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.

Biomass increases attributed to both faster tree growth and altered allometric relationships under long-term carbon dioxide enrichment at a temperate forest.

Increases in atmospheric carbon dioxide (CO2 ) concentrations are expected to lead to increases in the rate of tree biomass accumulation, at least temporarily. On the one hand, trees may simply grow faster under higher CO2 concentrations, preserving the allometric relations that prevailed under lower CO2 concentrations. Alternatively, the allometric relations themselves may change. In this study, the effects of elevated CO2 (eCO2 ) on tree biomass and allometric relations were jointly assessed. Over 100 trees, grown at Duke Forest, NC, USA, were harvested from eight plots. Half of the plots had been subjected to CO2 enrichment from 1996 to 2010. Several subplots had also been subjected to nitrogen fertilization from 2005 to 2010. Allometric equations were developed to predict tree height, stem volume, and aboveground biomass components for loblolly pine (Pinus taeda L.), the dominant tree species, and broad-leaved species. Using the same diameter-based allometric equations for biomass, it was estimated that plots with eCO2 contained 21% more aboveground biomass, consistent with previous studies. However, eCO2 significantly affected allometry, and these changes had an additional effect on biomass. In particular, P. taeda trees at a given diameter were observed to be taller under eCO2 than under ambient CO2 due to changes in both the allometric scaling exponent and intercept. Accounting for allometric change increased the treatment effect of eCO2 on aboveground biomass from a 21% to a 27% increase. No allometric changes for the nondominant broad-leaved species were identified, nor were allometric changes associated with nitrogen fertilization. For P. taeda, it is concluded that eCO2 affects allometries, and that knowledge of allometry changes is necessary to accurately compute biomass under eCO2 . Further observations are needed to determine whether this assessment holds for other taxa.

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.

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.

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.

Sustained effects of atmospheric [CO2] and nitrogen availability on forest soil CO2 efflux.

Soil CO2 efflux (Fsoil ) is the largest source of carbon from forests and reflects primary productivity as well as how carbon is allocated within forest ecosystems. Through early stages of stand development, both elevated [CO2] and availability of soil nitrogen (N; sum of mineralization, deposition, and fixation) have been shown to increase gross primary productivity, but the long-term effects of these factors on Fsoil are less clear. Expanding on previous studies at the Duke Free-Air CO2 Enrichment (FACE) site, we quantified the effects of elevated [CO2] and N fertilization on Fsoil using daily measurements from automated chambers over 10 years. Consistent with previous results, compared to ambient unfertilized plots, annual Fsoil increased under elevated [CO2] (ca. 17%) and decreased with N (ca. 21%). N fertilization under elevated [CO2] reduced Fsoil to values similar to untreated plots. Over the study period, base respiration rates increased with leaf productivity, but declined after productivity saturated. Despite treatment-induced differences in aboveground biomass, soil temperature and water content were similar among treatments. Interannually, low soil water content decreased annual Fsoil from potential values - estimated based on temperature alone assuming nonlimiting soil water content - by ca. 0.7% per 1.0% reduction in relative extractable water. This effect was only slightly ameliorated by elevated [CO2]. Variability in soil N availability among plots accounted for the spatial variability in Fsoil , showing a decrease of ca. 114 g C m(-2) yr(-1) per 1 g m(-2) increase in soil N availability, with consistently higher Fsoil in elevated [CO2] plots ca. 127 g C per 100 ppm [CO2] over the +200 ppm enrichment. Altogether, reflecting increased belowground carbon partitioning in response to greater plant nutritional needs, the effects of elevated [CO2] and N fertilization on Fsoil in this stand are sustained beyond the early stages of stand development and through stabilization of annual foliage production.

Nitrogen-addition effects on leaf traits and photosynthetic carbon gain of boreal forest understory shrubs.

Boreal coniferous forests are characterized by fairly open canopies where understory vegetation is an important component of ecosystem C and N cycling. We used an ecophysiological approach to study the effects of N additions on uptake and partitioning of C and N in two dominant understory shrubs: deciduous Vaccinium myrtillus in a Picea abies stand and evergreen Vaccinium vitis-idaea in a Pinus sylvestris stand in northern Sweden. N was added to these stands for 16 and 8 years, respectively, at rates of 0, 12.5, and 50 kg N ha(-1) year(-1). N addition at the highest rate increased foliar N and chlorophyll concentrations in both understory species. Canopy cover of P. abies also increased, decreasing light availability and leaf mass per area of V. myrtillus. Among leaves of either shrub, foliar N content did not explain variation in light-saturated CO2 exchange rates. Instead photosynthetic capacity varied with stomatal conductance possibly reflecting plant hydraulic properties and within-site variation in water availability. Moreover, likely due to increased shading under P. abies and due to water limitations in the sandy soil under P. sylvestris, individuals of the two shrubs did not increase their biomass or shift their allocation between above- and belowground parts in response to N additions. Altogether, our results indicate that the understory shrubs in these systems show little response to N additions in terms of photosynthetic physiology or growth and that changes in their performance are mostly associated with responses of the tree canopy.

