Effects of drought on mesophyll conductance and photosynthetic limitations at different tree canopy layers

In recent years, many studies have focused on the limiting role of mesophyll conductance (g_m) to photosynthesis (A_n) under water stress, but no studies have examined the effect of drought on g_m through the forest canopy. We investigated limitations to A_n on leaves at different heights in a mixed adult stand of sessile oak (Quercus petraea) and beech (Fagus sylvatica) trees during a moderately dry summer. Moderate drought decreased A_n of top and lowest beech canopy leaves much more than in leaves located in the mid canopy; whereas in oak, A_n of the lower canopy was decreased more than in sunlit leaves. The decrease of A_n was probably not due to leaf‐level biochemistry given that V_Cmax was generally unaffected by drought. The reduction in A_n was instead associated with reduction in stomatal and mesophyll conductances. Drought‐induced increases in stomatal limitations were largest in leaves from the top canopy, whereas drought‐induced increases in mesophyll limitations were largest in leaves from the lowest canopy. Sensitivity analysis highlighted the need to decompose the canopy into different leaf layers and to incorporate the limitation imposed by g_m when assessing the impact of drought on the gas exchange of tree canopies.     

Canopy structure and vertical patterns of photosynthesis and related leaf traits in a deciduous forest

Canopy structure and light interception were measured in an 18-m tall, closed canopy deciduous forest of sugar maple (Acer saccharum) in southwestern Wisconsin, USA, and related to leaf structural characteristics, N content, and leaf photosynthetic capacity. Light attenuation in the forest occurred primarily in the upper and middle portions of the canopy. Forest stand leaf area index (LAI) and its distribution with respect to canopy height were estimated from canopy transmittance values independently verified with a combined leaf litterfall and point-intersect method. Leaf mass, N and A _max per unit area (LMA, N/area and A _max/area, respectively) all decreased continuously by over two-fold from the upper to lower canopy, and these traits were strongly correlated with cumulative leaf area above the leaf position in the canopy. In contrast, neither N concentration nor A _max per unit mass varied significantly in relation to the vertical canopy gradient. Since leaf N concentration showed no consistent pattern with respect to canopy position, the observed vertical pattern in N/area is a direct consequence of vertical variation of LMA. N/area and LMA were strongly correlated with A _max/area among different canopy positions (r²=0.81 and r²=0.66, respectively), indicating that vertical variation in area-based photosynthetic capacity can also be attributed to variation in LMA. A model of whole-canopy photosynthesis was used to show that observed or hypothetical canopy mass distributions toward higher LMA (and hence higher N/area) in the upper portions of the canopy tended to increase integrated daily canopy photosynthesis over other LMA distribution patterns. Empirical relationships between leaf and canopy-level characteristics may help resolve problems associated with scaling gas exchange measurements made at the leaf level to the individual tree crown and forest canopy-level.     

Leaf gas exchange of trees in old-growth and young secondary forest stands in Sulawesi, Indonesia

In the tropics, old-growth forests are converted to other land cover types at a high rate and young secondary forest may gain in importance. Information on associated changes in leaf gas exchange and other leaf traits can be valuable for modelling biogeochemical fluxes under altered land-use patterns. We studied in situ photosynthetic parameters and stomatal conductance for water vapour in eight abundant tree species of young secondary forest and eight tree species of natural old-growth forest in Central Sulawesi, Indonesia. In sun leaves, the average maximal stomatal conductance (g _smax) in the secondary forest (SF) species was 2.1 times higher than in the old-growth forest (OGF) species. Species with a high g _smax reduced g _s sharply when vapour pressure deficit of the air increased, whereas species with a low g _smax were much less sensitive to air humidity. For area-based photosynthetic capacity (A _max-area), the SF species had a 2.3 times higher average than the OGF species. For both, g _smax and A _max-area the variation among species was higher in the OGF than in the SF. When all tree species (n=16) are considered, species means of specific leaf area (SLA), leaf N concentration and leaf P concentration were significantly correlated with g _smax and A _max-area. The strong correlation between A _max-area and foliar P (r ²=0.8) is remarkable as the alluvial soils in the study region are rich in nutrients. If the eight OGF species are analysed separately, the only significant correlation was observed between SLA and mass-based A _max; in the SF species strong correlations were found between leaf size and A _max-area and g _smax. These results show that the conversion of old-growth forest to young secondary forest in Sulawesi significantly alters tree leaf gas exchange characteristics and that chemical and structural leaf traits can be used for the prediction of these changes. The best correlations between leaf gas exchange parameters and leaf traits were obtained by different traits in the SF species, the OGF species and the entire pool of studied species.     

