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.     

Modelling Diurnal Courses of Photosynthesis and Transpiration of Leaves on the Basis of Stomatal and Non-Stomatal Responses,Including Photoinhibition

A mathematical model for photoinhibition of leaf photosynthesis was developed by formalising the assumptions that (1) the rate of photoinhibition is proportional to irradiance; and (2) the rate of recovery, derived from the formulae for a pseudo first-order process, is proportional to the extent of inhibition. The photoinhibition model to calculate initial photo yield is integrated into a photosynthesis-stomatal conductance (g _s) model that combines net photosynthetic rate (P _N), transpiration rate (E), and g _s, and also the leaf energy balance. The model was run to simulate the diurnal courses of P _N, E, g _s, photochemical efficiency, i.e., ratio of intercellular CO₂ concentration and CO₂ concentration over leaf surface (C _i/C _s), and leaf temperature (T ₁) under different irradiances, air temperature, and humidity separately with fixed time courses of others. When midday depression occurred under high temperature, g _s decreased the most and E the least. The duration of midday depression of g _s was the longest and that in E the shortest. E increased with increasing vapour pressure deficit (VPD) initially, but when VPD exceeded a certain value, it decreased with increasing VPD; this was caused by a rapid decrease in g _s. When air temperature exceeded a certain value, an increase in solar irradiance raised T ₁ and the degree of midday depression. High solar radiation caused large decrease in initial photon efficiency (). P _N, E, and g _s showed reasonable decreases under conditions causing photoinhibition compared with non-photoinhibition condition under high irradiance. The T ₁ under photoinhibition was higher than that under non-photoinhibition conditions, which was evident under high solar irradiance around noon. The decrease in C _i/C _s at midday implies that stomatal closure is a factor causing midday depression of photosynthesis.     

Modelling stomatal conductanceof field-grown sunflower under varying soil water content and leafenvironment: comparison of three models of stomatal response toleaf environment and coupling with an abscisic acid-based modelof stomatal response to soil drying

Stomatal conductance, g_s, responds both tothe immediate or local environment of the leaf, such as CO₂ partialpressure and irradiance, and to root‐sourced signals of water stress,particularly abscisic acid (ABA). Two models for the combined controlof g_s were formulated and tested in sunflower(Helianthus annuus). First, several empirical models weretested for the local control, demonstrating that the Ball–Berrymodel [Ball, Woodrow & Berry (in Progress in PhotosynthesisResearch Vol. 4, pp. 5.221–5.224: M. Nijhoff,Dordrecht, The Netherlands) 1987] is consistently amongthe most accurate. A problem of statistical non‐independence inthis model is shown to be minor. The model offers regularity ofparameter values among most species and, despite an oversimplicationin representing known humidity‐response mechanisms, it incorporates othersignalling loops from CO₂ and assimilation. In the firstcombined model, ABA as its concentration in xylem sap, [ABA]_xy,down‐regulates the slope, m, in the Ball–Berry modelby the factor g_fac = exp(– β[ABA]_xy).The ABA‐induced reduction in g_s decreases CO₂ assimilation andsurface humidity, thus appearing to induce the local‐control mechanismto amplify the ABA‐induced stomatal closure. In the second combinedmodel, g_s is estimated as the minimum of the local(Ball–Berry) response and the product g_fac g_s,max,with g_s,max as a maximal unstressed conductance.Both models can predict g_s from the external environmentalvariables with good accuracy (r² near 0·8 over20‐fold variations in g_s). Further analyses showthat g_s responds to humidity almost quadraticallyrather than linearly. It also responds to assimilation as a powerlaw with an exponent that is significantly less than 1. These limitations,shared by other models, suggest more research into biochemical signalling.     

