CO2 enrichment in a maturing pine forest: are CO2 exchange and water status in the canopy affected?

A _net, leaf net CO₂ assimilation
c_a, CO₂ concentration of air surrounding a leaf
c_i, leaf intercellular CO₂ concentration
Δ, ¹³C isotope discrimination
δ¹³C, relative stable carbon isotope content
?, ratio of A_net at c_a = 560μmol mol^–1 to A_net at c_a = 360 μmol mol^–1
FACE, free-air CO₂ enrichment
g_w, stomatal conductance to water vapour
Π_i, initial leaf osmotic potential
R_t, relative water content at incipient turgor loss
Ψ_l, xylem water potential of leaves
Ψ_m, soil matric potential

Elevated CO₂ is expected to reduce forest water use as a result of CO₂-induced stomatal closure, which has implications for ecosystem-scale phenomena controlled by water availability. Leaf-level CO₂ and H₂O exchange responses and plant and soil water relations were examined in a maturing loblolly pine (Pinus taeda L.) stand in a free-air CO₂ enrichment (FACE) experiment in North Carolina, USA to test if these parameters were affected by elevated CO₂. Current-year foliage in the canopy was continuously exposed to elevated CO₂ (ambient CO₂+200μmol mol^–1) in free-air during needle growth and development for up to 400 d. Photosynthesis in upper canopy foliage was stimulated by 50–60% by elevated CO₂ compared with ambient controls. This enhancement was similar in current-year, ambient-grown foliage temporarily measured at elevated CO₂ compared with long-term elevated CO₂ grown foliage. Significant photosynthetic enhancement by CO₂ was maintained over a range of conditions except during peak drought. There was no evidence of water savings in elevated CO₂ plots in FACE compared to ambient plots under drought and non-drought conditions. This was supported by evidence from three independent measures. First, stomatal conductance was not significantly different in elevated CO₂ versus ambient trees of P. taeda. Calculations of time-integrated c_i/c_a ratios from analysis of foliar δ¹³C showed that these ratios were maintained in foliage under elevated CO₂. Second, soil moisture was not significantly different between ambient and elevated CO₂ plots during drought. Third, pre-dawn and mid-day leaf water potentials were also unaffected by the seasonal CO₂ exposure, as were tissue osmotic potentials and turgor loss points. Together the results strongly support the hypothesis that maturing P. taeda trees have low stomatal responsiveness to elevated CO₂. Elevated CO₂ effects on water relations in loblolly pine-dominated forest ecosystems may be absent or small apart from those mediated by leaf area. Large photosynthetic enhancements in the upper canopy of P. taeda by elevated CO₂ indicate that this maturing forest may have a large carbon sequestration capacity with limiting water supply.     

Seasonal trends of light-saturated net photosynthesis and stomatal conductance of loblolly pine trees grown in contrasting environments of nutrition, water and carbon dioxide

Repeated measures analysis was used to evaluate the effect of long-term CO₂ enhancement on seasonal trends of light-saturated rates of net photosynthesis (A_sat) and stomatal conductance to water vapour (g_sat) of 9-year-old loblolly pine (Pinus taeda L.) trees grown in a 2 × 2 factorial experimental design of nutrition and water. A significant interaction effect of CO₂ and nutrition on mean A_sat was observed for juvenile foliage. Also, juvenile foliage exposed to +350 μmol mol^?1 CO₂ had a higher rate of increase of A_sat between late summer and early autumn. This would lead to a greater potential for recharging carbohydrate reserves for winter. Mature foliage was affected by CO_sat, water and nutrient treatments in two ways. First, A_sat was significantly increased as a result of elevated CO₂ in January, a period when stomatal conductance was only 47% of the maximum observed rate. Secondly, the rate of increase of A_sat from winter to early spring was accelerated as a result of both nutrient + water and + 350 μmol mol^?1 CO₂ treatments. This accelerated response resulted in a greater potential for photosynthate production during the period when growth initiation occurred. Nutrient, water or carbon dioxide treatments did not significantly alter trends in g_sat for mature or juvenile foliage. A significant nutrition × CO₂ interaction was observed for the mature foliage, suggesting that g_sat increased with increasing CO₂ and nutrition. These results may have important consequences for the determination of the water use efficiency of loblolly pine. In spite of low g_sat in the winter to early spring period, there was a substantial gain in A_sat attributable to elevated CO₂ concentrations.     

