Freshwater biota and rising pCO2?

Rising atmospheric carbon dioxide (CO₂) has caused a suite of environmental issues, however, little is known about how the partial pressure of CO₂ (pCO₂) in freshwater will be affected by climate change. Freshwater pCO₂ varies across systems and is controlled by a diverse array of factors, making it difficult to make predictions about future levels of pCO₂. Recent evidence suggests that increasing levels of atmospheric CO₂ may directly increase freshwater pCO₂ levels in lakes, but rising atmospheric CO₂ may also indirectly impact freshwater pCO₂ levels in a variety of systems by affecting other contributing factors such as soil respiration, terrestrial productivity and climate regimes. Although future freshwater pCO₂ levels remain uncertain, studies have considered the potential impacts of changes to pCO₂ levels on freshwater biota. Studies to date have focused on impacts of elevated pCO₂ on plankton and macrophytes, and have shown that phytoplankton nutritional quality is reduced, plankton community structure is altered, photosynthesis rates increase and macrophyte distribution shifts with increasing pCO₂. However, a number of key knowledge gaps remain and gaining a better understanding of how freshwater pCO₂ levels are regulated and how these levels may impact biota, will be important for predicting future responses to climate change.     

Interacting effects of ocean acidification and warming on growth and DMS‐production in the haptophyte coccolithophore Emiliania huxleyi

The production of the marine trace gas dimethyl sulfide (DMS) provides 90% of the marine biogenic sulfur in the atmosphere where it affects cloud formation and climate. The effects of increasing anthropogenic CO₂ and the resulting warming and ocean acidification on trace gas production in the oceans are poorly understood. Here we report the first measurements of DMS‐production and data on growth, DMSP and DMS concentrations in pH‐stated cultures of the phytoplankton haptophyte Emiliania huxleyi. Four different environmental conditions were tested: ambient, elevated CO₂ (+CO₂), elevated temperature (+T) and elevated temperature and CO₂ (+TCO₂). In comparison to the ambient treatment, average DMS production was about 50% lower in the +CO₂ treatment. Importantly, temperature had a strong effect on DMS production and the impacts outweighed the effects of a decrease in pH. As a result, the +T and +TCO₂ treatments showed significantly higher DMS production of 36.2 ± 2.58 and 31.5 ± 4.66 μmol L^?1 cell volume (CV) h^?1 in comparison with the +CO₂ treatment (14.9 ± 4.20 μmol L^?1 CV h^?1). As the cultures were aerated with an air/CO₂ mixture, DMS was effectively removed from the incubation bottles so that concentration remained relatively low (3.6–6.1 mmol L^?1 CV). Intracellular DMSP has been shown to increase in E. huxleyi as a result of elevated temperature and/or elevated CO₂ and our results are in agreement with this finding: the ambient and +CO₂ treatments showed 125 ± 20.4 and 162 ± 27.7 mmol L^?1 CV, whereas +T and +TCO₂ showed significantly increased intracellular DMSP concentrations of 195 ± 15.8 and 211 ± 28.2 mmol L^?1 CV respectively. Growth was unaffected by the treatments, but cell diameter decreased significantly under elevated temperature. These results indicate that DMS production is sensitive to CO₂ and temperature in E. huxleyi. Hence, global environmental change that manifests in ocean acidification and warming may not result in decreased DMS as suggested by earlier studies investigating the effect of elevated CO₂ in isolation.     

Land plants equilibrate O2 and CO2 concentrations in the atmosphere

The role of land plants in establishing our present day atmosphere is analysed. Before the evolution of land plants, photosynthesis by marine and fresh water organisms was not intensive enough to deplete CO₂ from the atmosphere, the concentration of which was more than the order of magnitude higher than present. With the appearance of land plants, the exudation of organic acids by roots, following respiratory and photorespiratory metabolism, led to phosphate weathering from rocks thus increasing aquatic productivity. Weathering also replaced silicates by carbonates, thus decreasing the atmospheric CO₂ concentration. As a result of both intensive photosynthesis and weathering, CO₂ was depleted from the atmosphere down to low values approaching the compensation point of land plants. During the same time period, the atmospheric O₂ concentration increased to maximum levels about 300 million years ago (Permo-Carboniferous boundary), establishing an O₂/CO₂ ratio above 1000. At this point, land plant productivity and weathering strongly decreased, exerting negative feedback on aquatic productivity. Increased CO₂ concentrations were triggered by asteroid impacts and volcanic activity and in the Mesozoic era could be related to the gymnosperm flora with lower metabolic and weathering rates. A high O₂/CO₂ ratio is metabolically linked to the formation of citrate and oxalate, the main factors causing weathering, and to the production of reactive oxygen species, which triggered mutations and stimulated the evolution of land plants. The development of angiosperms resulted in a decrease in CO₂ concentration during the Cenozoic era, which finally led to the glacial-interglacial oscillations in the Pleistocene epoch. Photorespiration, the rate of which is directly related to the O₂/CO₂ ratio, due to the dual function of Rubisco, may be an important mechanism in maintaining the limits of O₂ and CO₂ concentrations by restricting land plant productivity and weathering.     