Contrasting stomatal sensitivity to temperature and soil drought in mature alpine conifers

Conifers growing at high elevations need to optimize their stomatal conductance (g_s) for maximizing photosynthetic yield while minimizing water loss under less favourable thermal conditions. Yet the ability of high‐elevation conifers to adjust their g_s sensitivity to environmental drivers remains largely unexplored. We used 4 years of sap flow measurements to elucidate intraspecific and interspecific variability of g_s in Larix decidua Mill. and Picea abies (L.) Karst along an elevational gradient and contrasting soil moisture conditions. Site‐ and species‐specific g_s response to main environmental drivers were examined, including vapour pressure deficit, air temperature, solar irradiance, and soil water potential. Our results indicate that maximum g_s of L. decidua is >2 times higher, shows a more plastic response to temperature, and down‐regulates g_s stronger during atmospheric drought compared to P. abies. These differences allow L. decidua to exert more efficient water use, adjust to site‐specific thermal conditions, and reduce water loss during drought episodes. The stronger plasticity of g_s sensitivity to temperature and higher conductance of L. decidua compared to P. abies provide new insights into species‐specific water use strategies, which affect species' performance and should be considered when predicting terrestrial water dynamics under future climatic change.     

Factors influencing stomatal conductance in response to water availability in grapevine: a meta‐analysis

The main factors regulating grapevine response to decreasing water availability were assessed under statistical support using published data related to leaf water relations in an extensive range of scion and rootstock genotypes. Matching leaf water potential (Ψ_leaf) and stomatal conductance (g_s) data were collected from peer‐reviewed literature with associated information. The resulting database contained 718 data points from 26 different Vitis vinifera varieties investigated as scions, 15 non‐V. vinifera rootstock genotypes and 11 own‐rooted V. vinifera varieties. Linearised data were analysed using the univariate general linear model (GLM) with factorial design including biological (scion and rootstock genotypes), methodological and environmental (soil) fixed factors. The first GLM performed on the whole database explained 82.4% of the variability in data distribution having the rootstock genotype the greatest contribution to variability (19.1%) followed by the scion genotype (16.2%). A classification of scions and rootstocks according to their mean predicted g_s in response to moderate water stress was generated. This model also revealed that g_s data obtained using a porometer were in average 2.1 times higher than using an infra‐red gas analyser. The effect of soil water‐holding properties was evaluated in a second analysis on a restricted database and showed a scion‐dependant effect, which was dominant over rootstock effect, in predicting g_s values. Overall the results suggest that a continuum exists in the range of stomatal sensitivities to water stress in V. vinifera, rather than an isohydric–anisohydric dichotomy, that is further enriched by the diversity of scion‐rootstock combinations and their interaction with different soils.     

Mechanisms for minimizing height‐related stomatal conductance declines in tall vines

The ability to transport water through tall stems hydraulically limits stomatal conductance (g_s), thereby constraining photosynthesis and growth. However, some plants are able to minimize this height‐related decrease in g_s, regardless of path length. We hypothesized that kudzu (Pueraria lobata) prevents strong declines in g_s with height through appreciable structural and hydraulic compensative alterations. We observed only a 12% decline in maximum g_s 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 g_s 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.     

Seasonal variability of the parameters of the Ball–Berry model of stomatal conductance in maize (Zea mays L.) and sunflower (Helianthus annuus L.) under well‐watered and water‐stressed conditions

The Ball–Berry (BB) model of stomatal conductance (g_s) is frequently coupled with a model of assimilation to estimate water and carbon exchanges in plant canopies. The empirical slope (m) and ‘residual’ g_s (g₀) parameters of the BB model influence transpiration estimates, but the time‐intensive nature of measurement limits species‐specific data on seasonal and stress responses. We measured m and g₀ seasonally and under different water availability for maize and sunflower. The statistical method used to estimate parameters impacted values nominally when inter‐plant variability was low, but had substantial impact with larger inter‐plant variability. Values for maize (m = 4.53 ± 0.65; g₀ = 0.017 ± 0.016 mol m^?2 s^?1) were 40% higher than other published values. In maize, we found no seasonal changes in m or g₀, supporting the use of constant seasonal values, but water stress reduced both parameters. In sunflower, inter‐plant variability of m and g₀ was large (m = 8.84 ± 3.77; g₀ = 0.354 ± 0.226 mol m^?2 s^?1), presenting a challenge to clear interpretation of seasonal and water stress responses – m values were stable seasonally, even as g₀ values trended downward, and m values trended downward with water stress while g₀ values declined substantially.     