Stomatal closure induced by high vapor pressure deficit limited midday photosynthesis at the canopy top of <Emphasis Type="Italic">Fagus crenata</Emphasis> Blume on Naeba mountain in Japan

Diurnal changes in gas exchange, chlorophyll fluorescence and leaf water potential (_leaf) were measured to determine the environmental and physiological factors that limit carbon gain in the horizontal leaves of Fagus crenata Blume at the canopy top. Although midday depression of the net CO₂ assimilation rate (A_n) and stomatal conductance (gH₂O) were clearly evident on a fine day, the potential quantum yield of PS II (F_v/F_m) was fairly constant around 0.83 throughout the day. This result indicates that the leaves at the canopy top do not suffer from chronic photoinhibition, and the excess energy is dissipated safely. Large reversible increases in non-photochemical quenching (NPQ) were evident on fine days. Therefore, the non-radiative energy dissipation of excess light energy contributed to avoid chronic photoinhibition. The electron transfer rate (ETR) reached maximum during the midday depression, and thus there was no positive relation between ETR and A_n under high light conditions, indicating a high rate of photorespiration and the absence of non-stomatal effect during midday. The protective mechanisms such as non-radiative energy dissipation and photorespiration play an important role in preventing photoinhibitory damage, and stomatal limitation is the main factor of midday depression of A_n. To separate the effect of air to leaf vapor pressure deficit (ALVPD) and leaf temperature (T_leaf) on gas exchange, the dependencies of A_n and gH₂O on ALVPD and T_leaf were measured using detached branches under controlled conditions. A_n and gH₂O were insensitive to an increase in T_leaf. With the increase in ALVPD, A_n and gH₂O exhibited more than a 50% decrease even though water supply was optimum, suggesting the dominant role of high ALVPD in the midday depression of gH₂O. We conclude that midday depression of A_n results from the midday stomatal closure caused by high ALVPD.     

Seasonal and within‐canopy variation in shoot‐scale resource‐use efficiency trade‐offs in a Norway spruce stand

Previous leaf‐scale studies of carbon assimilation describe short‐term resource‐use efficiency (RUE) trade‐offs where high use efficiency of one resource requires low RUE of another. However, varying resource availabilities may cause long‐term RUE trade‐offs to differ from the short‐term patterns. This may have important implications for understanding canopy‐scale resource use and allocation. We used continuous gas exchange measurements collected at five levels within a Norway spruce, Picea abies (L.) karst., canopy over 3 years to assess seasonal differences in the interactions between shoot‐scale resource availability (light, water and nitrogen), net photosynthesis (A_n) and the use efficiencies of light (LUE), water (WUE) and nitrogen (NUE) for carbon assimilation. The continuous data set was used to develop and evaluate multiple regression models for predicting monthly shoot‐scale A_n. These models showed that shoot‐scale A_n was strongly dependent on light availability and was generally well described with simple one‐ or two‐parameter models. WUE peaked in spring, NUE in summer and LUE in autumn. However, the relative importance of LUE for carbon assimilation increased with canopy depth at all times. Our results suggest that accounting for seasonal and within‐canopy trade‐offs may be important for RUE‐based modelling of canopy carbon uptake.     

Coupled response of stomatal and mesophyll conductance to light enhances photosynthesis of shade leaves under sunflecks

Light gradients within tree canopies play a major role in the distribution of plant resources that define the photosynthetic capacity of sun and shade leaves. However, the biochemical and diffusional constraints on gas exchange in sun and shade leaves in response to light remain poorly quantified, but critical for predicting canopy carbon and water exchange. To investigate the CO₂ diffusion pathway of sun and shade leaves, leaf gas exchange was coupled with concurrent measurements of carbon isotope discrimination to measure net leaf photosynthesis (A_n), stomatal conductance (g_s) and mesophyll conductance (g_m) in Eucalyptus tereticornis trees grown in climate controlled whole‐tree chambers. Compared to sun leaves, shade leaves had lower A_n, g_m, leaf nitrogen and photosynthetic capacity (A_max) but g_s was similar. When light intensity was temporarily increased for shade leaves to match that of sun leaves, both g_s and g_m increased, and A_n increased to values greater than sun leaves. We show that dynamic physiological responses of shade leaves to altered light environments have implications for up‐scaling leaf level measurements and predicting whole canopy carbon gain. Despite exhibiting reduced photosynthetic capacity, the rapid up‐regulation of g_m with increased light enables shade leaves to respond quickly to sunflecks.     