Stomatal acclimation to increased CO2 concentration in a Florida scrub oak species Quercus myrtifolia Willd

Native scrub‐oak communities in Florida were exposed for three seasons in open top chambers to present atmospheric [CO₂] (approx. 350 μmol mol^?1) and to high [CO₂] (increased by 350 μmol mol^?1). Stomatal and photosynthetic acclimation to high [CO₂] of the dominant species Quercus myrtifolia was examined by leaf gas exchange of excised shoots. Stomatal conductance (g_s) was approximately 40% lower in the high‐ compared to low‐[CO₂]‐grown plants when measured at their respective growth concentrations. Reciprocal measurements of g_s in both high‐ and low‐[CO₂]‐grown plants showed that there was negative acclimation in the high‐[CO₂]‐grown plants (9–16% reduction in g_s when measured at 700 μmol mol^?1), but these were small compared to those for net CO₂ assimilation rate (A, 21–36%). Stomatal acclimation was more clearly evident in the curve of stomatal response to intercellular [CO₂] (c_i) which showed a reduction in stomatal sensitivity at low c_i in the high‐[CO₂]‐grown plants. Stomatal density showed no change in response to growth in high growth [CO₂]. Long‐term stomatal and photosynthetic acclimation to growth in high [CO₂] did not markedly change the 2·5‐ to 3‐fold increase in gas‐exchange‐derived water use efficiency caused by high [CO₂].     

Direct and acclimatory responses of stomatal conductance to elevated carbon dioxide in four herbaceous crop species in the field

In order to separate the net effect of growth at elevated [CO₂] on stomatal conductance (g_s) into direct and acclimatory responses, mid‐day values of g_s were measured for plants grown in field plots in open‐topped chambers at the current ambient [CO₂], which averaged 350 μmol mol^?1 in the daytime, and at ambient + 350 μmol mol^?1[CO₂] for winter wheat, winter barley, potato and sorghum. The acclimatory response was determined by comparing g_s measured at 700 μmol mol^?1[CO₂] for plants grown at the two [CO₂]. The direct effect of increasing [CO₂] from 350 to 700 μmol mol^?1 was determined for plants grown at the lower concentration. Photosynthetic rates were measured concurrently with g_s. For all species, growth at the higher [CO₂] significantly reduced g_s measured at 700 μmol mol^?1[CO₂]. The reduction in g_s caused by growth at the higher [CO₂] was larger for all species on days with low leaf to air water vapour pressure difference for a given temperature, which coincided with highest conductances and also the smallest direct effects of increased [CO₂] on conductance. For barley, there was no other evidence for stomatal acclimation, despite consistent down‐regulation of photosynthetic rate in plants grown at the higher [CO₂]. In wheat and potato, in addition to the vapour pressure difference interaction, the magnitude of stomatal acclimation varied directly in proportion to the magnitude of down‐regulation of photosynthetic rate through the season. In sorghum, g_s consistently exhibited acclimation, but there was no down‐regulation of photosynthetic rate. In none of the species except barley was the direct effect the larger component of the net reduction in g_s when averaged over measurement dates. The net effect of growth at elevated [CO₂] on mid‐day g_s resulted from unique combinations of direct and acclimatory responses in the various species.     

