Terrestrial biosphere model performance for inter‐annual variability of land‐atmosphere CO2 exchange

Interannual variability in biosphere‐atmosphere exchange of CO₂ is driven by a diverse range of biotic and abiotic factors. Replicating this variability thus represents the ‘acid test’ for terrestrial biosphere models. Although such models are commonly used to project responses to both normal and anomalous variability in climate, they are rarely tested explicitly against inter‐annual variability in observations. Herein, using standardized data from the North American Carbon Program, we assess the performance of 16 terrestrial biosphere models and 3 remote sensing products against long‐term measurements of biosphere‐atmosphere CO₂ exchange made with eddy‐covariance flux towers at 11 forested sites in North America. Instead of focusing on model‐data agreement we take a systematic, variability‐oriented approach and show that although the models tend to reproduce the mean magnitude of the observed annual flux variability, they fail to reproduce the timing. Large biases in modeled annual means are evident for all models. Observed interannual variability is found to commonly be on the order of magnitude of the mean fluxes. None of the models consistently reproduce observed interannual variability within measurement uncertainty. Underrepresentation of variability in spring phenology, soil thaw and snowpack melting, and difficulties in reproducing the lagged response to extreme climatic events are identified as systematic errors, common to all models included in this study.     

Water use of two Mojave Desert shrubs under elevated CO2

Plant responses to elevated atmospheric CO₂ have been characterized generally by stomatal closure and enhanced growth rates. These responses are being increasingly incorporated into global climate models that quantify interactions between the biosphere and atmosphere, altering climate predictions from simpler physically based models. However, current information on CO₂ responses has been gathered primarily from studies of crop and temperate forest species. In order to apply responses of vegetation to global predictions, CO₂ responses in other commonly occurring biomes must be studied. A Free Air CO₂ Enrichment (FACE) study is currently underway to examine plant responses to high CO₂ in a natural, undisturbed Mojave Desert ecosystem in Nevada, USA. Here we present findings from this study, and its companion glasshouse experiment, demonstrating that field‐grown Ephedra nevadensis and glasshouse‐grown Larrea tridentata responded to high CO₂ with reductions in the ratio of transpirational surface area to sapwood area (LSR) of 33% and 60%, respectively. Thus, leaf‐specific hydraulic conductivity increased and stomatal conductance remained constant or was increased under elevated CO₂. Field‐grown Larrea did not show a reduced LSR under high CO₂, and stomatal conductance was reduced in the high CO₂ treatment, although the effect was apparent only under conditions of unusually high soil moisture. Both findings suggest that the common paradigm of 20–50% reductions in stomatal conductance under high CO₂ may not be applicable to arid ecosystems under most conditions.     

Pre‐dawn stomatal opening does not substantially enhance early‐morning photosynthesis in Helianthus annuus

Most C₃ plant species have partially open stomata during the night especially in the 3–5 h before dawn. This pre‐dawn stomatal opening has been hypothesized to enhance early‐morning photosynthesis (A) by reducing diffusion limitations to CO₂ at dawn. We tested this hypothesis in cultivated Helianthus annuus using whole‐shoot gas exchange, leaf level gas exchange and modelling approaches. One hour pre‐dawn low‐humidity treatments were used to reduce pre‐dawn stomatal conductance (g). At the whole‐shoot level, a difference of pre‐dawn g (0.40 versus 0.17 mol m^?2 s^?1) did not significantly affect A during the first hour after dawn. Shorter term effects were investigated with leaf level gas exchange measurements and a difference of pre‐dawn g (0.10 versus 0.04 mol m^?2 s^?1) affected g and A for only 5 min after dawn. The potential effects of a wider range of stomatal apertures were explored with an empirical model of the relationship between A and intercellular CO₂ concentration during the half‐hour after dawn. Modelling results demonstrated that even extremely low pre‐dawn stomatal conductance values have only a minimal effect on early‐morning A for a few minutes after dawn. Thus, we found no evidence that pre‐dawn stomatal opening enhances A.     

A stomatal optimization theory to describe the effects of atmospheric CO2 on leaf photosynthesis and transpiration

Background and Aims

Global climate models predict decreases in leaf stomatal conductance and transpiration due to increases in atmospheric CO₂. The consequences of these reductions are increases in soil moisture availability and continental scale run-off at decadal time-scales. Thus, a theory explaining the differential sensitivity of stomata to changing atmospheric CO₂ and other environmental conditions must be identified. Here, these responses are investigated using optimality theory applied to stomatal conductance.

