar biome and so an alternative explanation for distribution of Arctic and Antarctic photoautotrophs is required. Here we identify how temporal, microclimate and evolutionary drivers of biogeography are important, rather than the macroclimate features that drive plant diversity patterns elsewhere. High polar ecosystems are biologically unique, with a more central role for bryophytes, lichens and microbial photoautotrophs over that of vascular plants. Constraints on vascular plants arise mainly due to stature and ontogenetic barriers. Conversely non-vascular plant and microbial photoautotroph distribution is correlated with favourable microclimates and the capacity for poikilohydric dormancy. Contemporary distribution also depends on evolutionary history, with adaptive and dispersal traits as well as legacy influencing biogeography. We highlight the relevance of these findings to predicting future impacts on polar plant diversity and to the current status of plants in Arctic and Antarctic conservation policy frameworks.

Most of the currently experienced global environmental changes (rising atmospheric CO2 concentrations, warming, altered amount and pattern of precipitation, and increased nutrient load) directly or indirectly affect ecosystem surface energy balance and plant transpiration. As a consequence, the relative humidity of the air surrounding the vegetation changes, thus creating a feedback loop whose net effect on transpiration and finally productivity is not trivial to quantify. Forcedly, in any global change experiment with the above drivers, we can only treat small plots, or 'islands', of vegetation. This means that the treated plots will likely experience the ambient humidity conditions influenced by the surrounding, non-treated vegetation. Experimental assessments of global change effects will thus systematically lack modifications originating from these potentially important feedback mechanisms, introducing a bias of unknown magnitude in all measurements of processes directly or indirectly depending on plant transpiration. We call this potential bias the 'island effect'. Here, we discuss its implications in various global change experiments with plants. We also suggest ways to complement experiments using modelling approaches and observational studies. Ultimately, there is no obvious solution to deal with the island effect in field experiments and only models can provide an estimate of modification of responses by these feedbacks. However, we suggest that increasing the awareness of the island effect among both experimental researchers and modellers will greatly improve the interpretation of vegetation responses to global change.

Dynamic vegetation models have been widely used for analyzing ecosystem dynamics and their interactions with climate. Their performance has been tested extensively against observations and by model intercomparison studies. In the present analysis, Lund-Potsdam-Jena General Ecosystem Simulator (LPJ-GUESS), a state-of-the-art ecosystem model, was evaluated by performing a global sensitivity analysis. The study aims at examining potential model limitations, particularly with regard to long-term applications. A detailed sensitivity analysis based on variance decomposition is presented to investigate structural model assumptions and to highlight processes and parameters that cause the highest variability in the output. First- and total-order sensitivity indices were calculated for selected parameters using Sobol's methodology. In order to elucidate the role of climate on model sensitivity, different climate forcings were used based on observations from Switzerland. The results clearly indicate a very high sensitivity of LPJ-GUESS to photosynthetic parameters. Intrinsic quantum efficiency alone is able to explain about 60% of the variability in vegetation carbon fluxes and pools for a wide range of climate forcings. Processes related to light harvesting were also found to be important together with parameters affecting forest structure (growth, establishment, and mortality). The model shows minor sensitivity to hydrological and soil texture parameters, questioning its skills in representing spatial vegetation heterogeneity at regional or watershed scales. In the light of these results, we discuss the deficiencies of LPJ-GUESS and possibly that of other, structurally similar, dynamic vegetation models and we highlight potential directions for further model improvements.

Aim Despite increasing evidence for plant growth often being limited by sink (meristem) activity rather than source (photosynthesis) activity, all currently available dynamic global vegetation models (DGVMs) simulate plant growth via source-limited processes. For a given climatic region, this may lead to an overestimation of carbon stock per unit surface area, particularly if a model fails to correctly predict forest cover. Our aim is to improve the Lund-Potsdam-Jena (LPJ) DGVM by replacing the source-limited (SoL) tree growth algorithm by a sink-limited (SiL) one. Location Our analysis focuses on the cold tree line at high latitudes and altitudes. We study two altitudinal transects in the Swiss Alps and the northern tree line. Methods We limit annual net primary productivity of the LPJ DGVM by an algorithm based on the annual sum of growing degree-days (GDD), assuming that maximum plant growth is reached asymptotically with increasing GDD. Results Comparing simulation results with observational data, we show that the locations of both the northern and the alpine tree line are estimated more accurately when using a SiL algorithm than when using the commonly employed SoL algorithm. Also, simulated carbon stocks decrease in a more realistic manner towards the tree line when the SiL algorithm is used. This has far-reaching implications for estimating and projecting present and future carbon stocks in temperature-limited ecosystems. Main conclusions In the range of 60-80 degrees N over Europe and Asia, carbon stored in vegetation is estimated to be c. 50% higher in the LPJ standard version (LPJ-SoL) compared with LPJ-SiL, resulting in a global difference in estimated biomass of 25 Pg (c. 5% of the global terrestrial standing biomass). Similarly, the simulated elevation of the upper tree line in the European Alps differs by c. 400 m between the two model versions, thus implying an additional overestimation of carbon stored in mountain forests around the world.