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

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

Stomatal sensitivity to vapour pressure difference over a subambient to elevated CO2 gradient in a C3/C4 grassland

In the present study the response of stomatal conductance (g_s) to increasing leaf‐to‐air vapour pressure difference (D) in early season C₃ (Bromus japonicus) and late season C₄ (Bothriochloa ischaemum) grasses grown in the field across a range of CO₂ (200–550 µmol mol^?1) was examined. Stomatal sensitivity to D was calculated as the slope of the response of g_s to the natural log of externally manipulated D (dg_s/dlnD). Increasing D and CO₂ significantly reduced g_s in both species. Increasing CO₂ caused a significant decrease in stomatal sensitivity to D in Br. japonicus, but not in Bo. ischaemum. The decrease in stomatal sensitivity to D at high CO₂ for Br. japonicus fit theoretical expectations of a hydraulic model of stomatal regulation, in which g_s varies to maintain constant transpiration and leaf water potential. The weaker stomatal sensitivity to D in Bo. ischaemum suggested that stomatal regulation of leaf water potential was poor in this species, or that non‐hydraulic signals influenced guard cell behaviour. Photosynthesis (A) declined with increasing D in both species, but analyses of the ratio of intercellular to atmospheric CO₂ (C_i/C_a) suggested that stomatal limitation of A occurred only in Br. japonicus. Rising CO₂ had the greatest effect on g_s and A in Br. japonicus at low D. In contrast, the strength of stomatal and photosynthetic responses to CO₂ were not affected by D in Bo. ischaemum. Carbon and water dynamics in this grassland are dominated by a seasonal transition from C₃ to C₄ photosynthesis. Interspecific variation in the response of g_s to D therefore has implications for predicting seasonal ecosystem responses to CO₂.     

Leaf photosynthesis and carbohydrate dynamics of soybeans grown throughout their life-cycle under Free-Air Carbon dioxide Enrichment

A lower than theoretically expected increase in leaf photosynthesis with long‐term elevation of carbon dioxide concentration ([CO₂]) is often attributed to limitations in the capacity of the plant to utilize the additional photosynthate, possibly resulting from restrictions in rooting volume, nitrogen supply or genetic constraints. Field‐grown, nitrogen‐fixing soybean with indeterminate flowering might therefore be expected to escape these limitations. Soybean was grown from emergence to grain maturity in ambient air (372 µmol mol^?1[CO₂]) and in air enriched with CO₂ (552 µmol mol^?1[CO₂]) using Free‐Air CO₂ Enrichment (FACE) technology. The diurnal courses of leaf CO₂ uptake (A) and stomatal conductance (g_s) for upper canopy leaves were followed throughout development from the appearance of the first true leaf to the completion of seed filling. Across the growing season the daily integrals of leaf photosynthetic CO₂ uptake (A′) increased by 24.6% in elevated [CO₂] and the average mid‐day g_s decreased by 21.9%. The increase in A′ was about half the 44.5% theoretical maximum increase calculated from Rubisco kinetics. There was no evidence that the stimulation of A was affected by time of day, as expected if elevated [CO₂] led to a large accumulation of leaf carbohydrates towards the end of the photoperiod. In general, the proportion of assimilated carbon that accumulated in the leaf as non‐structural carbohydrate over the photoperiod was small (< 10%) and independent of [CO₂] treatment. By contrast to A′, daily integrals of PSII electron transport measured by modulated chlorophyll fluorescence were not significantly increased by elevated [CO₂]. This indicates that A at elevated [CO₂] in these field conditions was predominantly ribulose‐1,5‐bisphosphate (RubP) limited rather than Rubisco limited. There was no evidence of any loss of stimulation toward the end of the growing season; the largest stimulation of A′ occurred during late seed filling. The stimulation of photosynthesis was, however, transiently lost for a brief period just before seed fill. At this point, daytime accumulation of foliar carbohydrates was maximal, and the hexose:sucrose ratio in plants grown at elevated [CO₂] was significantly larger than that in plants grown at current [CO₂]. The results show that even for a crop lacking the constraints that have been considered to limit the responses of C₃ plants to rising [CO₂] in the long term, the actual increase in A over the growing season is considerably less than the increase predicted from theory.     