Hydraulic limits on maximum plant transpiration and the emergence of the safety-efficiency trade-off.

Soil and plant hydraulics constrain ecosystem productivity by setting physical limits to water transport and hence carbon uptake by leaves. While more negative xylem water potentials provide a larger driving force for water transport, they also cause cavitation that limits hydraulic conductivity. An optimum balance between driving force and cavitation occurs at intermediate water potentials, thus defining the maximum transpiration rate the xylem can sustain (denoted as E(max)). The presence of this maximum raises the question as to whether plants regulate transpiration through stomata to function near E(max). To address this question, we calculated E(max) across plant functional types and climates using a hydraulic model and a global database of plant hydraulic traits. The predicted E(max) compared well with measured peak transpiration across plant sizes and growth conditions (R = 0.86, P < 0.001) and was relatively conserved among plant types (for a given plant size), while increasing across climates following the atmospheric evaporative demand. The fact that E(max) was roughly conserved across plant types and scales with the product of xylem saturated conductivity and water potential at 50% cavitation was used here to explain the safety-efficiency trade-off in plant xylem. Stomatal conductance allows maximum transpiration rates despite partial cavitation in the xylem thereby suggesting coordination between stomatal regulation and xylem hydraulic characteristics.

On the complementary relationship between marginal nitrogen and water-use efficiencies among Pinus taeda leaves grown under ambient and CO2-enriched environments.

BACKGROUND AND AIMS: Water and nitrogen (N) are two limiting resources for biomass production of terrestrial vegetation. Water losses in transpiration (E) can be decreased by reducing leaf stomatal conductance (g(s)) at the expense of lowering CO(2) uptake (A), resulting in increased water-use efficiency. However, with more N available, higher allocation of N to photosynthetic proteins improves A so that N-use efficiency is reduced when g(s) declines. Hence, a trade-off is expected between these two resource-use efficiencies. In this study it is hypothesized that when foliar concentration (N) varies on time scales much longer than g(s), an explicit complementary relationship between the marginal water- and N-use efficiency emerges. Furthermore, a shift in this relationship is anticipated with increasing atmospheric CO(2) concentration (c(a)). METHODS: Optimization theory is employed to quantify interactions between resource-use efficiencies under elevated c(a) and soil N amendments. The analyses are based on marginal water- and N-use efficiencies, λ = (∂A/∂g(s))/(∂E/∂g(s)) and η = ∂A/∂N, respectively. The relationship between the two efficiencies and related variation in intercellular CO(2) concentration (c(i)) were examined using A/c(i) curves and foliar N measured on Pinus taeda needles collected at various canopy locations at the Duke Forest Free Air CO(2) Enrichment experiment (North Carolina, USA). KEY RESULTS: Optimality theory allowed the definition of a novel, explicit relationship between two intrinsic leaf-scale properties where η is complementary to the square-root of λ. The data support the model predictions that elevated c(a) increased η and λ, and at given c(a) and needle age-class, the two quantities varied among needles in an approximately complementary manner. CONCLUSIONS: The derived analytical expressions can be employed in scaling-up carbon, water and N fluxes from leaf to ecosystem, but also to derive transpiration estimates from those of η, and assist in predicting how increasing c(a) influences ecosystem water use.

Causality and persistence in ecological systems: a nonparametric spectral granger causality approach.

Abstract Directionality in coupling, defined as the linkage relating causes to their effects at a later time, can be used to explain the core dynamics of ecological systems by untangling direct and feedback relationships between the different components of the systems. Inferring causality from measured ecological variables sampled through time remains a formidable challenge further made difficult by the action of periodic drivers overlapping the natural dynamics of the system. Periodicity in the drivers can often mask the self-sustained oscillations originating from the autonomous dynamics. While linear and direct causal relationships are commonly addressed in the time domain, using the well-established machinery of Granger causality (G-causality), the presence of periodic forcing requires frequency-based statistics (e.g., the Fourier transform), able to distinguish coupling induced by oscillations in external drivers from genuine endogenous interactions. Recent nonparametric spectral extensions of G-causality to the frequency domain pave the way for the scale-by-scale decomposition of causality, which can improve our ability to link oscillatory behaviors of ecological networks to causal mechanisms. The performance of both spectral G-causality and its conditional extension for multivariate systems is explored in quantifying causal interactions within ecological networks. Through two case studies involving synthetic and actual time series, it is demonstrated that conditional G-causality outperforms standard G-causality in identifying causal links and their concomitant timescales.

