Seasonal dynamics and punctuated carbon sink reduction suggest photosynthetic capacity of boreal silver birch is reduced by the accumulation of hexose.

The 'assimilates inhibition hypothesis' posits that accumulation of nonstructural carbohydrates (NSCs) in leaves reduces leaf net photosynthetic rate, thus internally regulating photosynthesis. Experimental work provides equivocal support mostly under controlled conditions without identifying a particular NSC as involved in the regulation. We combined 3-yr in situ leaf gas exchange observations (natural dynamics) in the upper crown of mature Betula pendula simultaneously with measurements of concentrations of sucrose, hexoses (glucose and fructose), and starch, and similar measurements during several one-day shoot girdling (perturbation dynamics). Leaf water potential and water and nitrogen content were measured to account for their possible contribution to photosynthesis regulation. Leaf photosynthetic capacity (A/Ci) was temporally negatively correlated with NSC accumulation under both natural and perturbation states. For developed leaves, leaf hexose concentration explained A/Ci variation better than environmental variables (temperature history and daylength); the opposite was observed for developing leaves. The weaker correlations between NSCs and A/Ci in developing leaves may reflect their strong internal sink strength for carbohydrates. By contrast, the strong decline in photosynthetic capacity with NSCs accumulation in mature leaves, observed most clearly with hexose, and even more tightly with its constituents, provides support for the role of assimilates in regulating photosynthesis under natural conditions.

Partitioning seasonal stem carbon dioxide efflux into stem respiration, bark photosynthesis and transport-related flux.

Stem CO2 efflux is an important component of the carbon balance in forests. The efflux is considered to principally reflect the net result of two dominating and opposing processes: stem respiration and stem photosynthesis. In addition, transport of CO2 in xylem sap is thought to play an appreciable role in affecting the net flux. This work presents an approach to partition stem CO2 efflux among these processes using sap-flux data and CO2-exchange measurements from dark and transparent chambers placed on mature Scots pine (Pinus sylvestris) trees. Seasonal changes and monthly parameters describing the studied processes were determined. Respiration contributed most to stem net CO2 flux, reaching up to 79% (considering the sum of the absolute values of stem respiration, stem photosynthesis and flux from CO2 transported in xylem sap to be 100%) in June, when stem growth was greatest. Photosynthesis contribution accounted for up to 13 % of the stem net CO2 flux, increasing over the monitoring period. CO2 transported axially with sap flow, decreased towards the end of the growing season. At a reference temperature, respiration decreased starting around midsummer, while its temperature sensitivity increased during the summer. A decline was observed for photosynthetic quantum yield around midsummer together with decreasing light-saturation point. The proposed approach facilitates modeling net stem CO2 flux at a range of time scales.

Hyposensitive canopy conductance renders ecosystems vulnerable to meteorological droughts.

Increased meteorological drought intensity with rising atmospheric demand for water (hereafter vapor pressure deficit [VPD]) increases the risk of tree mortality and ecosystem dysfunction worldwide. Ecosystem-scale water-use strategy is increasingly recognized as a key factor in regulating drought-related ecosystem responses. However, the link between water-use strategy and ecosystem vulnerability to meteorological droughts is poorly established. Using the global flux observations, historic hydroclimatic data, remote-sensing products, and plant functional-trait archive, we identified potentially vulnerable ecosystems, examining how ecosystem water-use strategy, quantified by the percentage bias (δ) of the empirical canopy conductance sensitivity to VPD relative to the theoretical value, mediated ecosystem responses to droughts. We found that prevailing soil water availability substantially impacted δ in dryland regions where ecosystems with insufficient soil moisture usually showed conservative water-use strategy, while ecosystems in humid regions exhibited more pronounced climatic adaptability. Hyposensitive and hypersensitive ecosystems, classified based on δ falling below or above the theoretical sensitivity, respectively, achieved similar net ecosystem productivity during droughts, employing different structural and functional strategies. However, hyposensitive ecosystems, risking their hydraulic system with a permissive water-use strategy, were unable to recover from droughts as quickly as hypersensitive ones. Our findings highlight that processed-based models predicting current functions and future performance of vegetation should account for the greater vulnerability of hyposensitive ecosystems to intensifying atmospheric and soil droughts.

The response of coarse root biomass to long-term CO2 enrichment and nitrogen application in a maturing Pinus taeda stand with a large broadleaved component.