Effects of instantaneous and growth CO2 levels and abscisic acid on stomatal and mesophyll conductances

C₃ photosynthesis is often limited by CO₂ diffusivity or stomatal (g_s) and mesophyll (g_m) conductances. To characterize effects of stomatal closure induced by either high CO₂ or abscisic acid (ABA) application on g_m, we examined g_s and g_m in the wild type (Col‐0) and ost1 and slac1‐2 mutants of Arabidopsis thaliana grown at 390 or 780 μmol mol^?1 CO₂. Stomata of these mutants were reported to be insensitive to both high CO₂ and ABA. When the ambient CO₂ increased instantaneously, g_m decreased in all these plants, whereas g_s in ost1 and slac1‐2 was unchanged. Therefore, the decrease in g_m in response to high CO₂ occurred irrespective of the responses of g_s. g_m was mainly determined by the instantaneous CO₂ concentration during the measurement and not markedly by the CO₂ concentration during the growth. Exogenous application of ABA to Col‐0 caused the decrease in the intercellular CO₂ concentration (C_i). With the decrease in C_i, g_m did not increase but decreased, indicating that the response of g_m to CO₂ and that to ABA are differently regulated and that ABA content in the leaves plays an important role in the regulation of g_m.     

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.     

Diurnal and seasonal variations in stomatal conductance of rice at elevated atmospheric CO2 under fully open‐air conditions

Understanding of leaf stomatal responses to the atmospheric CO₂ concentration, [CO₂], is essential for accurate prediction of plant water use under future climates. However, limited information is available for the diurnal and seasonal changes in stomatal conductance (g_s) under elevated [CO₂]. We examined the factors responsible for variations in g_s under elevated [CO₂] with three rice cultivars grown in an open‐field environment under flooded conditions during two growing seasons (a total of 2140 individual measurements). Conductance of all cultivars was generally higher in the morning and around noon than in the afternoon, and elevated [CO₂] decreased g_s by up to 64% over the 2 years (significantly on 26 out of 38 measurement days), with a mean g_s decrease of 23%. We plotted the g_s variations against three parameters from the Ball‐Berry model and two revised versions of the model, and all parameters explained the g_s variations well at each [CO₂] in the morning and around noon (R² > 0.68), but could not explain these variations in the afternoon (R² < 0.33). The present results provide an important basis for modelling future water use in rice production.     

Stomatal conductance and not stomatal density determines the long-term reduction in leaf transpiration of poplar in elevated CO<Subscript>2</Subscript>

Using a free-air CO₂ enrichment (FACE) experiment, poplar trees (Populus × euramericana clone I214) were exposed to either ambient or elevated [CO₂] from planting, for a 5-year period during canopy development, closure, coppice and re-growth. In each year, measurements were taken of stomatal density (SD, number mm⁻²) and stomatal index (SI, the proportion of epidermal cells forming stomata). In year 5, measurements were also taken of leaf stomatal conductance (g _s, μmol m⁻² s⁻¹), photosynthetic CO₂ fixation (A, mmol m⁻² s⁻¹), instantaneous water-use efficiency (A/E) and the ratio of intercellular to atmospheric CO₂ (C_i:C_a). Elevated [CO₂] caused reductions in SI in the first year, and in SD in the first 2 years, when the canopy was largely open. In following years, when the canopy had closed, elevated [CO₂] had no detectable effects on stomatal numbers or index. In contrast, even after 5 years of exposure to elevated [CO₂], g _s was reduced, A/E was stimulated, and C_i:C_a was reduced relative to ambient [CO₂]. These outcomes from the long-term realistic field conditions of this forest FACE experiment suggest that stomatal numbers (SD and SI) had no role in determining the improved instantaneous leaf-level efficiency of water use under elevated [CO₂]. We propose that altered cuticular development during canopy closure may partially explain the changing response of stomata to elevated [CO₂], although the mechanism for this remains obscure.     