Quantitative estimates of changes in marine and terrestrial primary productivity over the past 300 million years

Changes in marine primary production over geological time have influenced a network of global biogeochemical cycles with corresponding feedbacks on climate. However, these changes continue to remain largely unquantified because of uncertainties in calculating global estimates from sedimentary palaeoproductivity indicators. I therefore describe a new approach to the problem using a mass balance analysis of the stable isotopes (¹⁸O/¹⁶O) of oxygen with modelled O₂ fluxes and isotopic exchanges by terrestrial vegetation for 300, 150, 100 and 50 million years before present, and the treatment of the Earth as a closed system, with respect to the cycling of O₂. Calculated in this way, oceanic net primary productivity was low in the Carboniferous but high (up to four times that of modern oceans) during the Late Jurassic, mid-Cretaceous and early Eocene greenhouse eras with a greater requirement for key nutrients. Such a requirement would be compatible with accelerated rates of continental weathering under the greenhouse conditions of the Mesozoic and early Tertiary. These results indicate possible changes in the strength of a key component of the oceanic carbon (organic and carbonate) pump in the geological past, with a corresponding feedback on atmospheric CO₂ and climate, and provide an improved framework for understanding the role of ocean biota in the evolution of the global biogeochemical cycles of C, N and P.     

Trends in tuna carbon isotopes suggest global changes in pelagic phytoplankton communities

Considerable uncertainty remains over how increasing atmospheric CO₂ and anthropogenic climate changes are affecting open‐ocean marine ecosystems from phytoplankton to top predators. Biological time series data are thus urgently needed for the world's oceans. Here, we use the carbon stable isotope composition of tuna to provide a first insight into the existence of global trends in complex ecosystem dynamics and changes in the oceanic carbon cycle. From 2000 to 2015, considerable declines in δ¹³C values of 0.8‰–2.5‰ were observed across three tuna species sampled globally, with more substantial changes in the Pacific Ocean compared to the Atlantic and Indian Oceans. Tuna recorded not only the Suess effect, that is, fossil fuel‐derived and isotopically light carbon being incorporated into marine ecosystems, but also recorded profound changes at the base of marine food webs. We suggest a global shift in phytoplankton community structure, for example, a reduction in ¹³C‐rich phytoplankton such as diatoms, and/or a change in phytoplankton physiology during this period, although this does not rule out other concomitant changes at higher levels in the food webs. Our study establishes tuna δ¹³C values as a candidate essential ocean variable to assess complex ecosystem responses to climate change at regional to global scales and over decadal timescales. Finally, this time series will be invaluable in calibrating and validating global earth system models to project changes in marine biota.     

How uncertainties in future climate change predictions translate into future terrestrial carbon fluxes

We forced a global terrestrial carbon cycle model by climate fields of 14 ocean and atmosphere general circulation models (OAGCMs) to simulate the response of terrestrial carbon pools and fluxes to climate change over the next century. These models participated in the second phase of the Coupled Model Intercomparison Project (CMIP2), where a 1% per year increase of atmospheric CO₂ was prescribed. We obtain a reduction in net land uptake because of climate change ranging between 1.4 and 5.7 Gt C yr^?1 at the time of atmospheric CO₂ doubling. Such a reduction in terrestrial carbon sinks is largely dominated by the response of tropical ecosystems, where soil water stress occurs. The uncertainty in the simulated land carbon cycle response is the consequence of discrepancies in land temperature and precipitation changes simulated by the OAGCMs. We use a statistical approach to assess the coherence of the land carbon fluxes response to climate change. The biospheric carbon fluxes and pools changes have a coherent response in the tropics, in the Mediterranean region and in high latitudes of the Northern Hemisphere. This is because of a good coherence of soil water content change in the first two regions and of temperature change in the high latitudes of the Northern Hemisphere. Then we evaluate the carbon uptake uncertainties to the assumptions on plant productivity sensitivity to atmospheric CO₂ and on decomposition rate sensitivity to temperature. We show that these uncertainties are on the same order of magnitude than the uncertainty because of climate change. Finally, we find that the OAGCMs having the largest climate sensitivities to CO₂ are the ones with the largest soil drying in the tropics, and therefore with the largest reduction of carbon uptake.     