Elevated atmospheric CO2 (eCO2 ) typically increases aboveground growth in both growth chamber and free-air carbon enrichment (FACE) studies. Here we report on the impacts of eCO2 and nitrogen amendment on coarse root biomass and net primary productivity (NPP) at the Duke FACE study, where half of the eight plots in a 30-year-old loblolly pine (Pinus taeda, L.) plantation, including competing naturally regenerated broadleaved species, were subjected to eCO2 (ambient, aCO2 plus 200 ppm) for 15-17 years, combined with annual nitrogen amendments (11.2 g N m-2 ) for 6 years. Allometric equations were developed following harvest to estimate coarse root (>2 mm diameter) biomass. Pine root biomass under eCO2 increased 32%, 1.80 kg m-2 above the 5.66 kg m-2 observed in aCO2 , largely accumulating in the top 30 cm of soil. In contrast, eCO2 increased broadleaved root biomass more than twofold (aCO2 : 0.81, eCO2 : 2.07 kg m-2 ), primarily accumulating in the 30-60 cm soil depth. Combined, pine and broadleaved root biomass increased 3.08 kg m-2 over aCO2 of 6.46 kg m-2 , a 48% increase. Elevated CO2 did not increase pine root:shoot ratio (average 0.24) but increased the ratio from 0.57 to 1.12 in broadleaved species. Averaged over the study (1997-2010), eCO2 increased pine, broadleaved and total coarse root NPP by 49%, 373% and 86% respectively. Nitrogen amendment had smaller effects on any component, singly or interacting with eCO2 . A sustained increase in root NPP under eCO2 over the study period indicates that soil nutrients were sufficient to maintain root growth response to eCO2 . These responses must be considered in computing coarse root carbon sequestration of the extensive southern pine and similar forests, and in modelling the responses of coarse root biomass of pine-broadleaved forests to CO2 concentration over a range of soil N availability.

How tree species, tree size, and topographical location influenced tree transpiration in northern boreal forests during the historic 2018 drought.

Trees in northern latitude ecosystems are projected to experience increasing drought stress as a result of rising air temperatures and changes in precipitation patterns in northern latitude ecosystems. However, most drought-related studies on high-latitude boreal forests (>50°N) have been conducted in North America, with few studies quantifying the response in European and Eurasian boreal forests. Here, we tested how daily whole-tree transpiration (Q, Liters day-1 ) and Q normalized for mean daytime vapor pressure deficit (QDZ , Liters day-1 kPa-1 ) were affected by the historic 2018 drought in Europe. More specifically, we examined how tree species, size, and topographic position affected drought response in high-latitude mature boreal forest trees. We monitored 30 Pinus sylvestris (pine) and 30 Picea abies (spruce) trees distributed across a topographic gradient in northern Sweden. In general, pine showed a greater QDZ control compared to spruce during periods of severe drought (standardized precipitation-evapotranspiration index: SPEI < -1.5), suggesting that the latter are more sensitive to drought. Overall, QDZ reductions (using non-drought QDZ as reference) were less pronounced in larger trees during severe drought, but there was a species-specific pattern: QDZ reductions were greater in pine trees at high elevations and greater in spruce trees at lower elevations. Despite lower QDZ during severe drought, drought spells were interspersed with small precipitation events and overcast conditions, and QDZ returned to pre-drought conditions relatively quickly. This study highlights unique species-specific responses to drought, which are additionally driven by a codependent interaction among tree size, relative topographic position, and unique regional climate conditions.

Estimating canopy gross primary production by combining phloem stable isotopes with canopy and mesophyll conductances.

Gross primary production (GPP) is a key component of the forest carbon cycle. However, our knowledge of GPP at the stand scale remains uncertain, because estimates derived from eddy covariance (EC) rely on semi-empirical modelling and the assumptions of the EC technique are sometimes not fully met. We propose using the sap flux/isotope method as an alternative way to estimate canopy GPP, termed GPPiso/SF , at the stand scale and at daily resolution. It is based on canopy conductance inferred from sap flux and intrinsic water-use efficiency estimated from the stable carbon isotope composition of phloem contents. The GPPiso/SF estimate was further corrected for seasonal variations in photosynthetic capacity and mesophyll conductance. We compared our estimate of GPPiso/SF to the GPP derived from PRELES, a model parameterized with EC data. The comparisons were performed in a highly instrumented, boreal Scots pine forest in northern Sweden, including a nitrogen fertilized and a reference plot. The resulting annual and daily GPPiso/SF estimates agreed well with PRELES, in the fertilized plot and the reference plot. We discuss the GPPiso/SF method as an alternative which can be widely applied without terrain restrictions, where the assumptions of EC are not met.