We review observational, experimental, and model results on how plants respond to extreme climatic conditions induced by changing climatic variability. Distinguishing between impacts of changing mean climatic conditions and changing climatic variability on terrestrial ecosystems is generally underrated in current studies. The goals of our review are thus (1) to identify plant processes that are vulnerable to changes in the variability of climatic variables rather than to changes in their mean, and (2) to depict/evaluate available study designs to quantify responses of plants to changing climatic variability. We find that phenology is largely affected by changing mean climate but also that impacts of climatic variability are much less studied, although potentially damaging. We note that plant water relations seem to be very vulnerable to extremes driven by changes in temperature and precipitation and that heatwaves and flooding have stronger impacts on physiological processes than changing mean climate. Moreover, interacting phenological and physiological processes are likely to further complicate plant responses to changing climatic variability. Phenological and physiological processes and their interactions culminate in even more sophisticated responses to changing mean climate and climatic variability at the species and community level. Generally, observational studies are well suited to study plant responses to changing mean climate, but less suitable to gain a mechanistic understanding of plant responses to climatic variability. Experiments seem best suited to simulate extreme events. In models, temporal resolution and model structure are crucial to capture plant responses to changing climatic variability. We highlight that a combination of experimental, observational, and/or modeling studies have the potential to overcome important caveats of the respective individual approaches.

Climatic changes, including altered precipitation regimes, will affect key ecosystem processes, such as plant productivity and biodiversity for many terrestrial ecosystems. Past and ongoing precipitation experiments have been conducted to quantify these potential changes. An analysis of these experiments indicates that they have provided important information on how water regulates ecosystem processes. However, they do not adequately represent global biomes nor forecasted precipitation scenarios and their potential contribution to advance our understanding of ecosystem responses to precipitation changes is therefore limited, as is their potential value for the development and testing of ecosystem models. This highlights the need for new precipitation experiments in biomes and ambient climatic conditions hitherto poorly studied applying relevant complex scenarios including changes in precipitation frequency and amplitude, seasonality, extremity and interactions with other global change drivers. A systematic and holistic approach to investigate how soil and plant community characteristics change with altered precipitation regimes and the consequent effects on ecosystem processes and functioning within these experiments will greatly increase their value to the climate change and ecosystem research communities. Experiments should specifically test how changes in precipitation leading to exceedance of biological thresholds affect ecosystem resilience and acclimation.

Climatic changes, including altered precipitation regimes, will affect key ecosystem processes, such as plant productivity and biodiversity for many terrestrial ecosystems. Past and ongoing precipitation experiments have been conducted to quantify these potential changes. An analysis of these experiments indicates that they have provided important information on how water regulates ecosystem processes. However, they do not adequately represent global biomes nor forecasted precipitation scenarios and their potential contribution to advance our understanding of ecosystem responses to precipitation changes is therefore limited, as is their potential value for the development and testing of ecosystem models. This highlights the need for new precipitation experiments in biomes and ambient climatic conditions hitherto poorly studied applying relevant complex scenarios including changes in precipitation frequency and amplitude, seasonality, extremity and interactions with other global change drivers. A systematic and holistic approach to investigate how soil and plant community characteristics change with altered precipitation regimes and the consequent effects on ecosystem processes and functioning within these experiments will greatly increase their value to the climate change and ecosystem research communities. Experiments should specifically test how changes in precipitation leading to exceedance of biological thresholds affect ecosystem resilience and acclimation.

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 result

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 result

In recent decades, many climate manipulation experiments have investigated biosphere responses to global change. These experiments typically examined effects of elevated atmospheric CO2, warming or drought (driver variables) on ecosystem processes such as the carbon and water cycle (response variables). Because experiments are inevitably constrained in the number of driver variables tested simultaneously, as well as in time and space, a key question is how results are scaled up to predict net ecosystem responses. In this review, we argue that there might be a general trend for the magnitude of the responses to decline with higher-order interactions, longer time periods and larger spatial scales. This means that on average, both positive and negative global change impacts on the biosphere might be dampened more than previously assumed.