Methods

An analytical model for stomatal conductance is proposed based on: (a) Fickian mass transfer of CO₂ and H₂O through stomata; (b) a biochemical photosynthesis model that relates intercellular CO₂ to net photosynthesis; and (c) a stomatal model based on optimization for maximizing carbon gains when water losses represent a cost. Comparisons between the optimization-based model and empirical relationships widely used in climate models were made using an extensive gas exchange dataset collected in a maturing pine (Pinus taeda) forest under ambient and enriched atmospheric CO₂.

Key Results and Conclusion

In this interpretation, it is proposed that an individual leaf optimally and autonomously regulates stomatal opening on short-term (approx. 10-min time-scale) rather than on daily or longer time-scales. The derived equations are analytical with explicit expressions for conductance, photosynthesis and intercellular CO₂, thereby making the approach useful for climate models. Using a gas exchange dataset collected in a pine forest, it is shown that (a) the cost of unit water loss λ (a measure of marginal water-use efficiency) increases with atmospheric CO₂; (b) the new formulation correctly predicts the condition under which CO₂-enriched atmosphere will cause increasing assimilation and decreasing stomatal conductance.     

Guard cell photosynthesis is critical for stomatal turgor production,yet does not directly mediate CO2‐ and ABA‐induced stomatal closing

Stomata mediate gas exchange between the inter‐cellular spaces of leaves and the atmosphere. CO₂ levels in leaves (Ci) are determined by respiration, photosynthesis, stomatal conductance and atmospheric [CO₂]. [CO₂] in leaves mediates stomatal movements. The role of guard cell photosynthesis in stomatal conductance responses is a matter of debate, and genetic approaches are needed. We have generated transgenic Arabidopsis plants that are chlorophyll‐deficient in guard cells only, expressing a constitutively active chlorophyllase in a guard cell specific enhancer trap line. Our data show that more than 90% of guard cells were chlorophyll‐deficient. Interestingly, approximately 45% of stomata had an unusual, previously not‐described, morphology of thin‐shaped chlorophyll‐less stomata. Nevertheless, stomatal size, stomatal index, plant morphology, and whole‐leaf photosynthetic parameters (PSII, qP, qN, F_V′/F_M′) were comparable with wild‐type plants. Time‐resolved intact leaf gas‐exchange analyses showed a reduction in stomatal conductance and CO₂‐assimilation rates of the transgenic plants. Normalization of CO₂ responses showed that stomata of transgenic plants respond to [CO₂] shifts. Detailed stomatal aperture measurements of normal kidney‐shaped stomata, which lack chlorophyll, showed stomatal closing responses to [CO₂] elevation and abscisic acid (ABA), while thin‐shaped stomata were continuously closed. Our present findings show that stomatal movement responses to [CO₂] and ABA are functional in guard cells that lack chlorophyll. These data suggest that guard cell CO₂ and ABA signal transduction are not directly modulated by guard cell photosynthesis/electron transport. Moreover, the finding that chlorophyll‐less stomata cause a ‘deflated’ thin‐shaped phenotype, suggests that photosynthesis in guard cells is critical for energization and guard cell turgor production.     

Photosynthesis and isoprene emission from trees along an urban–rural gradient in Texas

Isoprene emission is an important mechanism for improving the thermotolerance of plant photosystems as temperatures increase. In this study, we measured photosynthesis and isoprene emission in trees along an urban–rural gradient that serves as a proxy for climate change, to understand daily and seasonal responses to changes in temperature and other environmental variables. Leaf‐level gas exchange and basal isoprene emission of post oak (Quercus stellata) and sweet gum (Liquidambar styraciflua) were recorded at regular intervals over an entire growing season at urban, suburban, and rural sites in eastern Texas. In addition, the temperature and atmospheric carbon dioxide concentration experienced by leaves were experimentally manipulated in spring, early summer, and late summer. We found that trees experienced lower stomatal conductance and photosynthesis and higher isoprene emission, at the urban and suburban sites compared to the rural site. Path analysis indicated a daily positive effect of isoprene emission on photosynthesis, but unexpectedly, higher isoprene emission from urban trees was not associated with improved photosynthesis as temperatures increased during the growing season. Furthermore, urban trees experienced relatively higher isoprene emission at high CO₂ concentrations, while isoprene emission was suppressed at the other sites. These results suggest that isoprene emission may be less beneficial in urban, and potentially future, environmental conditions, particularly if higher temperatures override the suppressive effects of high CO₂ on isoprene emission. These are important considerations for modeling future biosphere–atmosphere interactions and for understanding tree physiological responses to climate change.     