Dynamic changes of canopy-scale mesophyll conductance to CO₂ diffusion of sunflower as affected by CO₂ concentration and abscisic acid

Leaf‐level measurements have shown that mesophyll conductance (g_m) can vary rapidly in response to CO₂ and other environmental factors, but similar studies at the canopy‐scale are missing. Here, we report the effect of short‐term variation of CO₂ concentration on canopy‐scale g_m and other CO₂ exchange parameters of sunflower (Helianthus annuus L.) stands in the presence and absence of abscisic acid (ABA) in their nutrient solution. g_m was estimated from gas exchange and on‐line carbon isotope discrimination (Δ_obs) in a ¹³CO₂/¹²CO₂ gas exchange mesocosm. The isotopic contribution of (photo)respiration to stand‐scale Δ_obs was determined with the experimental approach of Tcherkez et al. Without ABA, short‐term exposures to different CO₂ concentrations (C_a 100 to 900 µmol mol^?1) had little effect on canopy‐scale g_m. But, addition of ABA strongly altered the CO₂‐response: g_m was high (approx. 0.5 mol CO₂ m^?2 s^?1) at C_a < 200 µmol mol^?1 and decreased to <0.1 mol CO₂ m^?2 s^?1 at C_a >400 µmol mol^?1. In the absence of ABA, the contribution of (photo)respiration to stand‐scale Δ_obs was high at low C_a (7.2‰) and decreased to <2‰ at C_a > 400 µmol mol^?1. Treatment with ABA halved this effect at all C_a.     

The influence of elevated temperature,elevated atmospheric CO2 concentration and water stress on net photosynthesis of loblolly pine (Pinus taeda L.) at northern,central and southern sites in its native range

We investigated the effect of elevated [CO₂] (700 μmol mol^?1), elevated temperature (+2 °C above ambient) and decreased soil water availability on net photosynthesis (A_net) and water relations of one‐year old potted loblolly pine (Pinus taeda L.) seedlings grown in treatment chambers with high fertility at three sites along a north‐south transect covering a large portion of the species native range. At each location (Blairsville, Athens and Tifton, GA) we constructed four treatment chambers and randomly assigned each chamber one of four treatments: ambient [CO₂] and ambient temperature, elevated [CO₂] and ambient temperature, ambient [CO₂] and elevated temperature, or elevated [CO₂] and elevated temperature. Within each chamber half of the seedlings were well watered and half received much less water (1/4 that of the well watered). Measurements of net photosynthesis (A_net), stomatal conductance (g_s), leaf water potential and leaf fluorescence were made in June and September, 2008. We observed a significant increase in A_net in response to elevated [CO₂] regardless of site or temperature treatment in June and September. An increase in air temperature of over 2 °C had no significant effect on A_net at any of the sites in June or September despite over a 6 °C difference in mean annual temperature between the sites. Decreased water availability significantly reduced A_net in all treatments at each site in June. The effects of elevated [CO₂] and temperature on g_s followed a similar trend. The temperature, [CO₂] and water treatments did not significantly affect leaf water potential or chlorophyll fluorescence. Our findings suggest that predicted increases in [CO₂] will significantly increase A_net, while predicted increases in air temperature will have little effect on A_net across the native range of loblolly pine. Potential decreases in precipitation will likely cause a significant reduction in A_net, though this may be mitigated by increased [CO₂].     

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

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

Stomatal acclimation over a subambient to elevated CO2 gradient in a C3/C4 grassland