Increases in the flux of carbon belowground stimulate nitrogen uptake and sustain the long-term enhancement of forest productivity under elevated CO2.

The earth's future climate state is highly dependent upon changes in terrestrial C storage in response to rising concentrations of atmospheric CO2. Here we show that consistently enhanced rates of net primary production (NPP) are sustained by a C-cascade through the root-microbe-soil system; increases in the flux of C belowground under elevated CO2 stimulated microbial activity, accelerated the rate of soil organic matter decomposition and stimulated tree uptake of N bound to this SOM. This process set into motion a positive feedback maintaining greater C gain under elevated CO2 as a result of increases in canopy N content and higher photosynthetic N-use efficiency. The ecosystem-level consequence of the enhanced requirement for N and the exchange of plant C for N belowground is the dominance of C storage in tree biomass but the preclusion of a large C sink in the soil.

Effects of stomatal delays on the economics of leaf gas exchange under intermittent light regimes.

• Understory plants are subjected to highly intermittent light availability and their leaf gas exchanges are mediated by delayed responses of stomata and leaf biochemistry to light fluctuations. In this article, the patterns in stomatal delays across biomes and plant functional types were studied and their effects on leaf carbon gains and water losses were quantified. • A database of more than 60 published datasets on stomatal responses to light fluctuations was assembled. To interpret these experimental observations, a leaf gas exchange model was developed and coupled to a novel formulation of stomatal movement energetics. The model was used to test whether stomatal delays optimize light capture for photosynthesis, whilst limiting transpiration and carbon costs for stomatal movement. • The data analysis showed that stomatal opening and closing delays occurred over a limited range of values and were strongly correlated. Plant functional type and climate were the most important drivers of stomatal delays, with faster responses in graminoids and species from dry climates. • Although perfectly tracking stomata would maximize photosynthesis and minimize transpiration at the expense of large opening costs, the observed combinations of opening and closure times appeared to be consistent with a near-optimal balance of carbon gain, water loss and movement costs.

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 lambda (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.

Physiological responses of Norway spruce (Picea abies) seedlings to drought stress.

Four-year-old seedlings of Picea abies [L.] Karst (Norway spruce) were grown in semi-controlled conditions with three watering regimes. The seedlings in the control group (c) were watered to prevent any dehydration effect. The two remaining groups were subjected to mild (ms) and severe water stress (ss), respectively. The following physiological variables were monitored until ss seedlings began to die: leaf water potential (psi(L)), stomatal conductance (g(s)), CO(2) exchange (P(N)), free proline content (Pro), total chlorophyll (a + b) concentration (Chl(t)) and the maximal photochemical efficiency of photosystem II (F(v)/F(m)). The results indicate that not all observed physiological parameters display the same degree of sensitivity to dehydration. After Day 12 of dehydration, psi(L) of ss seedlings was already significantly lower than that of the two other groups. On Day 26, significant differences in psi(L) were recorded among all treatments. Decreasing values of water potential were accompanied by early changes in P(N), g(s) and Pro. A significant decrease in Chl(t) and F(v)/F(m) were only observed during the more advanced stages of dehydration. These results demonstrate that the drought response of P. abies seedlings include a number of parallel physiological and biochemical changes in concert, enhancing the capability of plants to survive and grow during drought periods, but only to a point.

Leaf stomatal responses to vapour pressure deficit under current and CO(2)-enriched atmosphere explained by the economics of gas exchange.

Using the economics of gas exchange, early studies derived an expression of stomatal conductance (g) assuming that water cost per unit carbon is constant as the daily loss of water in transpiration (f(e)) is minimized for a given gain in photosynthesis (f(c)). Other studies reached identical results, yet assumed different forms for the underlying functions and defined the daily cost parameter as carbon cost per unit water. We demonstrated that the solution can be recovered when optimization is formulated at time scales commensurate with the response time of g to environmental stimuli. The optimization theory produced three emergent gas exchange responses that are consistent with observed behaviour: (1) the sensitivity of g to vapour pressure deficit (D) is similar to that obtained from a previous synthesis of more than 40 species showing g to scale as 1 - m log(D), where m is in [0.5,0.6], (2) the theory is consistent with the onset of an apparent 'feed-forward' mechanism in g, and (3) the emergent non-linear relationship between the ratio of intercellular to atmospheric [CO(2)] (c(i)/c(a)) and D agrees with the results available on this response. We extended the theory to diagnosing experimental results on the sensitivity of g to D under varying c(a).