Canopy‐scale relationships between stomatal conductance and photosynthesis in irrigated rice

Modeling stomatal behavior is critical in research on land–atmosphere interactions and climate change. The most common model uses an existing relationship between photosynthesis and stomatal conductance. However, its parameters have been determined using infrequent and leaf‐scale gas‐exchange measurements and may not be representative of the whole canopy in time and space. In this study, we used a top‐down approach based on a double‐source canopy model and eddy flux measurements throughout the growing season. Using this approach, we quantified the canopy‐scale relationship between gross photosynthesis and stomatal conductance for 3 years and their relationships with leaf nitrogen content throughout each growing season above a paddy rice canopy in Japan. The canopy‐averaged stomatal conductance (g_sc) increased with increasing gross photosynthesis per unit green leaf area (A_g), as was the case with leaf‐scale measurements, and 41–90% of its variation was explained by variations in A_g adjusted to account for the leaf‐to‐air vapor‐pressure deficit and CO₂ concentration using the Leuning model. The slope (m) in this model (g_sc versus the adjusted A_g) was almost constant within a 15‐day period, but changed seasonally. The m values determined using an ensemble dataset for two mid‐growing‐season 15‐day periods were 30.8 (SE = 0.5), 29.9 (SE = 0.7), and 29.9 (SE = 0.6) in 2004, 2005, and 2006, respectively; the overall mid‐season value was 30.3 and did not greatly differ among the 3 years. However, m appeared to be higher during the early and late growing seasons. The ontogenic changes in leaf nitrogen content strongly affected A_g and thus g_sc. In addition, we have discussed the agronomic impacts of the interactions between leaf nitrogen content and g_sc. Despite limitations in the observations and modeling, our canopy‐scale results emphasize the importance of continuous, season‐long estimates of stomatal model parameters for crops using top‐down approaches.     

Stomatal function,density and pattern,and CO2 assimilation in Arabidopsis thaliana tmm1 and sdd1‐1 mutants

• Stomata modulate the exchange of water and CO₂ between plant and atmosphere. Although stomatal density is known to affect CO₂ diffusion into the leaf and thus photosynthetic rate, the effect of stomatal density and patterning on CO₂ assimilation is not fully understood.
• We used wild types Col‐0 and C24 and stomatal mutants sdd1‐1 and tmm1 of Arabidopsis thaliana, differing in stomatal density and pattern, to study the effects of these variations on both stomatal and mesophyll conductance and CO₂ assimilation rate. Anatomical parameters of stomata, leaf temperature and carbon isotope discrimination were also assessed.
• Our results indicate that increased stomatal density enhanced stomatal conductance in sdd1‐1 plants, with no effect on photosynthesis, due to both unchanged photosynthetic capacity and decreased mesophyll conductance. Clustering (abnormal patterning formed by clusters of two or more stomata) and a highly unequal distribution of stomata between the adaxial and abaxial leaf sides in tmm1 mutants also had no effect on photosynthesis.
• Except at very high stomatal densities, stomatal conductance and water loss were proportional to stomatal density. Stomatal formation in clusters reduced stomatal dynamics and their operational range as well as the efficiency of CO₂ transport.