An investigation to determine whether stomatal acclimation to [CO₂] occurred in C₃/C₄ grassland plants grown across a range of [CO₂] (200–550 µmol mol^?1) in the field was carried out. Acclimation was assessed by measuring the response of stomatal conductance (g_s) to a range of intercellular CO₂ (a g_s–C_i curve) at each growth [CO₂] in the third and fourth growing seasons of the treatment. The g_s–C_i response curves for Solanum dimidiatum (C₃ perennial forb) differed significantly across [CO₂] treatments, suggesting that stomatal acclimation had occurred. Evidence of non–linear stomatal acclimation to [CO₂] in this species was also found as maximum g_s (g_smax; g_s measured at the lowest C_i) increased with decreasing growth [CO₂] only below 400 µmol mol^?1. The substantial increase in g_s at subambient [CO₂] for S. dimidiatum was weakly correlated with the maximum velocity of carboxylation (V_cmax; r² = 0·27) and was not associated with CO₂ saturated photosynthesis (A_max). The response of g_s to C_i did not vary with growth [CO₂] in Bromus japonicus (C₃ annual grass) or Bothriochloa ischaemum (C₄ perennial grass), suggesting that stomatal acclimation had not occurred in these species. Stomatal density, which increased with rising [CO₂] in both C₃ species, was not correlated with g_s. Larger stomatal size at subambient [CO₂], however, may be associated with stomatal acclimation in S. dimidiatum. Incorporating stomatal acclimation into modelling studies could improve the ability to predict changes in ecosystem water fluxes and water availability with rising CO₂ and to understand their magnitudes relative to the past.     

Chloride as a macronutrient increases water‐use efficiency by anatomically driven reduced stomatal conductance and increased mesophyll diffusion to CO2

Chloride (Cl^?) has been recently described as a beneficial macronutrient, playing specific roles in promoting plant growth and water‐use efficiency (WUE). However, it is still unclear how Cl^? could be beneficial, especially in comparison with nitrate (NO₃^?), an essential source of nitrogen that shares with Cl^? similar physical and osmotic properties, as well as common transport mechanisms. In tobacco plants, macronutrient levels of Cl^? specifically reduce stomatal conductance (g_s) without a concomitant reduction in the net photosynthesis rate (A_N). As stomata‐mediated water loss through transpiration is inherent in the need of C₃ plants to capture CO₂, simultaneous increase in photosynthesis and WUE is of great relevance to achieve a sustainable increase in C₃ crop productivity. Our results showed that Cl^?‐mediated stimulation of larger leaf cells leads to a reduction in stomatal density, which in turn reduces g_s and water consumption. Conversely, Cl^? improves mesophyll diffusion conductance to CO₂ (g_m) and photosynthetic performance due to a higher surface area of chloroplasts exposed to the intercellular airspace of mesophyll cells, possibly as a consequence of the stimulation of chloroplast biogenesis. A key finding of this study is the simultaneous improvement of A_N and WUE due to macronutrient Cl^? nutrition. This work identifies relevant and specific functions in which Cl^? participates as a beneficial macronutrient for higher plants, uncovering a sustainable approach to improve crop yield.     

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.     

Will intra‐specific differences in transpiration efficiency in wheat be maintained in a high CO2 world? A FACE study

This study evaluates whether the target breeding trait of superior leaf level transpiration efficiency is still appropriate under increasing carbon dioxide levels of a future climate using a semi‐arid cropping system as a model. Specifically, we investigated whether physiological traits governing leaf level transpiration efficiency, such as net assimilation rates (A_net), stomatal conductance (g_s) or stomatal sensitivity were affected differently between two Triticum aestivum L. cultivars differing in transpiration efficiency (cv. Drysdale, superior; cv. Hartog, low). Plants were grown under Free Air Carbon dioxide Enrichment (FACE, approximately 550 µmol mol^?1 or ambient CO₂ concentrations (approximately 390 µmol mol^?1). Mean A_net (approximately 15% increase) and g_s (approximately 25% decrease) were less affected by elevated [CO₂] than previously found in FACE‐grown wheat (approximately 25% increase and approximately 32% decrease, respectively), potentially reflecting growth in a dry‐land cropping system. In contrast to previous FACE studies, analyses of the Ball et al. model revealed an elevated [CO₂] effect on the slope of the linear regression by 12% indicating a decrease in stomatal sensitivity to the combination of [CO₂], photosynthesis rate and humidity. Differences between cultivars indicated greater transpiration efficiency for Drysdale with growth under elevated [CO₂] potentially increasing the response of this trait. This knowledge adds valuable information for crop germplasm improvement for future climates.     