Stomatal dynamics are limited by leaf hydraulics in ferns and conifers: results from simultaneous measurements of liquid and vapour fluxes in leaves

Stomatal responsiveness to vapour pressure deficit (VPD) results in continuous regulation of daytime gas‐exchange directly influencing leaf water status and carbon gain. Current models can reasonably predict steady‐state stomatal conductance (g_s) to changes in VPD but the g_s dynamics between steady‐states are poorly known. Here, we used a diverse sample of conifers and ferns to show that leaf hydraulic architecture, in particular leaf capacitance, has a major role in determining the g_s response time to perturbations in VPD. By using simultaneous measurements of liquid and vapour fluxes into and out of leaves, the in situ fluctuations in leaf water balance were calculated and appeared to be closely tracked by changes in g_s thus supporting a passive model of stomatal control. Indeed, good agreement was found between observed and predicted g_s when using a hydropassive model based on hydraulic traits. We contend that a simple passive hydraulic control of stomata in response to changes in leaf water status provides for efficient stomatal responses to VPD in ferns and conifers, leading to closure rates as fast or faster than those seen in most angiosperms.     

Observations and model simulations link stomatal inhibition to impaired hydraulic conductance following ozone exposure in cotton

Ozone (O₃) inhibits plant gas exchange and productivity. Vapour phase (g_s) and liquid or hydraulic phase (K) conductances to water flux are often correlated as both change with environmental parameters. Exposure of cotton plants to tropospheric O₃ reduces g_s through reversible short-term mechanisms and by irreversible long-term disruption of biomass allocation to roots which reduces K. We hypothesize that chronic effects of O₃ on gas exchange can be mediated by effects on K without a direct effect of O₃ on g_s or carbon assimilation (A). Experimental observations from diverse field and exposure chamber studies, and simulations with a model of mass and energy transport, support this hypothesis. O₃ inhibition of K leads to realistic simulated diurnal courses of g_s that reproduce observations at low ambient O₃ concentration and maintain the positive correlation between midday g_s and K observed experimentally at higher O₃ concentrations. Effects mediated by reduced K may interact with more rapid responses of g_s and A to yield the observed suite of oxidant impacts on vegetation. The model extends these physiological impacts to the extensive canopy scale. Simulated magnitudes and diurnal time courses of canopy-scale fluxes of H₂O and O₃ match observations under low ambient concentrations of O₃. With greater simulated concentrations of O₃ during plant development, the model suggests potential reductions of canopy-scale water fluxes and O₃ deposition. This could represent a potentially unfavourable positive feedback on tropospheric O₃ concentrations associated with biosphere–atmosphere exchange.     

Late-stage canopy tree species with extremely low δ13C and high stomatal sensitivity to seasonal soil drought in the tropical rainforest of French Guiana

We assessed the daily time‐courses of CO₂ assimilation rate (A), leaf transpiration rate (E), stomatal conductance for water vapour (g_s), leaf water potential ( Ψ _w) and tree transpiration in a wet and a dry season for three late‐stage canopy rainforest tree species in French Guiana differing in leaf carbon isotope composition ( δ ¹³C). The lower sunlit leaf δ ¹³C values found in Virola surinamensis ( ? 29·9‰) and in Diplotropis purpurea ( ? 30·9‰), two light‐demanding species, as compared to Eperua falcata ( ? 28·6‰), a shade‐semi‐tolerant species, were clearly associated with higher maximum g_s values of sunlit leaves in the two former species. These two species were also characterized by a high sensitivity of g_s, sap flow density (Ju) and canopy conductance (g_c) to seasonal soil drought, allowing maintenance of high midday Ψ _w values in the dry season. The data for Diplotropis provided an original picture of increasing midday Ψ _w with increasing soil drought. In Virola, stomata were extremely sensitive to seasonal soil drought, leading to a dramatic decrease in leaf and tree transpiration in the dry season, whereas midday Ψ _w remained close to ? 0·3 MPa. The mechanisms underlying such an extremely high sensitivity of stomata to soil drought remain unknown. In Eperua, g_s of sunlit leaves was non‐responsive to seasonal drought, whereas Ju and g_c were lower in the dry season. This suggests a higher stomatal sensitivity to seasonal drought in shaded leaves than in sunlit ones in this species.     