     

Towards physiologically meaningful water‐use efficiency estimates from eddy covariance data

Intrinsic water‐use efficiency (iWUE) characterizes the physiological control on the simultaneous exchange of water and carbon dioxide in terrestrial ecosystems. Knowledge of iWUE is commonly gained from leaf‐level gas exchange measurements, which are inevitably restricted in their spatial and temporal coverage. Flux measurements based on the eddy covariance (EC) technique can overcome these limitations, as they provide continuous and long‐term records of carbon and water fluxes at the ecosystem scale. However, vegetation gas exchange parameters derived from EC data are subject to scale‐dependent and method‐specific uncertainties that compromise their ecophysiological interpretation as well as their comparability among ecosystems and across spatial scales. Here, we use estimates of canopy conductance and gross primary productivity (GPP) derived from EC data to calculate a measure of iWUE (G₁, “stomatal slope”) at the ecosystem level at six sites comprising tropical, Mediterranean, temperate, and boreal forests. We assess the following six mechanisms potentially causing discrepancies between leaf and ecosystem‐level estimates of G₁: (i) non‐transpirational water fluxes; (ii) aerodynamic conductance; (iii) meteorological deviations between measurement height and canopy surface; (iv) energy balance non‐closure; (v) uncertainties in net ecosystem exchange partitioning; and (vi) physiological within‐canopy gradients. Our results demonstrate that an unclosed energy balance caused the largest uncertainties, in particular if it was associated with erroneous latent heat flux estimates. The effect of aerodynamic conductance on G₁ was sufficiently captured with a simple representation. G₁ was found to be less sensitive to meteorological deviations between canopy surface and measurement height and, given that data are appropriately filtered, to non‐transpirational water fluxes. Uncertainties in the derived GPP and physiological within‐canopy gradients and their implications for parameter estimates at leaf and ecosystem level are discussed. Our results highlight the importance of adequately considering the sources of uncertainty outlined here when EC‐derived water‐use efficiency is interpreted in an ecophysiological context.     

Future carbon dioxide concentration decreases canopy evapotranspiration and soil water depletion by field‐grown maize

Maize, in rotation with soybean, forms the largest continuous ecosystem in temperate North America, therefore changes to the biosphere‐atmosphere exchange of water vapor and energy of these crops are likely to have an impact on the Midwestern US climate and hydrological cycle. As a C₄ crop, maize photosynthesis is already CO₂‐saturated at current CO₂ concentrations ([CO₂]) and the primary response of maize to elevated [CO₂] is decreased stomatal conductance (g_s). If maize photosynthesis is not stimulated in elevated [CO₂], then reduced g_s is not offset by greater canopy leaf area, which could potentially result in a greater ET reduction relative to that previously reported in soybean, a C₃ species. The objective of this study is to quantify the impact of elevated [CO₂] on canopy energy and water fluxes of maize (Zea mays). Maize was grown under ambient and elevated [CO₂] (550 μmol mol^?1 during 2004 and 2006 and 585 μmol mol^?1 during 2010) using Free Air Concentration Enrichment (FACE) technology at the SoyFACE facility in Urbana, Illinois. Maize ET was determined using a residual energy balance approach based on measurements of sensible (H) and soil heat fluxes, and net radiation. Relative to control, elevated [CO₂] decreased maize ET (7–11%; P < 0.01) along with lesser soil moisture depletion, while H increased (25–30 W m^?2; P < 0.01) along with higher canopy temperature (0.5–0.6 °C). This reduction in maize ET in elevated [CO₂] is approximately half that previously reported for soybean. A partitioning analysis showed that transpiration contributed less to total ET for maize compared to soybean, indicating a smaller role of stomata in dictating the ET response to elevated [CO₂]. Nonetheless, both maize and soybean had significantly decreased ET and increased H, highlighting the critical role of elevated [CO₂] in altering future hydrology and climate of the region that is extensively cropped with these species.     

State of the science in reconciling top‐down and bottom‐up approaches for terrestrial CO2 budget

Robust estimates of CO₂ budget, CO₂ exchanged between the atmosphere and terrestrial biosphere, are necessary to better understand the role of the terrestrial biosphere in mitigating anthropogenic CO₂ emissions. Over the past decade, this field of research has advanced through understanding of the differences and similarities of two fundamentally different approaches: “top‐down” atmospheric inversions and “bottom‐up” biosphere models. Since the first studies were undertaken, these approaches have shown an increasing level of agreement, but disagreements in some regions still persist, in part because they do not estimate the same quantity of atmosphere–biosphere CO₂ exchange. Here, we conducted a thorough comparison of CO₂ budgets at multiple scales and from multiple methods to assess the current state of the science in estimating CO₂ budgets. Our set of atmospheric inversions and biosphere models, which were adjusted for a consistent flux definition, showed a high level of agreement for global and hemispheric CO₂ budgets in the 2000s. Regionally, improved agreement in CO₂ budgets was notable for North America and Southeast Asia. However, large gaps between the two methods remained in East Asia and South America. In other regions, Europe, boreal Asia, Africa, South Asia, and Oceania, it was difficult to determine whether those regions act as a net sink or source because of the large spread in estimates from atmospheric inversions. These results highlight two research directions to improve the robustness of CO₂ budgets: (a) to increase representation of processes in biosphere models that could contribute to fill the budget gaps, such as forest regrowth and forest degradation; and (b) to reduce sink–source compensation between regions (dipoles) in atmospheric inversion so that their estimates become more comparable. Advancements on both research areas will increase the level of agreement between the top‐down and bottom‐up approaches and yield more robust knowledge of regional CO₂ budgets.     