Anatomical changes with needle length are correlated with leaf structural and physiological traits across five Pinus species.

The genus Pinus has wide geographical range and includes species that are the most economically valued among forest trees worldwide. Pine needle length varies greatly among species, but the effects of needle length on anatomy, function, and coordination and trade-offs among traits are poorly understood. We examined variation in leaf morphological, anatomical, mechanical, chemical, and physiological characteristics among five southern pine species: Pinus echinata, Pinus elliottii, Pinus palustris, Pinus taeda, and Pinus virginiana. We found that increasing needle length contributed to a trade-off between the relative fractions of support versus photosynthetic tissue (mesophyll) across species. From the shortest (7 cm) to the longest (36 cm) needles, mechanical tissue fraction increased by 50%, whereas needle dry density decreased by 21%, revealing multiple adjustments to a greater need for mechanical support in longer needles. We also found a fourfold increase in leaf hydraulic conductance over the range of needle length across species, associated with weaker upward trends in stomatal conductance and photosynthetic capacity. Our results suggest that the leaf size strongly influences their anatomical traits, which, in turn, are reflected in leaf mechanical support and physiological capacity.

Mechanisms for minimizing height-related stomatal conductance declines in tall vines.

The ability to transport water through tall stems hydraulically limits stomatal conductance (gs ), thereby constraining photosynthesis and growth. However, some plants are able to minimize this height-related decrease in gs , regardless of path length. We hypothesized that kudzu (Pueraria lobata) prevents strong declines in gs with height through appreciable structural and hydraulic compensative alterations. We observed only a 12% decline in maximum gs along 15-m-long stems and were able to model this empirical trend. Increasing resistance with transport distance was not compensated by increasing sapwood-to-leaf-area ratio. Compensating for increasing leaf area by adjusting the driving force would require water potential reaching -1.9 MPa, far below the wilting point (-1.2 MPa). The negative effect of stem length was compensated for by decreasing petiole hydraulic resistance and by increasing stem sapwood area and water storage, with capacitive discharge representing 8-12% of the water flux. In addition, large lateral (petiole, leaves) relative to axial hydraulic resistance helped improve water flow distribution to top leaves. These results indicate that gs of distal leaves can be similar to that of basal leaves, provided that resistance is highest in petioles, and sufficient amounts of water storage can be used to subsidize the transpiration stream.

Evapotranspiration and water yield of a pine-broadleaf forest are not altered by long-term atmospheric [CO2 ] enrichment under native or enhanced soil fertility.

Changes in evapotranspiration (ET) from terrestrial ecosystems affect their water yield (WY), with considerable ecological and economic consequences. Increases in surface runoff observed over the past century have been attributed to increasing atmospheric CO2 concentrations resulting in reduced ET by terrestrial ecosystems. Here, we evaluate the water balance of a Pinus taeda (L.) forest with a broadleaf component that was exposed to atmospheric [CO2 ] enrichment (ECO2 ; +200 ppm) for over 17 years and fertilization for 6 years, monitored with hundreds of environmental and sap flux sensors on a half-hourly basis. These measurements were synthesized using a one-dimensional Richard's equation model to evaluate treatment differences in transpiration (T), evaporation (E), ET, and WY. We found that ECO2 did not create significant differences in stand T, ET, or WY under either native or enhanced soil fertility, despite a 20% and 13% increase in leaf area index, respectively. While T, ET, and WY responded to fertilization, this response was weak (<3% of mean annual precipitation). Likewise, while E responded to ECO2 in the first 7 years of the study, this effect was of negligible magnitude (<1% mean annual precipitation). Given the global range of conifers similar to P. taeda, our results imply that recent observations of increased global streamflow cannot be attributed to decreases in ET across all ecosystems, demonstrating a great need for model-data synthesis activities to incorporate our current understanding of terrestrial vegetation in global water cycle models.

The carbon bonus of organic nitrogen enhances nitrogen use efficiency of plants.

The importance of organic nitrogen (N) for plant nutrition and productivity is increasingly being recognized. Here we show that it is not only the availability in the soil that matters, but also the effects on plant growth. The chemical form of N taken up, whether inorganic (such as nitrate) or organic (such as amino acids), may significantly influence plant shoot and root growth, and nitrogen use efficiency (NUE). We analysed these effects by synthesizing results from multiple laboratory experiments on small seedlings (Arabidopsis, poplar, pine and spruce) based on a tractable plant growth model. A key point is that the carbon cost of assimilating organic N into proteins is lower than that of inorganic N, mainly because of its carbon content. This carbon bonus makes it more beneficial for plants to take up organic than inorganic N, even when its availability to the roots is much lower - up to 70% lower for Arabidopsis seedlings. At equal growth rate, root:shoot ratio was up to three times higher and nitrogen productivity up to 20% higher for organic than inorganic N, which both are factors that may contribute to higher NUE in crop production.