Midday stomatal conductance is more related to stem rather than leaf water status in subtropical deciduous and evergreen broadleaf trees

Midday depressions in stomatal conductance (g_s) and photosynthesis are common in plants. The aim of this study was to understand the hydraulic determinants of midday g_s, the coordination between leaf and stem hydraulics and whether regulation of midday g_s differed between deciduous and evergreen broadleaf tree species in a subtropical cloud forest of Southwest (SW) China. We investigated leaf and stem hydraulics, midday leaf and stem water potentials, as well as midday g_s of co‐occurring deciduous and evergreen tree species. Midday g_s was correlated positively with midday stem water potential across both groups of species, but not with midday leaf water potential. Species with higher stem hydraulic conductivity and greater daily reliance on stem hydraulic capacitance were able to maintain higher stem water potential and higher g_s at midday. Deciduous species exhibited significantly higher stem hydraulic conductivity, greater reliance on stem capacitance, higher stem water potential and g_s at midday than evergreen species. Our results suggest that midday g_s is more associated with midday stem than with leaf water status, and that the functional significance of stomatal regulation in these broadleaf tree species is probably for preventing stem xylem dysfunction.     

Mycorrhizal promotion of host stomatal conductance in relation to irradiance and temperature

Colonization of roots and soil by arbuscular mycorrhizal (AM) fungi sometimes promotes stomatal conductance (g _s) of the host plant, but scientists have had difficulty predicting or manipulating the response. Our objective was to test whether the magnitude of AM influence on g _s is related to environmental conditions: irradiance, air temperature or leaf temperature. Stomatal conductances of two groups of uncolonized sorghum plants were compared to g _s of plants colonized by Glomus intraradices (Gi) or Gigaspora margarita (Gm) in 31 morning and afternoon periods under naturally varying greenhouse conditions. Stomatal conductance of Gi and Gm plants was often markedly higher than g _s of similarly sized nonAM plants. AM promotion of g _s was minimal at the lowest irradiances and lowest air and leaf temperatures, but was substantial at intermediate irradiance and temperatures. AM promotion was again low or absent at the highest irradiances and temperatures. Magnitude of AM promotion of g _s was not a function of absolute g _s. Promotion of g _s by Gi and Gm was remarkably similar. Differing phosphorus fertilization did not affect g _s.     

Stomatal acclimation to vapour pressure deficit doubles transpiration of small tree seedlings with warming

Future climate change is expected to increase temperature (T) and atmospheric vapour pressure deficit (VPD) in many regions, but the effect of persistent warming on plant stomatal behaviour is highly uncertain. We investigated the effect of experimental warming of 1.9–5.1 °C and increased VPD of 0.5–1.3 kPa on transpiration and stomatal conductance (g_s) of tree seedlings in the temperate forest understory (Duke Forest, North Carolina, USA). We observed peaked responses of transpiration to VPD in all seedlings, and the optimum VPD for transpiration (D_opt) shifted proportionally with increasing chamber VPD. Warming increased mean water use of Carya by 140% and Quercus by 150%, but had no significant effect on water use of Acer. Increased water use of ring‐porous species was attributed to (1) higher air T and (2) stomatal acclimation to VPD resulting in higher g_s and more sensitive stomata, and thereby less efficient water use. Stomatal acclimation maintained homeostasis of leaf T and carbon gain despite increased VPD, revealing that short‐term stomatal responses to VPD may not be representative of long‐term exposure. Acclimation responses differ from expectations of decreasing g_s with increasing VPD and may necessitate revision of current models based on this assumption.     