Spatial variability and temporal trends in water‐use efficiency of European forests

The increasing carbon dioxide (CO₂) concentration in the atmosphere in combination with climatic changes throughout the last century are likely to have had a profound effect on the physiology of trees: altering the carbon and water fluxes passing through the stomatal pores. However, the magnitude and spatial patterns of such changes in natural forests remain highly uncertain. Here, stable carbon isotope ratios from a network of 35 tree‐ring sites located across Europe are investigated to determine the intrinsic water‐use efficiency (iWUE), the ratio of photosynthesis to stomatal conductance from 1901 to 2000. The results were compared with simulations of a dynamic vegetation model (LPX‐Bern 1.0) that integrates numerous ecosystem and land–atmosphere exchange processes in a theoretical framework. The spatial pattern of tree‐ring derived iWUE of the investigated coniferous and deciduous species and the model results agreed significantly with a clear south‐to‐north gradient, as well as a general increase in iWUE over the 20th century. The magnitude of the iWUE increase was not spatially uniform, with the strongest increase observed and modelled for temperate forests in Central Europe, a region where summer soil‐water availability decreased over the last century. We were able to demonstrate that the combined effects of increasing CO₂ and climate change leading to soil drying have resulted in an accelerated increase in iWUE. These findings will help to reduce uncertainties in the land surface schemes of global climate models, where vegetation–climate feedbacks are currently still poorly constrained by observational data.     

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.     

Modelling the soil-plant-atmosphere continuum in a Quercus–Acer stand at Harvard Forest: the regulation of stomatal conductance by light, nitrogen and soil/plant hydraulic properties

Our objective is to describe a multi-layer model of C₃-canopy processes that effectively simulates hourly CO₂ and latent energy (LE) fluxes in a mixed deciduous Quercus-Acer (oak–maple) stand in central Massachusetts, USA. The key hypothesis governing the biological component of the model is that stomatal conductance (g_s) is varied so that daily carbon uptake per unit of foliar nitrogen is maximized within the limitations of canopy water availability. The hydraulic system is modelled as an analogue to simple electrical circuits in parallel, including a separate soil hydraulic resistance, plant resistance and plant capacitance for each canopy layer. Stomatal opening is initially controlled to conserve plant water stores and delay the onset of water stress. Stomatal closure at a threshold minimum leaf water potential prevents xylem cavitation and controls the maximum rate of water flux through the hydraulic system. We show a strong correlation between predicted hourly CO₂ exchange rate (r²= 0.86) and LE (r²= 0.87) with independent whole-forest measurements made by the eddy correlation method during the summer of 1992. Our theoretical derivation shows that observed relationships between CO₂ assimilation and LE flux can be explained on the basis of stomatal behaviour optimizing carbon gain, and provides an explicit link between canopy structure, soil properties, atmospheric conditions and stomatal conductance.     

Stomatal action directly feeds back on leaf turgor: new insights into the regulation of the plant water status from non‐invasive pressure probe measurements