Dynamics of soil CO2 efflux under varying atmospheric CO2 concentrations reveal dominance of slow processes.

We evaluated the effect on soil CO2 efflux (FCO2 ) of sudden changes in photosynthetic rates by altering CO2 concentration in plots subjected to +200 ppmv for 15 years. Five-day intervals of exposure to elevated CO2 (eCO2 ) ranging 1.0-1.8 times ambient did not affect FCO2 . FCO2 did not decrease until 4 months after termination of the long-term eCO2 treatment, longer than the 10 days observed for decrease of FCO2 after experimental blocking of C flow to belowground, but shorter than the ~13 months it took for increase of FCO2 following the initiation of eCO2 . The reduction of FCO2 upon termination of enrichment (~35%) cannot be explained by the reduction of leaf area (~15%) and associated carbohydrate production and allocation, suggesting a disproportionate contraction of the belowground ecosystem components; this was consistent with the reductions in base respiration and FCO2 -temperature sensitivity. These asymmetric responses pose a tractable challenge to process-based models attempting to isolate the effect of individual processes on FCO2 .

Informing climate models with rapid chamber measurements of forest carbon uptake.

Models predicting ecosystem carbon dioxide (CO2 ) exchange under future climate change rely on relatively few real-world tests of their assumptions and outputs. Here, we demonstrate a rapid and cost-effective method to estimate CO2 exchange from intact vegetation patches under varying atmospheric CO2 concentrations. We find that net ecosystem CO2 uptake (NEE) in a boreal forest rose linearly by 4.7 ± 0.2% of the current ambient rate for every 10 ppm CO2 increase, with no detectable influence of foliar biomass, season, or nitrogen (N) fertilization. The lack of any clear short-term NEE response to fertilization in such an N-limited system is inconsistent with the instantaneous downregulation of photosynthesis formalized in many global models. Incorporating an alternative mechanism with considerable empirical support - diversion of excess carbon to storage compounds - into an existing earth system model brings the model output into closer agreement with our field measurements. A global simulation incorporating this modified model reduces a long-standing mismatch between the modeled and observed seasonal amplitude of atmospheric CO2 . Wider application of this chamber approach would provide critical data needed to further improve modeled projections of biosphere-atmosphere CO2 exchange in a changing climate.

Annual climate variation modifies nitrogen induced carbon accumulation of Pinus sylvestris forests.

We report results from long-term simulated external nitrogen (N) input experiments in three northern Pinus sylvestris forests, two of moderately high and one of moderately low productivity, assessing effects on annual net primary production (NPP) of woody mass and its interannual variation in response to variability in weather conditions. A sigmoidal response of wood NPP to external N inputs was observed in the both higher and lower productivity stands, reaching a maximum of ~65% enhancement regardless of the native site productivity, saturating at an external N input of 4-5 g N·m-2 ·yr-1 . The rate of increase in wood NPP and the N response efficiency (REN , increase in wood NPP per external N input) were maximized at an external N input of ~3 g N·m-2 ·yr-1 , regardless of site productivity. The maximum REN was greater in the higher productivity than the lower productivity stand (~20 vs. ~14 g C/g N). The N-induced enhancement of wood NPP and its REN were, however, markedly contingent on climatic variables. In both of the higher and lower productivity stands, wood NPP increased with growing season precipitation (P), but only up to ~400 mm. The sensitivity of the response to P increased with increasing external N inputs. Increasing growing season temperature (T) somewhat increased the N-induced drought effect, whereas decreasing T reduced the drought effect. These responses of wood NPP infused a large temporal variation to REN , making the use of a fixed value unadvisable. Based on these results, we suggest that regional climate conditions and future climate scenarios should be considered when modeling carbon sequestration in response to N deposition in boreal P. sylvestris, and possibly other forests.

Ecophysiological variation of transpiration of pine forests: synthesis of new and published results.