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.     

Towards a coupled paradigm of NH3‐CO2 biosphere–atmosphere exchange modelling

Stomatal conductance, one of the major plant physiological controls within NH₃ biosphere–atmosphere exchange models, is commonly estimated from semi‐empirical multiplicative schemes or simple light‐ and temperature‐response functions. However, due to their inherent parameterization on meteorological proxy variables, instead of a direct measure of stomatal opening, they are unfit for the use in climate change scenarios and of limited value for interpreting field‐scale measurements. Alternatives based on H₂O flux measurements suffer from uncertainties in the partitioning of evapotranspiration at humid sites, as well as a potential decoupling of transpiration from stomatal opening in the presence of hygroscopic particles on leaf surfaces. We argue that these problems may be avoided by directly deriving stomatal conductance from CO₂ fluxes instead. We reanalysed a data set of NH₃ flux measurements based on CO₂‐derived stomatal conductance, confirming the hypothesis that the increasing relevance of stomatal exchange with the onset of vegetation activity caused a rapid decrease of observed NH₃ deposition velocities. Finally, we argue that developing more mechanistic representations of NH₃ biosphere–atmosphere exchange can be of great benefit in many applications. These range from model‐based flux partitioning, over deposition monitoring using low‐cost samplers and inferential modelling, to a direct response of NH₃ exchange to climate change.     

An In Situ Leaf and Branch Warming Experiment in the Amazon

Leaves and branches of mature trees, lianas, and gap species were warmed in an Amazonian forest for 4 mo to observe the effect of warming on photosynthesis, stomatal conductance, and transpiration. Electric resistance heaters increased air temperatures near the leaves by approximately 2°C. Sunlit leaf temperatures increased by 2–3°C on average, but during some periods leaf temperatures increased by >5°C. Maximum photosynthesis (A_max) decreased significantly in the warmed leaves vs. the control leaves over the 13‐wk study period with an average decrease in A_max of 1.4 μmol/m²s (19% decrease from a mean A_max of 7.2 μmol/m²s) when measured at 30°C and there were no signs of acclimation to higher temperatures within existing leaves. The decline in A_max was likely due to irreversible temperature damage caused by very high leaf temperatures and not due to C_i limitation of carboxylation. Warming had a larger negative impact on A_max in canopy level tree species than other tested functional groups such as lianas or gap species. Transpiration did not significantly increase in the warmed leaves compared with the control group. This study indicates that increased temperatures due to global warming could potentially decrease future tropical forest carbon uptake by a significant amount. Abstract in Portuguese is available at http://www.blackwell‐synergy.com/loi/btp .     

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.     

Microclimate and fluxes of water vapour, sensible heat and carbon dioxide in structurally differing subalpine plant communities in the Central Caucasus

Effects of canopy structure on microclimate, energy budget and CO₂ exchange were analysed in a pasture, two hay meadows, a tall herb community and a dwarf shrub community in the subalpine belt of the Central Caucasus. The results show that canopy structure exerts a marked influence on the distribution of photon flux density, temperature and canopy photosynthesis A_c. Three canopy types were distinguished. Type 1 (pasture) has a small LAI (leaf area index) and more than two-thirds of the phytomass is concentrated in the lowest few cm of the canopy, mainly as planophile leaves. This results in (1) a low degree of utilization of photosynthetic photon flux density (PPFD) by assimilatory plant components, (2) high leaf temperatures and a high soil heat flux during the phase of incoming radiation, and (3) a relatively low A_c/ LAI ratio. Type 2 (meadows), in spite of its erect leaves, which at high solar elevations permit light to penetrate to the lower canopy layers, is characterized by (1) marked effects of mutual shading in the lower canopy layers for most of the day, and thus (2) only slight variations in air and leaf temperatures and (3) a comparatively low A_c/LAI ratio. In canopies of type 3 (tall herb and dwarf shrub communities), there is a concentration of flat leaves in the upper layers. This results in (1) very good utilization of PPFD; (2) no strong fluctuations in canopy temperature as the flat leaves act as a buffer, reducing the amounts of incoming and outgoing radiation in lower canopy layers, and (3) high values of the A_c/LAI ratio. The energy budgets of the canopies investigated are governed not so much by their spatial structure, but rather indirectly by LAI and the degree of coupling of the canopy with the atmosphere.     