Upward increase of kerogen δ13C in the Peru Margin Upper Oligocene: possible implications for the Cenozoic evolution of atmospheric CO2

Carbon isotopic composition of predominantly marine kerogen in latest Oligocene mudstones of the Peru Margin ODP 682A Hole shows an about 3.5‰ increase with decreasing age. Py-GC and elemental (C/N ratio) analysis of the kerogen plus sulphur isotopic study together with earlier knowledge on geological setting and organic geochemistry results in a better understanding of depositionary environment and allows to separation of the influence of concentration of water dissolved carbon dioxide (c_e) on kerogen δ¹³C from that of other factors (bacterial degradation, sea surface temperature, DIC δ¹³C, productivity, and admixture of land plant OM). Based on this analysis, the major part of the kerogen shift is considered as a result of the latest Oligocene decrease of marine photosynthetic carbon isotopic fractionation in the Peru Margin photic zone, which in turn possibly reflects a simultaneous drop in atmospheric CO₂ level. Uncertainties in the evaluation of the factors affecting the marine photosynthetic carbon isotopic fractionation and the extent of ocean–atmosphere disequilibrium do not permit calculation of the decrease of the atmospheric CO₂.     

Increase of atmospheric CO2 promotes phytoplankton productivity

It is usually thought that unlike terrestrial plants, phytoplankton will not show a significant response to an increase of atmospheric CO₂. Here we suggest that this view may be biased by a neglect of the effects of carbon (C) assimilation on the pH and the dissociation of the C species. We show that under eutrophic conditions, productivity may double as a result of doubling of the atmospheric CO₂ concentration. Although in practice productivity increase will usually be less, we still predict a productivity increase of up to 40% in marine species with a low affinity for bicarbonate. In eutrophic freshwater systems doubling of atmospheric CO₂ may result in an increase of the productivity of more than 50%. Freshwaters with low alkalinity appeared to be very sensitive to atmospheric CO₂ elevation. Our results suggest that the aquatic C sink may increase more than expected, and that nuisance phytoplankton blooms may be aggravated at elevated atmospheric CO₂ concentrations.     

Modified future diurnal variability of the global surface ocean CO2 system

Our understanding of how increasing atmospheric CO₂ and climate change influences the marine CO₂ system and in turn ecosystems has increasingly focused on perturbations to carbonate chemistry variability. This variability can affect ocean-climate feedbacks and has been shown to influence marine ecosystems. The seasonal variability of the ocean CO₂ system has already changed, with enhanced seasonal variations in the surface ocean pCO₂ over recent decades and further amplification projected by models over the 21st century. Mesocosm studies and CO₂ vent sites indicate that diurnal variability of the CO₂ system, the amplitude of which in extreme events can exceed that of mean seasonal variability, is also likely to be altered by climate change. Here, we modified a global ocean biogeochemical model to resolve physically and biologically driven diurnal variability of the ocean CO₂ system. Forcing the model with 3-h atmospheric outputs derived from an Earth system model, we explore how surface ocean diurnal variability responds to historical changes and project how it changes under two contrasting 21st-century emission scenarios. Compared to preindustrial values, the global mean diurnal amplitude of pCO₂ increases by 4.8 μatm (+226%) in the high-emission scenario but only 1.2 μatm (+55%) in the high-mitigation scenario. The probability of extreme diurnal amplitudes of pCO₂ and [H⁺] is also affected, with 30- to 60-fold increases relative to the preindustrial under high 21st-century emissions. The main driver of heightened pCO₂ diurnal variability is the enhanced sensitivity of pCO₂ to changes in temperature as the ocean absorbs atmospheric CO₂. Our projections suggest that organisms in the future ocean will be exposed to enhanced diurnal variability in pCO₂ and [H⁺], with likely increases in the associated metabolic cost that such variability imposes.     