Uptake of CO₂ by the leaf is associated with loss of water. Control of stomatal aperture by volume changes of guard cell pairs optimizes the efficiency of water use. Under water stress, the protein kinase OPEN STOMATA 1 (OST1) activates the guard‐cell anion release channel SLOW ANION CHANNEL‐ASSOCIATED 1 (SLAC1), and thereby triggers stomatal closure. Plants with mutated OST1 and SLAC1 are defective in guard‐cell turgor regulation. To study the effect of stomatal movement on leaf turgor using intact leaves of Arabidopsis, we used a new pressure probe to monitor transpiration and turgor pressure simultaneously and non‐invasively. This probe permits routine easy access to parameters related to water status and stomatal conductance under physiological conditions using the model plant Arabidopsis thaliana. Long‐term leaf turgor pressure recordings over several weeks showed a drop in turgor during the day and recovery at night. Thus pressure changes directly correlated with the degree of plant transpiration. Leaf turgor of wild‐type plants responded to CO₂, light, humidity, ozone and abscisic acid (ABA) in a guard cell‐specific manner. Pressure probe measurements of mutants lacking OST1 and SLAC1 function indicated impairment in stomatal responses to light and humidity. In contrast to wild‐type plants, leaves from well‐watered ost1 plants exposed to a dry atmosphere wilted after light‐induced stomatal opening. Experiments with open stomata mutants indicated that the hydraulic conductance of leaf stomata is higher than that of the root–shoot continuum. Thus leaf turgor appears to rely to a large extent on the anion channel activity of autonomously regulated stomatal guard cells.     

High precision measurements of atmospheric concentrations and plant exchange rates of carbonyl sulfide using mid‐IR quantum cascade laser

Linking measurements of carbonyl sulfide (COS) with those of carbon dioxide (CO₂) has a potential in providing a powerful tracer of gross CO₂ fluxes between the atmosphere and land plants, a critical element in understanding the response of the land biosphere to global change. A new application of online COS, CO₂ and water vapor measurements based on newly designed mid‐infrared (IR) dual quantum cascade laser (QCL) spectrometer measures COS and CO₂ (at 2056 cm^?1) and water vapor (at 2190 cm^?1), with detectors cooled thermoelectrically (at ?43 °C) or with liquid nitrogen (?197 °C) for improved precision. Using the cryogenic detectors with averaging time of 1 s, precision was 50 pmol mol^?1, 0.4 μmol mol^?1 and 0.01 mmol mol^?1 for COS, CO₂ and water vapor, respectively (14, 0.2 and 0.003, respectively, for 60 s averaging time). We measured COS concentrations in ambient air, and changes in the rates of COS, CO₂ and water vapor exchange of attached leaves in response to changes in light intensity and ambient COS concentrations. The results were consistent with those of nononline gas chromatography–mass spectrometry for COS and IR gas analyzer for CO₂ and water vapor, with a high linear correlation for a broad range of concentrations (R²= 0.85 for COS and R²= 1.00 for CO₂ and water vapor). The new methodology opens the way for lab and field explorations of COS fluxes as a powerful new tracer for CO₂ exchange in the land biosphere.     

Short-chained oxygenated VOC emissions in <Emphasis Type="Italic">Pinus halepensis</Emphasis> in response to changes in water availability

Short-chained oxygenated VOC (oxVOCs) emissions from Pinus halepensis saplings were monitored in response to changes in water availability. Online measurements were made with a proton transfer reaction—mass spectrometer under controlled conditions, together with CO₂ and H₂O exchange measurements. Masses corresponding to methanol and acetone were the most emitted oxVOCs. All the oxVOC exchanges, except that of acetone (M59), were significantly related to stomatal conductance and transpiration. Acetaldehyde (M45) emission showed, moreover, a strong dependence on the concentration of acetaldehyde in the ambient: stomatal opening (stomatal conductance above 75 mmol m⁻² s⁻¹) only allowed increased emissions when external concentration were below 6 ppb. Acetone (M59) presented an important peak of emission following light and stomatal opening in the morning when plants were water stressed. Thus, the alterations in oxVOC emissions in P. halepensis caused by the water deficit seem to be mainly driven by water stress effect on stomatal closure and oxVOC air concentrations.     