Influence of vegetation and soil CO2 exchange on the concentration and stable oxygen isotope ratio of atmospheric CO2 within a Pinus resinosa canopy

Measurements were made of the concentration and stable oxygen isotopic ratio of carbon dioxide in air samples collected on a diurnal basis at two heights within a Pinus resinosa canopy. Large changes in CO₂ concentration and isotopic composition were observed during diurnal time courses on all three symple dates. In addition, there was strong vertical stratification in the forest canopy, with higher CO₂ concentrations and more negative ¹⁸O values observed closer to the soil surface. The observed daily increases in ¹⁸O values of forest CO₂ were dependent on relative humidity consistent with the modelled predictions of isotopic fractionation during photosynthetic gas exchange. During photosynthetic gas exchange, a portion of the CO₂ that enters the leaf and equilibrates with leaf water is not fixed and diffuses back out of the leaf with an altered oxygen isotopic ratio. The oxygen isotope ratio of CO₂ diffusing out of a leaf depends primarily on the ¹⁸O content of leaf water which changes in response to relative humidity. In contrast, soil respiration caused a decline in the ¹⁸O values of forest CO₂ at night, because CO₂ released from the soil has equilibrated with soil water which has a lower ¹⁸O content than leaf water. The observed relationship between diurnal changes in CO₂ concentration and oxygen isotopic composition in the forest environment were consistent with a gas mixing model that considered the relative magnitudes of CO₂ fluxes associated with photosynthesis, respiration and turbulent exchange between the forest and the bulk atmosphere.     

Is distribution of hydraulic constraints within tree crowns reflected in photosynthetic water-use efficiency? An example of <Emphasis Type="Italic">Betula pendula</Emphasis>

Spatial and daily variation in photosynthetic water-use efficiency was examined in leaves of Betula pendula Roth with respect to distribution of hydraulic conductance within the crown, morphological properties of stomata, and water availability. Intrinsic water-use efficiency (A _n/g _s) was determined from gas-exchange measurements performed both in situ in a natural forest stand and on detached shoots under laboratory conditions. In intact foliage, sun leaves demonstrated significantly higher (P < 0.001) A _n/g _s than shade leaves, as photosynthesis in the lower canopy was chronically limited by low light availability. However, this difference reversed in the mid-day period under sufficient irradiance (I > 800 μmol m⁻² s⁻¹): A _n/g _s averaged 28.8 and 24.0 μmol mol⁻¹ (P < 0.01) for shade and sun leaves, respectively. This last finding coincided with the data obtained in laboratory conditions: under equivalent leaf water supply and light, A _n/g _s in shade foliage was greater (P < 0.001) than in sun foliage across a wide range of irradiance. Thus, shade foliage of B. pendula is characterized by inherently higher A _n/g _s than sun foliage, associated with more conservative stomatal behavior, and lower soil-to-leaf (K _T) and leaf hydraulic conductances. Under unlimited light conditions, a within-crown trade-off between A _n/g _s and K _T becomes apparent. Differences in stomatal conductance between the detached shoots from sunlit and shaded canopy layers were largely attributable to the variation in stomatal morphology; significant relationships were established with characteristics combining stomatal size and density (relative stomatal surface, stomatal pore area index). Stomatal morphology is very likely involved in long-term adjustment of photosynthetic WUE.     

Effects of seasonal and interannual variations in leaf photosynthesis and canopy leaf area index on gross primary production of a cool-temperate deciduous broadleaf forest in Takayama, Japan