Significance of ocean carbonate budgets for the global carbon cycle

Changes in the trace gas composition of the atmosphere over glacial–interglacial cycles are linked to changes in the oceanic carbon cycle. This paper examines the role of biologically driven fluxes of organic and inorganic carbon in modifying the carbon dioxide chemistry of the oceans, and the corresponding implications for the partitioning of CO₂ between the atmosphere and ocean. Relevant details of the marine carbon system are presented together with an assessment of the significance of remineralization and dissolution processes. Recent estimates of the marine carbonate fluxes show significant uncertainties and inconsistencies which must be resolved in order to assess fully the role of the oceans' biota in the marine carbon system. Various types of ocean carbon cycle models have been developed in order to interpret the changes in past atmospheric carbon dioxide. Some take account of the role of the oceans' biota, focussing in the main on the cycling of organic matter. Relatively few have considered the role of the carbonate pump and the subtle interactions between organic and inorganic carbon cycling. The significance of carbonate formation and dissolution, and of the effects of global change on the marine carbonate system, for air–sea fluxes of CO₂ are discussed. Finally some recommendations for future research are made in order to improve our understanding of how spatial and temporal variation in marine carbonate fluxes, in conjunction with processes determining the oxidation and burial of organic matter in the oceans, affect levels of CO₂ in the atmosphere.     

Causes of slowing‐down seasonal CO2 amplitude at Mauna Loa

Changing amplitude of the seasonal cycle of atmospheric CO₂ (SCA) in the northern hemisphere is an emerging carbon cycle property. Mauna Loa (MLO) station (20°N, 156°W), which has the longest continuous northern hemisphere CO₂ record, shows an increasing SCA before the 1980s (p < .01), followed by no significant change thereafter. We analyzed the potential driving factors of SCA slowing‐down, with an ensemble of dynamic global vegetation models (DGVMs) coupled with an atmospheric transport model. We found that slowing‐down of SCA at MLO is primarily explained by response of net biome productivity (NBP) to climate change, and by changes in atmospheric circulations. Through NBP, climate change increases SCA at MLO before the 1980s and decreases it afterwards. The effect of climate change on the slowing‐down of SCA at MLO is mainly exerted by intensified drought stress acting to offset the acceleration driven by CO₂ fertilization. This challenges the view that CO₂ fertilization is the dominant cause of emergent SCA trends at northern sites south of 40°N. The contribution of agricultural intensification on the deceleration of SCA at MLO was elusive according to land–atmosphere CO₂ flux estimated by DGVMs and atmospheric inversions. Our results also show the necessity to adequately account for changing circulation patterns in understanding carbon cycle dynamics observed from atmospheric observations and in using these observations to benchmark DGVMs.     

Understorey productivity in temperate grassy woodland responds to soil water availability but not to elevated [CO2]

Rising atmospheric [CO₂] and associated climate change are expected to modify primary productivity across a range of ecosystems globally. Increasing aridity is predicted to reduce grassland productivity, although rising [CO₂] and associated increases in plant water use efficiency may partially offset the effect of drying on growth. Difficulties arise in predicting the direction and magnitude of future changes in ecosystem productivity, due to limited field experimentation investigating climate and CO₂ interactions. We use repeat near‐surface digital photography to quantify the effects of water availability and experimentally manipulated elevated [CO₂] (eCO₂) on understorey live foliage cover and biomass over three growing seasons in a temperate grassy woodland in south‐eastern Australia. We hypothesised that (i) understorey herbaceous productivity is dependent upon soil water availability, and (ii) that eCO₂ will increase productivity, with greatest stimulation occurring under conditions of low water availability. Soil volumetric water content (VWC) determined foliage cover and growth rates over the length of the growing season (August to March), with low VWC (<0.1 m³ m^?3) reducing productivity. However, eCO₂ did not increase herbaceous cover and biomass over the duration of the experiment, or mitigate the effects of low water availability on understorey growth rates and cover. Our findings suggest that projected increases in aridity in temperate woodlands are likely to lead to reduced understorey productivity, with little scope for eCO₂ to offset these changes.     