Elevated CO2 affects photosynthetic responses in canopy pine and subcanopy deciduous trees over 10 years: a synthesis from Duke FACE

Leaf responses to elevated atmospheric CO₂ concentration (C_a) are central to models of forest CO₂ exchange with the atmosphere and constrain the magnitude of the future carbon sink. Estimating the magnitude of primary productivity enhancement of forests in elevated C_a requires an understanding of how photosynthesis is regulated by diffusional and biochemical components and up‐scaled to entire canopies. To test the sensitivity of leaf photosynthesis and stomatal conductance to elevated C_a in time and space, we compiled a comprehensive dataset measured over 10 years for a temperate pine forest of Pinus taeda, but also including deciduous species, primarily Liquidambar styraciflua. We combined over one thousand controlled‐response curves of photosynthesis as a function of environmental drivers (light, air C_a and temperature) measured at canopy heights up to 20 m over 11 years (1996–2006) to generate parameterizations for leaf‐scale models for the Duke free‐air CO₂ enrichment (FACE) experiment. The enhancement of leaf net photosynthesis (A_net) in P. taeda by elevated C_a of +200 μmol mol^?1 was 67% for current‐year needles in the upper crown in summer conditions over 10 years. Photosynthetic enhancement of P. taeda at the leaf‐scale increased by two‐fold from the driest to wettest growing seasons. Current‐year pine foliage A_net was sensitive to temporal variation, whereas previous‐year foliage A_net was less responsive and overall showed less enhancement (+30%). Photosynthetic downregulation in overwintering upper canopy pine needles was small at average leaf N (N_area), but statistically significant. In contrast, co‐dominant and subcanopy L. styraciflua trees showed A_net enhancement of 62% and no A_net–N_area adjustments. Various understory deciduous tree species showed an average A_net enhancement of 42%. Differences in photosynthetic responses between overwintering pine needles and subcanopy deciduous leaves suggest that increased C_a has the potential to enhance the mixed‐species composition of planted pine stands and, by extension, naturally regenerating pine‐dominated stands.     

Will photosynthesis of maize (Zea mays) in the US Corn Belt increase in future [CO2] rich atmospheres? An analysis of diurnal courses of CO2 uptake under free‐air concentration enrichment (FACE)

The C₄ grass Zea mays (maize or corn) is the third most important food crop globally in terms of production and demand is predicted to increase 45% from 1997 to 2020. However, the effects of rising [CO₂] upon C₄ plants, and Z. mays specifically, are not sufficiently understood to allow accurate predictions of future crop production. A rainfed, field experiment utilizing free‐air concentration enrichment (FACE) technology in the primary area of global corn production (US Corn Belt) was undertaken to determine the effects of elevated [CO₂] on corn. FACE technology allows experimental treatments to be imposed upon a complete soil–plant–atmosphere continuum with none of the effects of experimental enclosures on plant microclimate. Crop performance was compared at ambient [CO₂] (354 μ mol mol^?1) and the elevated [CO₂] (549 μmol mol^?1) predicted for 2050. Previous laboratory studies suggest that under favorable growing conditions C₄ photosynthesis is not typically enhanced by elevated [CO₂]. However, stomatal conductance and transpiration are decreased, which can indirectly increase photosynthesis in dry climates. Given the deep soils and relatively high rainfall of the US Corn Belt, it was predicted that photosynthesis would not be enhanced by elevated [CO₂]. The diurnal course of gas exchange of upper canopy leaves was measured in situ across the growing season of 2002. Contrary to the prediction, growth at elevated [CO₂] significantly increased leaf photosynthetic CO₂ uptake rate (A) by up to 41%, and 10% on average. Greater A was associated with greater intercellular [CO₂], lower stomatal conductance and lower transpiration. Summer rainfall during 2002 was very close to the 50‐year average for this site, indicating that the year was not atypical or a drought year. The results call for a reassessment of the established view that C₄ photosynthesis is insensitive to elevated [CO₂] under favorable growing conditions and that the production potential of corn in the US Corn Belt will not be affected by the global rise in [CO₂].