There is evidence of continued stimulation of foliage photosynthesis in trees exposed to elevated atmospheric CO2 concentrations; however, this is mostly without a proportional growth response. Consequently, we lack information on the fate of this extra carbon (C) acquired. By a steady application of a 13CO2 label in a free air CO2 enrichment (FACE) experiment, we traced the fate of C in 37-m-tall, ca. 110-year-old Picea abies trees in a natural forest in Switzerland. Hence, we are not reporting tree responses to elevated CO2 (which would require equally 13C labeled controls), but are providing insights into assimilate processing in such trees. Sunlit needles and branchlets grow almost exclusively from current assimilates, whereas shaded parts of the crowns also rely on stored C. Only 2.5years after FACE initiation, tree rings contained 100% new C. Stem-respiratory CO2 averaged 50% of new C over the entire FACE period. Fine roots and mycorrhizal fungi contained 49-56 and 26-43% new C, respectively, after 2.5years. The isotopic signals in soil CO2 arrived 12days after the onset of FACE, yet it contained only ca. 15% new C thereafter. We conclude that new C first feeds into fast turnover C pools in the canopy and becomes increasingly mixed with older C sources as one moves away (downward) from the crown. We speculate that enhanced C turnover (its metabolic cost) along the phloem path, as evidenced by basipetal isotope signal depletion, explains part of the ‘missing carbon' in trees that assimilated more C under elevated CO2.

An organism's pattern of resource allocation to reproduction and growth over time critically impacts on its lifetime reproductive success. During times of low resource availability, there are two fundamental, mutually exclusive strategies of energy investment: maintenance of somatic tissues to support survival and later reproduction or investment into an immediate reproductive event at the risk of subsequent death. Here, we examine energy allocation patterns in the coral Montipora digitata to determine whether energy investment during periods of resource shortage favours growth or reproduction in a sessile, modular marine species. We manipulated light regimes (two levels of shading) on plots within a shallow reef flat habitat (Orpheus Island, Great Barrier Reef, Australia) and quantified energy uptake (rates of net photosynthesis), energy investment into reproduction (E R ), tissue growth per unit surface area (E T ) and energy channelled into calcification (E C ). With declining resource availability (i.e. reduced photosynthesis), relative energy investment shifted from high (~80%) allocation to tissue growth (E R :E T :E C =11:81:8%) to an increasing proportion channelled into reproduction and skeletal growth (20:31:49%). At the lowest light regime, calcification was maintained but reproduction was halted and thus energy content per unit surface area of tissue declined, although no mortality was observed. The changing hierarchy in energy allocation among life functions with increasing resource limitation found here for an autotrophic coral, culminating in cessation of reproduction when limitations are severe, stands in contrast to observations from annual plants. However, the strategy may be optimal for maximising fitness components (growth, reproduction and survival) through time in marine modular animals

Stomatal conductance (g s) of mature trees exposed to elevated CO2 concentrations was examined in a diverse deciduous forest stand in NW Switzerland. Measurements of g s were carried out on upper canopy foliage before noon, over four growing seasons, including an exceptionally dry summer (2003). Across all species reductions in stomatal conductance were smaller than 25% most likely around 10%, with much variation among species and trees. Given the large heterogeneity in light conditions within a tree crown, this signal was not statistically significant, but the responses within species were surprisingly consistent throughout the study period. Except during a severe drought, stomatal conductance was always lower in trees of Carpinus betulus exposed to elevated CO2 compared to Carpinus trees in ambient air, but the difference was only statistically significant on 2 out of 15 days. In contrast, stomatal responses in Fagus sylvatica and Quercus petraea varied around zero with no consistent trend in relation to CO2 treatment. During the 2003 drought in the third treatment year, the CO2 effect became reversed in Carpinus, resulting in higher g s in trees exposed to elevated CO2 compared to control trees, most likely due to better water supply because of the previous soil water savings. This was supported by less negative predawn leaf water potential in CO2 enriched Carpinus trees, indicating an improved water status. These findings illustrate (1) smaller than expected CO2-effects on stomata of mature deciduous forest trees, and (2) the possibility of soil moisture feedback on canopy water relations under elevated CO2

Over the past 25years, countless experiments have been conducted on the impact of increased atmospheric CO2 concentration on various plants and ecosystems. While this research was motivated to better understand and predict how rising CO2 will affect the structure and function of ecosystems in the future, it also shed light on some general, CO2-research independent, aspects in ecological research. Interestingly, it is these general aspects that continue to create confusion and lead to misinterpretation. Here, we focus on seven interrelated key issues including (1) the confusion between fluxes and pools, (2) the stoichiometric aspects of growth and biomass production, (3) resource allocation within organisms, (4) data scaling and the choice of a reference metric, (5) the consideration of time and timing (experimental duration, ontogenetic shifts), (6) confounding and second-order (indirect or feedback) effects, and (7) the key role of biodiversity. The principles deriving from addressing these issues relate strongly to each other. Their concurrent consideration requires experimenters and modellers to likewise maintain a broad, holistic perspective. In this synthesis, we attempt to show how appropriate consideration of these principles can greatly enhance the assessment of the validity, plausibility and generality of experimental and modelling results. We conclude that neglecting to adequately address these key issues in ecological research may lead to overestimations of measured responses and/or simplistic interpretations. Our examples mostly originate from research on plant responses to elevated atmospheric CO2, but are also applicable to other areas of ecological research. We provide a checklist for the planning of ecological field experiments and the interpretation of their results that may help in avoiding common pitfalls