Responses of CAM species to increasing atmospheric CO2 concentrations

Crassulacean acid metabolism (CAM) species show an average increase in biomass productivity of 35% in response to a doubled atmospheric CO₂ concentration. Daily net CO₂ uptake is similarly enhanced, reflecting in part an increase in chlorenchyma thickness and accompanied by an even greater increase in water‐use efficiency. The responses of net CO₂ uptake in CAM species to increasing atmospheric CO₂ concentrations are similar to those for C₃ species and much greater than those for C₄ species. Increases in net daily CO₂ uptake by CAM plants under elevated atmospheric CO₂ concentrations reflect increases in both Rubisco‐mediated daytime CO₂ uptake and phosphoenolpyruvate carboxylase (PEPCase)‐mediated night‐time CO₂ uptake, the latter resulting in increased nocturnal malate accumulation. Chlorophyll contents and the activities of Rubisco and PEPCase decrease under elevated atmospheric CO₂, but the activated percentage for Rubisco increases and the K_M(HCO₃ ^? ) for PEPCase decreases, resulting in more efficient photosynthesis. Increases in root:shoot ratios and the formation of additional photosynthetic organs, together with increases in sucrose‐Pi synthase and starch synthase activity in these organs under elevated atmospheric CO₂ concentrations, decrease the potential feedback inhibition of photosynthesis. Longer‐term studies for several CAM species show no downward acclimatization of photosynthesis in response to elevated atmospheric CO₂ concentrations. With increasing temperature and drought duration, the percentage enhancement of daily net CO₂ uptake caused by elevated atmospheric CO₂ concentrations increases. Thus net CO₂ uptake, productivity, and the potential area for cultivation of CAM species will be enhanced by the increasing atmospheric CO₂ concentrations and the increasing temperatures associated with global climate change.     

Global response of terrestrial ecosystem structure and function to CO2 and climate change: results from six dynamic global vegetation models

The possible responses of ecosystem processes to rising atmospheric CO₂ concentration and climate change are illustrated using six dynamic global vegetation models that explicitly represent the interactions of ecosystem carbon and water exchanges with vegetation dynamics. The models are driven by the IPCC IS92a scenario of rising CO₂ ( Wigley et al. 1991 ), and by climate changes resulting from effective CO₂ concentrations corresponding to IS92a, simulated by the coupled ocean atmosphere model HadCM2‐SUL. Simulations with changing CO₂ alone show a widely distributed terrestrial carbon sink of 1.4–3.8 Pg C y^?1 during the 1990s, rising to 3.7–8.6 Pg C y^?1 a century later. Simulations including climate change show a reduced sink both today (0.6–3.0 Pg C y^?1) and a century later (0.3–6.6 Pg C y^?1) as a result of the impacts of climate change on NEP of tropical and southern hemisphere ecosystems. In all models, the rate of increase of NEP begins to level off around 2030 as a consequence of the ‘diminishing return’ of physiological CO₂ effects at high CO₂ concentrations. Four out of the six models show a further, climate‐induced decline in NEP resulting from increased heterotrophic respiration and declining tropical NPP after 2050. Changes in vegetation structure influence the magnitude and spatial pattern of the carbon sink and, in combination with changing climate, also freshwater availability (runoff). It is shown that these changes, once set in motion, would continue to evolve for at least a century even if atmospheric CO₂ concentration and climate could be instantaneously stabilized. The results should be considered illustrative in the sense that the choice of CO₂ concentration scenario was arbitrary and only one climate model scenario was used. However, the results serve to indicate a range of possible biospheric responses to CO₂ and climate change. They reveal major uncertainties about the response of NEP to climate change resulting, primarily, from differences in the way that modelled global NPP responds to a changing climate. The simulations illustrate, however, that the magnitude of possible biospheric influences on the carbon balance requires that this factor is taken into account for future scenarios of atmospheric CO₂ and climate change.     

Changes in microbial activity and composition in a pasture ecosystem exposed to elevated atmospheric carbon dioxide

Elevated atmospheric CO₂ increases aboveground plant growth and productivity. However, carbon dioxide-induced alterations in plant growth are also likely to affect belowground processes, including the composition of soil biota. We investigated the influence of increased atmospheric CO₂on bacterial numbers and activity, and on soil microbial community composition in a pasture ecosystem under Free-Air Carbon Dioxide Enrichment (FACE). Composition of the soil microbial communities, in rhizosphere and bulk soil, under two atmospheric CO₂ levels was evaluated by using phospholipid fatty acid analysis (PLFA), and total and respiring bacteria counts were determined by epifluorescence microscopy. While populations increased with elevated atmospheric CO₂ in bulk soil of white clover (Trifolium repens L.), a higher atmospheric CO₂ concentration did not affect total or metabolically active bacteria in bulk soil of perennial ryegrass (Lolium perenne L.). There was no effect of atmospheric CO₂ on total bacteria populations per gram of rhizosphere soil. The combined effect of elevated CO₂ on total root length of each species and the bacterial population in these rhizospheres, however, resulted in an 85% increase in total rhizosphere bacteria and a 170% increase in respiring rhizosphere bacteria for the two plant species, when assessed on a per unit land area basis. Differences in microbial community composition between rhizosphere and bulk soil were evident in samples from white clover, and these communities changed in response to CO₂ enrichment. Results of this study indicate that changes in soil microbial activity, numbers, and community composition are likely to occur under elevated atmospheric CO₂, but the extent of those changes depend on plant species and the distance that microbes are from the immediate vicinity of the plant root surface.     