Canopy transpiration (EC ) is a large fraction of evapotranspiration, integrating physical and biological processes within the energy, water, and carbon cycles of forests. Quantifying EC is of both scientific and practical importance, providing information relevant to questions ranging from energy partitioning to ecosystem services, such as primary productivity and water yield. We estimated EC of four pine stands differing in age and growing on sandy soils. The stands consisted of two wide-ranging conifer species: Pinus taeda and Pinus sylvestris, in temperate and boreal zones, respectively. Combining results from these and published studies on all soil types, we derived an approach to estimate daily EC of pine forests, representing a wide range of conditions from 35° S to 64° N latitude. During the growing season and under moist soils, maximum daily EC (ECm ) at day-length normalized vapor pressure deficit of 1 kPa (ECm-ref ) increased by 0.55 ± 0.02 (mean ± SE) mm/d for each unit increase of leaf area index (L) up to L = ~5, showing no sign of saturation within this range of quickly rising mutual shading. The initial rise of ECm with atmospheric demand was linearly related to ECm-ref . Both relations were unaffected by soil type. Consistent with theoretical prediction, daily EC was sensitive to decreasing soil moisture at an earlier point of relative extractable water in loamy than sandy soils. Our finding facilitates the estimation of daily EC of wide-ranging pine forests using remotely sensed L and meteorological data. We advocate an assembly of worldwide sap flux database for further evaluation of this approach.

Observed increase in local cooling effect of deforestation at higher latitudes.

Deforestation in mid- to high latitudes is hypothesized to have the potential to cool the Earth's surface by altering biophysical processes. In climate models of continental-scale land clearing, the cooling is triggered by increases in surface albedo and is reinforced by a land albedo-sea ice feedback. This feedback is crucial in the model predictions; without it other biophysical processes may overwhelm the albedo effect to generate warming instead. Ongoing land-use activities, such as land management for climate mitigation, are occurring at local scales (hectares) presumably too small to generate the feedback, and it is not known whether the intrinsic biophysical mechanism on its own can change the surface temperature in a consistent manner. Nor has the effect of deforestation on climate been demonstrated over large areas from direct observations. Here we show that surface air temperature is lower in open land than in nearby forested land. The effect is 0.85 ± 0.44 K (mean ± one standard deviation) northwards of 45° N and 0.21 ± 0.53 K southwards. Below 35° N there is weak evidence that deforestation leads to warming. Results are based on comparisons of temperature at forested eddy covariance towers in the USA and Canada and, as a proxy for small areas of cleared land, nearby surface weather stations. Night-time temperature changes unrelated to changes in surface albedo are an important contributor to the overall cooling effect. The observed latitudinal dependence is consistent with theoretical expectation of changes in energy loss from convection and radiation across latitudes in both the daytime and night-time phase of the diurnal cycle, the latter of which remains uncertain in climate models.

Greater carbon allocation to mycorrhizal fungi reduces tree nitrogen uptake in a boreal forest.

The central role that ectomycorrhizal (EM) symbioses play in the structure and function of boreal forests pivots around the common assumption that carbon (C) and nitrogen (N) are exchanged at rates favorable for plant growth. However, this may not always be the case. It has been hypothesized that the benefits mycorrhizal fungi convey to their host plants strongly depends upon the availability of C and N, both of which are rapidly changing as a result of intensified human land use and climate change. Using large-scale shading and N addition treatments, we assessed the independent and interactive effects of changes in C and N supply on the transfer of N in intact EM associations with -15 yr. old Scots pine trees. To assess the dynamics of N transfer in EM symbioses, we added trace amounts of highly enriched 5NO3(-) label to the EM-dominated mor-layer and followed the fate of the 15N label in tree foliage, fungal chitin on EM root tips, and EM sporocarps. Despite no change in leaf biomass, shading resulted in reduced tree C uptake, ca. 40% lower fungal biomass on EM root tips, and greater 15N label in tree foliage compared to unshaded control plots, where more 15N label was found in fungal biomass on EM colonized root tips. Short-term addition of N shifted the incorporation of 15N label from EM fungi to tree foliage, despite no significant changes in below-ground tree C allocation to EM fungi. Contrary to the common assumption that C and N are exchanged at rates favorable for plant growth, our results show for the first time that under N-limited conditions greater C allocation to EM fungi in the field results in reduced, not increased, N transfer to host trees. Moreover, given the ubiquitous nature of mycorrhizal symbioses, our results stress the need to incorporate mycorrhizal dynamics into process-based ecosystem models to better predict forest C and N cycles in light of global climate change.

Response to CO2 enrichment of understory vegetation in the shade of forests.