Impact of CO2 concentration changes on the biosphere-atmosphere system of West Africa

Vegetation dynamics plays a critical role in causing the decadal variability of precipitation over the Sahel region of West Africa. However, the potential impact of changes in CO₂ concentration on vegetation dynamics and precipitation variability of this region has not been addressed by previous studies. In this paper, we explore the role of CO₂ concentration in the regional climate system of West Africa using a zonally symmetric, synchronously coupled biosphere‐atmosphere model. We first document the response of precipitation and vegetation to incremental changes of CO₂ concentration; the impact of CO₂ concentration on the variability of the regional biosphere‐atmosphere system is then addressed using the second half of the twentieth century as an example. An increase of CO₂ concentration causes the regional biosphere‐atmosphere system to become wetter and greener, with the radiative effect of CO₂ and improved plant‐water relation dominant in the Sahelian grassland region and the direct enhancement of leaf carbon assimilation dominant in the tree‐covered region to the south. Driven by the observed sea surface temperature (SST) of the tropical Atlantic Ocean during the period 1950–97 and with CO₂ concentration prescribed at a pre‐industrial level 300ppmv, the model simulates a persistent Sahel drought during the period of 1960s?1990s. The simulated drought takes place in the form of a transition of the coupled biosphere‐atmosphere system from a wet/green regime in the 1950s to a dry/barren regime after the 1960s. This climate transition is triggered by SST forcing and materialized through vegetation‐climate interactions. The same SST forcing does not produce such a persistent drought when a constant modern CO₂ concentration of 350ppmv is specified, indicating that the biosphere‐atmosphere system at higher CO₂ level is more resilient to drought‐inducing external forcings. This finding suggests that the regional climate in Sahel, which tends to alternate between dry and wet spells, may experience longer (or more frequent) wet episodes and shorter (or less frequent) dry episodes in the future than in the past. Our study has significant implications regarding the impact of climate change on regional socio‐economic development.     

CO2-forced evolution of plant gas exchange capacity and water-use efficiency over the Phanerozoic

The capacity of plants to fix carbon is ultimately constrained by two core plant attributes: photosynthetic biochemistry and the conductance to CO₂ diffusion from the atmosphere to sites of carboxylation in chloroplasts, predominantly stomatal conductance. Analysis of fossilized plant remains shows that stomatal density (number per unit area, D) and size (length by width, S) have fluctuated widely over the Phanerozoic Eon, indicating changes in maximum stomatal conductance. Parallel changes are likely to have taken place in leaf photosynthetic biochemistry, of which maximal rubisco carboxylation rate, V_cmax is a central element. We used measurements of S and D from fossilized plant remains spanning the last 400 Myr (most of the Phanerozoic), together with leaf gas exchange data and modeled Phanerozoic trends in atmospheric CO₂ concentration, [CO₂]_a, to calibrate a [CO₂]_a‐driven model of the long‐term environmental influences on S, D and V_cmax. We show that over the Phanerozoic large changes in [CO₂]_a forced S, D and V_cmax to co‐vary so as to reduce the impact of the change in [CO₂]_a on leaf CO₂ assimilation for minimal energetic cost and reduced nitrogen requirements. Underlying this is a general negative correlation between S and D, and a positive correlation between water‐use efficiency and [CO₂]_a. Furthermore, the calculated steady rise in stomatal conductance over the Phanerozoic is consistent with independent evidence for the evolution of plant hydraulic capacity, implying coordinated and sustained increase in gas exchange capacity and hydraulic capacity parallel long‐term increases in land plant diversity.     

Physiological acclimation dampens initial effects of elevated temperature and atmospheric CO2 concentration in mature boreal Norway spruce

Physiological processes of terrestrial plants regulate the land–atmosphere exchange of carbon, water, and energy, yet few studies have explored the acclimation responses of mature boreal conifer trees to climate change. Here we explored the acclimation responses of photosynthesis, respiration, and stomatal conductance to elevated temperature and/or CO₂ concentration ([CO₂]) in a 3‐year field experiment with mature boreal Norway spruce. We found that elevated [CO₂] decreased photosynthetic carboxylation capacity (?23% at 25 °C) and increased shoot respiration (+64% at 15 °C), while warming had no significant effects. Shoot respiration, but not photosynthetic capacity, exhibited seasonal acclimation. Stomatal conductance at light saturation and a vapour pressure deficit of 1 kPa was unaffected by elevated [CO₂] but significantly decreased (?27%) by warming, and the ratio of intercellular to ambient [CO₂] was enhanced (+17%) by elevated [CO₂] and decreased (?12%) by warming. Many of these responses differ from those typically observed in temperate tree species. Our results show that long‐term physiological acclimation dampens the initial stimulation of plant net carbon assimilation to elevated [CO₂], and of plant water use to warming. Models that do not account for these responses may thus overestimate the impacts of climate change on future boreal vegetation–atmosphere interactions.