Implications of CO2 fertilization for future climate change in a coupled climate–carbon model

The terrestrial carbon cycle plays a critical role in determining levels of atmospheric CO₂ that result from anthropogenic carbon emissions. Elevated atmospheric CO₂ is thought to stimulate terrestrial carbon uptake, through the process of CO₂ fertilization of vegetation productivity. This negative carbon cycle feedback results in reduced atmospheric CO₂ growth, and has likely accounted for a substantial portion of the historical terrestrial carbon sink. However, the future strength of CO₂ fertilization in response to continued carbon emissions and atmospheric CO₂ rise is highly uncertain. In this paper, the ramifications of CO₂ fertilization in simulations of future climate change are explored, using an intermediate complexity coupled climate–carbon model. It is shown that the absence of future CO₂ fertilization results in substantially higher future CO₂ levels in the atmosphere, as this removes the dominant contributor to future terrestrial carbon uptake in the model. As a result, climate changes are larger, though the radiative effect of higher CO₂ on surface temperatures in the model is offset by about 30% due to reduced positive dynamic vegetation feedbacks; that is, the removal of CO₂ fertilization results in less vegetation expansion in the model, which would otherwise constitute an important positive surface albedo‐temperature feedback. However, the effect of larger climate changes has other important implications for the carbon cycle – notably to further weaken remaining carbon sinks in the model. As a result, positive climate–carbon cycle feedbacks are larger when CO₂ fertilization is absent. This creates an interesting synergism of terrestrial carbon cycle feedbacks, whereby positive (climate–carbon cycle) feedbacks are amplified when a negative (CO₂ fertilization) feedback is removed.     

Carbon fixation and carbon availability in marine phytoplankton

It is widely believed that inorganic C does not limit the rate of short-term photosynthesis, the net productivity, or the maximum biomass, of marine phytoplankton. This lack of inorganic C restriction is less widely believed to hold for phytoplankton in many low alkalinity freshwaters or for seaweed in nutrient-enriched rock pools. These views are examined in the context of the physical chemistry of the inorganic C system in natural waters and of the ways in which various taxa of phytoplankton deal with inorganic C and discriminate between ¹²C and ¹³C. Using this information to interpret data obtained in the ocean or in freshwater suggests that short-term photosynthesis, production rate, and achieved biomass, of phytoplankton are rarely limited by inorganic C supply but, rather, that the widely suggested factors of limited light, nitrogen or phosphorus supply are the resource inputs which restrict productivity. Global change, by increasing atmospheric CO₂ partial pressure and global mean temperatures, is likely to increase the mean CO₂ concentration in the atmosphere, but the corresponding change in the oceans will be much less. There are, however, genotypic differences in the handling of inorganic C among the diversity of marine phytoplankton, and in impact on use of limiting nutrients, so increases in the mean CO₂ and HCO₃ ^- concentrations in surface ocean waters could cause changes in species composition. However, the rarity of inorganic C limitation of marine phytoplankton short-term photosynthesis, net productivity, or the maximum biomass, in today's ocean means that global change is unlikely to increase these three values in the ocean.     