Responses of forest ecosystems to increased atmospheric CO2 concentration have been studied in few free-air CO2 enrichment (FACE) experiments during last two decades. Most studies focused principally on the overstory trees with little attention given to understory vegetation. Despite its small contribution to total productivity of an ecosystem, understory vegetation plays an important role in predicting successional dynamics and future plant community composition. Thus, the response of understory vegetation in Pinus taeda plantation at the Duke Forest FACE site after 15-17 years of exposure to elevated CO2 , 6-13 of which with nitrogen (N) amendment, was examined. Aboveground biomass and density of the understory decreased across all treatments with increasing overstory leaf area index (LAI). However, the CO2 and N treatments had no effect on aboveground biomass, tree density, community composition, and the fraction of shade-tolerant species. The increases of overstory LAI (~28%) under elevated CO2 resulted in a reduction of light available to the understory (~18%) sufficient to nullify the expected growth-enhancing effect of elevated CO2 on understory vegetation.

Increases in atmospheric CO2 have little influence on transpiration of a temperate forest canopy.

Models of forest energy, water and carbon cycles assume decreased stomatal conductance with elevated atmospheric CO2 concentration ([CO2]) based on leaf-scale measurements, a response not directly translatable to canopies. Where canopy-atmosphere are well-coupled, [CO2 ]-induced structural changes, such as increasing leaf-area index (LD), may cause, or compensate for, reduced mean canopy stomatal conductance (GS), keeping transpiration (EC) and, hence, runoff unaltered. We investigated GS responses to increasing [CO2] of conifer and broadleaved trees in a temperate forest subjected to 17-yr free-air CO2 enrichment (FACE; + 200 µmol mol(-1)). During the final phase of the experiment, we employed step changes of [CO2] in four elevated-[CO2 ] plots, separating direct response to changing [CO2] in the leaf-internal air-space from indirect effects of slow changes via leaf hydraulic adjustments and canopy development. Short-term manipulations caused no direct response up to 1.8 × ambient [CO2], suggesting that the observed long-term 21% reduction of GS was an indirect effect of decreased leaf hydraulic conductance and increased leaf shading. Thus, EC was unaffected by [CO2] because 19% higher canopy LD nullified the effect of leaf hydraulic acclimation on GS . We advocate long-term experiments of duration sufficient for slow responses to manifest, and modifying models predicting forest water, energy and carbon cycles accordingly.

The space-time continuum: the effects of elevated CO2 and temperature on trees and the importance of scaling.

To predict how forests will respond to rising temperatures and atmospheric CO2 concentrations, we need to understand how trees respond to both of these environmental factors. In this review, we discuss the importance of scaling, moving from leaf-level responses to those of the canopy, and from short-term to long-term responses of vegetation to climate change. While our knowledge of leaf-level, instantaneous responses of photosynthesis, respiration, stomatal conductance, transpiration and water-use efficiency to elevated CO2 and temperature is quite good, our ability to scale these responses up to larger spatial and temporal scales is less developed. We highlight which physiological processes are least understood at various levels of study, and discuss how ignoring differences in the spatial or temporal scale of a physiological process impedes our ability to predict how forest carbon and water fluxes forests will be altered in the future. We also synthesize data from the literature to show that light respiration follows a generalized temperature response across studies, and that the light compensation point of photosynthesis is reduced by elevated growth CO2. Lastly, we emphasize the need to move beyond single factorial experiments whenever possible, and to combine both CO2 and temperature treatments in studies of tree performance.

Role of aquaporins in determining transpiration and photosynthesis in water-stressed plants: crop water-use efficiency, growth and yield.

The global shortage of fresh water is one of our most severe agricultural problems, leading to dry and saline lands that reduce plant growth and crop yield. Here we review recent work highlighting the molecular mechanisms allowing some plant species and genotypes to maintain productivity under water stress conditions, and suggest molecular modifications to equip plants for greater production in water-limited environments. Aquaporins (AQPs) are thought to be the main transporters of water, small and uncharged solutes, and CO2 through plant cell membranes, thus linking leaf CO2 uptake from the intercellular airspaces to the chloroplast with water loss pathways. AQPs appear to play a role in regulating dynamic changes of root, stem and leaf hydraulic conductivity, especially in response to environmental changes, opening the door to using AQP expression to regulate plant water-use efficiency. We highlight the role of vascular AQPs in regulating leaf hydraulic conductivity and raise questions regarding their role (as well as tonoplast AQPs) in determining the plant isohydric threshold, growth rate, fruit yield production and harvest index. The tissue- or cell-specific expression of AQPs is discussed as a tool to increase yield relative to control plants under both normal and water-stressed conditions.