Revealing the seasonal and interannual variations in forest canopy photosynthesis is a critical issue in understanding the ecological mechanisms underlying the dynamics of carbon dioxide exchange between the atmosphere and deciduous forests. This study examined the effects of temporal variations of canopy leaf area index (LAI) and leaf photosynthetic capacity [the maximum velocity of carboxylation (V _cmax)] on gross primary production (GPP) of a cool-temperate deciduous broadleaf forest for 5 years in Takayama AsiaFlux site, central Japan. We made two estimations to examine the effects of canopy properties on GPP; one is to incorporate the in situ observation of V _cmax and LAI throughout the growing season, and another considers seasonality of LAI but constantly high V _cmax. The simulations indicated that variation in V _cmax and LAI, especially in the leaf expansion period, had remarkable effects on GPP, and if V _cmax was assumed constant GPP will be overestimated by 15%. Monthly examination of air temperature, radiation, LAI and GPP suggested that spring temperature could affect canopy phenology, and also that GPP in summer was determined mainly by incoming radiation. However, the consequences among these factors responsible for interannual changes of GPP are not straightforward since leaf expansion and senescence patterns and summer meteorological conditions influence GPP independently. This simulation based on in situ ecophysiological research suggests the importance of intensive consideration and understanding of the phenology of leaf photosynthetic capacity and LAI to analyze and predict carbon fixation in forest ecosystems.     

Strong thermal acclimation of photosynthesis in tropical and temperate wet‐forest tree species: the importance of altered Rubisco content

Understanding of the extent of acclimation of light‐saturated net photosynthesis (A_n) to temperature (T), and associated underlying mechanisms, remains limited. This is a key knowledge gap given the importance of thermal acclimation for plant functioning, both under current and future higher temperatures, limiting the accuracy and realism of Earth system model (ESM) predictions. Given this, we analysed and modelled T‐dependent changes in photosynthetic capacity in 10 wet‐forest tree species: six from temperate forests and four from tropical forests. Temperate and tropical species were each acclimated to three daytime growth temperatures (T_growth): temperate – 15, 20 and 25 °C; tropical – 25, 30 and 35 °C. CO₂ response curves of A_n were used to model maximal rates of RuBP (ribulose‐1,5‐bisphosphate) carboxylation (V_cmax) and electron transport (J_max) at each treatment's respective T_growth and at a common measurement T (25 °C). SDS‐PAGE gels were used to determine abundance of the CO₂‐fixing enzyme, Rubisco. Leaf chlorophyll, nitrogen (N) and mass per unit leaf area (LMA) were also determined. For all species and T_growth, A_n at current atmospheric CO₂ partial pressure was Rubisco‐limited. Across all species, LMA decreased with increasing T_growth. Similarly, area‐based rates of V_cmax at a measurement T of 25 °C (V_cmax²⁵) linearly declined with increasing T_growth, linked to a concomitant decline in total leaf protein per unit leaf area and Rubisco as a percentage of leaf N. The decline in Rubisco constrained V_cmax and A_n for leaves developed at higher T_growth and resulted in poor predictions of photosynthesis by currently widely used models that do not account for T_growth‐mediated changes in Rubisco abundance that underpin the thermal acclimation response of photosynthesis in wet‐forest tree species. A new model is proposed that accounts for the effect of T_growth‐mediated declines in V_cmax²⁵ on A_n, complementing current photosynthetic thermal acclimation models that do not account for T sensitivity of V_cmax²⁵.     

Impact of insect defoliation on forest carbon balance as assessed with a canopy assimilation model

Disturbances such as fire, hurricanes, and herbivory often result in the net release of CO₂ from forests to the atmosphere, but the magnitude of carbon (C) loss is poorly quantified and difficult to predict. Here, we investigate the carbon balance of an oak/pine forest in the New Jersey Pine Barrens using the Canopy Conductance Constrained Carbon Assimilation (4C‐A) model. The 4C‐A model utilizes whole‐tree sap‐flux and leaf‐level photosynthetic gas exchange measurements at distinct canopy levels to estimate canopy assimilation. After model parameterization, sensitivity analyses, and evaluation against eddy flux measurements made in 2006, the model was used to predict C assimilation for an undisturbed year in 2005, and in 2007 when the stand was completely defoliated for 2–3 weeks during an infestation of gypsy moths (Lymantria dispar L.). Following defoliation, only 50% of the foliage reemerged in a second flush. In 2007, canopy net assimilation (A_nC), as modeled with the 4C‐A, was reduced to approximately 75% of A_nC in 2006 (940 vs. 1240 g C m^?2 a^?1). Overall, net primary production (NPP) in 2007 was approximately 240 g C m^?2 a^?1 (vs. 250 g C m^?2 a^?1 in 2006), with 60% of NPP allocated to foliage production, a short‐term carbon pool. Woody biomass accumulation, a long‐term carbon pool, was reduced by 20% compared with the previous year (72 vs. 57 g C m^?2 a^?1 in 2006 and 2007, respectively). The overall impact of the defoliation spanned 21% of upland forests (320 km²) in the New Jersey Pine Barrens, representing a significant amount of overall C not being taken up from the atmosphere by the forest, thus not accumulated in the biosphere.     