Water use and yield of bioenergy poplar in future climates: modelling the interactive effects of elevated atmospheric CO2 and climate on productivity and water use

Vegetation exerts large control on global biogeochemical cycles through the processes of photosynthesis and transpiration that exchange CO₂ and water between the land and the atmosphere. Increasing atmospheric CO₂ concentrations exert direct effects on vegetation through enhanced photosynthesis and reduced stomatal conductance, and indirect effects through changes in climatic variables that drive these processes. How these direct and indirect CO₂ impacts interact with each other to affect plant productivity and water use has not been explicitly analysed and remains unclear, yet is important to fully understand the response of the global carbon cycle to future climate change. Here, we use a set of factorial modelling experiments to quantify the direct and indirect impacts of atmospheric CO₂ and their interaction on yield and water use in bioenergy short rotation coppice poplar, in addition to quantifying the impact of other environmental drivers such as soil type. We use the JULES land‐surface model forced with a ten‐member ensemble of projected climate change for 2100 with atmospheric CO₂ concentrations representative of the A1B emissions scenario. We show that the simulated response of plant productivity to future climate change was nonadditive in JULES, however this nonadditivity was not apparent for plant transpiration. The responses of both growth and transpiration under all experimental scenarios were highly variable between sites, highlighting the complexity of interactions between direct physiological CO₂ effects and indirect climate effects. As a result, no general pattern explaining the response of bioenergy poplar water use and yield to future climate change could be discerned across sites. This study suggests attempts to infer future climate change impacts on the land biosphere from studies that force with either the direct or indirect CO₂ effects in isolation from each other may lead to incorrect conclusions in terms of both the direction and magnitude of plant response to future climate change.     

Defoliation reduces soil biota – and modifies stimulating effects of elevated CO2

To understand the responses to external disturbance such as defoliation and possible feedback mechanisms at global change in terrestrial ecosystems, it is necessary to examine the extent and nature of effects on aboveground–belowground interactions. We studied a temperate heathland system subjected to experimental climate and atmospheric factors based on prognoses for year 2075 and further exposed to defoliation. By defoliating plants, we were able to study how global change modifies the interactions of the plant–soil system. Shoot production, root biomass, microbial biomass, and nematode abundance were assessed in the rhizosphere of manually defoliated patches of Deschampsia flexuosa in June in a full‐factorial FACE experiment with the treatments: increased atmospheric CO₂, increased nighttime temperatures, summer droughts, and all of their combinations. We found a negative effect of defoliation on microbial biomass that was not apparently affected by global change. The negative effect of defoliation cascades through to soil nematodes as dependent on CO₂ and drought. At ambient CO₂, drought and defoliation each reduced nematodes. In contrast, at elevated CO₂, a combination of drought and defoliation was needed to reduce nematodes. We found positive effects of CO₂ on root density and microbial biomass. Defoliation affected soil biota negatively, whereas elevated CO₂ stimulated the plant–soil system. This effect seen in June is contrasted by the effects seen in September at the same site. Late season defoliation increased activity and biomass of soil biota and more so at elevated CO₂. Based on soil biota responses, plants defoliated in active growth therefore conserve resources, whereas defoliation after termination of growth results in release of resources. This result challenges the idea that plants via exudation of organic carbon stimulate their rhizosphere biota when in apparent need of nutrients for growth.     

Modelling the recent historical impacts of atmospheric CO2 and climate change on Mediterranean vegetation

During the past century, annual mean temperature has increased by 0.75°C and precipitation has shown marked variation throughout the Mediterranean basin. These historical climate changes may have had significant, but presently undefined, impacts on the productivity and structure of sclerophyllous shrubland, an important vegetation type in the region. We used a vegetation model for this functional type to examine climate change impacts, and their interaction with the concurrent historical rise in atmospheric CO₂. Using only climate and soil texture as data inputs, model predictions showed good agreement with observations of seasonal and regional variation in leaf and canopy physiology, net primary productivity (NPP), leaf area index (LAI) and soil water. Model simulations for shrubland sites indicated that potential NPP has risen by 25% and LAI by 7% during the past century, although the absolute increase in LAI was small. Sensitivity analysis suggested that the increase in atmospheric CO₂ since 1900 was the primary cause of these changes, and that simulated climate change alone had negative impacts on both NPP and LAI. Effects of rising CO₂ were mediated by significant increases in the efficiency of water‐use in NPP throughout the region, as a consequence of the direct effect of CO₂ on leaf gas exchange. This increase in efficiency compensated for limitation of NPP by drought, except in areas where drought was most severe. However, while water was used more efficiently, total canopy water loss rose slightly or remained unaffected in model simulations, because increases in LAI with CO₂ counteracted the effects of reduced stomatal conductance on transpiration. Model simulations for the Mediterranean region indicate that the recent rise in atmospheric CO₂ may already have had significant impacts on productivity, structure and water relations of sclerophyllous shrub vegetation, which tended to offset the detrimental effects of climate change in the region.