Exposure to an enriched CO2 atmosphere alters carbon assimilation and allocation in a pine forest ecosystem

We linked a leaf-level CO₂ assimilation model with a model that accounts for light attenuation in the canopy and measurements of sap-flux-based canopy conductance into a new canopy conductance-constrained carbon assimilation (4C-A) model. We estimated canopy CO₂ uptake (A_nC) at the Duke Forest free-air CO₂ enrichment (FACE) study. Rates of A_nC estimated from the 4C-A model agreed well with leaf gas exchange measurements (A_net) in both CO₂ treatments. Under ambient conditions, monthly sums of net CO₂ uptake by the canopy (A_nC) were 13% higher than estimates based on eddy-covariance and chamber measurements. Annual estimates of A_nC were only 3% higher than carbon (C) accumulations and losses estimated from ground-based measurements for the entire stand. The C budget for the Pinus taeda component was well constrained (within 1% of ground-based measurements). Although the closure of the C budget for the broadleaf species was poorer (within 20%), these species are a minor component of the forest. Under elevated CO₂, the C used annually for growth, turnover, and respiration balanced only 80% of the A_nC. Of the extra 700 g C m⁻² a⁻¹ (1999 and 2000 average), 86% is attributable to surface soil CO₂ efflux. This suggests that the production and turnover of fine roots was underestimated or that mycorrhizae and rhizodeposition became an increasingly important component of the C balance. Under elevated CO₂, net ecosystem production increased by 272 g C m⁻² a⁻¹: 44% greater than under ambient CO₂. The majority (87%) of this C was sequestered in a moderately long-term C pool in wood, with the remainder in the forest floor–soil subsystem.     

Interactive effect of elevated temperature and O<Subscript>3</Subscript> on antioxidant capacity and gas exchange in <Emphasis Type="Italic">Betula pendula</Emphasis> saplings

We studied the effects of slightly elevated temperature (T), O₃ concentration (O₃) and their combination (T + O₃) on the antioxidant defense, gas exchange and total leaf area of Betula pendula saplings in field conditions. During the second year of the experiment, T enhanced the total leaf area, net photosynthesis (P _n) and maximum capacity of carboxylation, redox state of ascorbate and total antioxidant capacity in the apoplast. O₃ did not affect the total leaf area, but P _n was slightly and g _s significantly reduced. The saplings responded to elevated O₃ level by closing the stomata and by developing leaves with a lower leaf area per mass, rather than by accumulating ascorbate in the apoplast. The effects of T and O₃ on total leaf area and P _n were counteractive. Elevated O₃ reduced the saplings’ ability to utilize the warmer growth environment by increasing the stomatal limitation for photosynthesis and by reducing the redox state of ascorbate in the apoplast in the combination treatment as compared to T alone.     

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.     

Leaf age does not justify its early removal in Carica papaya L

This study evaluated gas exchange and therefore sink/source condition of papaya leaves as they age in order to determine the right moment for leaf blade removal under two different strategies of climate control in greenhouses of Almería (SE Spain): Passive control (PCC; climate control of the greenhouse consisting of natural ventilation by only lateral and roof windows) versus active control (ACC; in addition to the natural ventilation, the greenhouse also incorporated a nebulization and a heating system). The effects of improved climate control were determined on papaya leaves distributed in five different positions in the canopy: S1, S2, S3, S4, S5, where S1 represented the youngest leaf, whereas S5 was the oldest (senescent) leaf present in the plant canopy. We observed that all age leaves presented positive values of net photosynthesis (A_net) despite reduced light availability under plastic greenhouses. In order to separate the effects of limited light availability from the leaf age (position in the canopy), we then repeated gas exchange measurements using artificial light (400 μmol m² s⁻¹) equal for all treatments and observed that the leaves of the ACC increased up to 96% their photosynthetic rates with respect to natural light. ACC plants presented also higher A_net/g_s, showing better intrinsic water use efficiency. Based on these results, we discourage blade removal of papaya leaves when light availability above 400 μmol m² s⁻¹ is assured by correct plant spacing and cultural management.