Most stomatal closure in woody species under moderate drought can be explained by stomatal responses to leaf turgor

Reduced stomatal conductance (g_s) during soil drought in angiosperms may result from effects of leaf turgor on stomata and/or factors that do not directly depend on leaf turgor, including root‐derived abscisic acid (ABA) signals. To quantify the roles of leaf turgor‐mediated and leaf turgor‐independent mechanisms in g_s decline during drought, we measured drought responses of g_s and water relations in three woody species (almond, grapevine and olive) under a range of conditions designed to generate independent variation in leaf and root turgor, including diurnal variation in evaporative demand and changes in plant hydraulic conductance and leaf osmotic pressure. We then applied these data to a process‐based g_s model and used a novel method to partition observed declines in g_s during drought into contributions from each parameter in the model. Soil drought reduced g_s by 63–84% across species, and the model reproduced these changes well (r² = 0.91, P < 0.0001, n = 44) despite having only a single fitted parameter. Our analysis concluded that responses mediated by leaf turgor could explain over 87% of the observed decline in g_s across species, adding to a growing body of evidence that challenges the root ABA‐centric model of stomatal responses to drought.     

The response of stomatal conductance to seasonal drought in tropical forests

Stomata regulate CO₂ uptake for photosynthesis and water loss through transpiration. The approaches used to represent stomatal conductance (g_s) in models vary. In particular, current understanding of drivers of the variation in a key parameter in those models, the slope parameter (i.e. a measure of intrinsic plant water‐use‐efficiency), is still limited, particularly in the tropics. Here we collected diurnal measurements of leaf gas exchange and leaf water potential (Ψ_leaf), and a suite of plant traits from the upper canopy of 15 tropical trees in two contrasting Panamanian forests throughout the dry season of the 2016 El Niño. The plant traits included wood density, leaf‐mass‐per‐area (LMA), leaf carboxylation capacity (V_c,max), leaf water content, the degree of isohydry, and predawn Ψ_leaf. We first investigated how the choice of four commonly used leaf‐level g_s models with and without the inclusion of Ψ_leaf as an additional predictor variable influence the ability to predict g_s, and then explored the abiotic (i.e. month, site‐month interaction) and biotic (i.e. tree‐species‐specific characteristics) drivers of slope parameter variation. Our results show that the inclusion of Ψ_leaf did not improve model performance and that the models that represent the response of g_s to vapor pressure deficit performed better than corresponding models that respond to relative humidity. Within each g_s model, we found large variation in the slope parameter, and this variation was attributable to the biotic driver, rather than abiotic drivers. We further investigated potential relationships between the slope parameter and the six available plant traits mentioned above, and found that only one trait, LMA, had a significant correlation with the slope parameter (R² = 0.66, n = 15), highlighting a potential path towards improved model parameterization. This study advances understanding of g_s dynamics over seasonal drought, and identifies a practical, trait‐based approach to improve modeling of carbon and water exchange in tropical forests.     

Effects of light history on primary productivity in a phytoplankton community dominated by the toxic cyanobacterium Cylindrospermopsis raciborskii

1. The effects of instantaneous irradiance and short‐term light history on primary production were determined for samples from a subtropical water reservoir dominated by the toxic cyanobacterium Cylindrospermopsis raciborskii. ¹⁴C‐bicarbonate uptake incubations were conducted on water samples from the reservoir, for irradiance (photosynthetically active radiation) ranging from 0 to 1654 μmol quanta m⁻² s⁻¹. Prior to the ¹⁴C incubations, cells were pre‐treated at irradiance levels ranging from 0 to 1006 μmol quanta m⁻² s⁻¹. 2. The average irradiance experienced by cells during the 2–2.5 h pre‐treatment incubations affected the productivity–irradiance (P–I) parameters: exposure to high light in pre‐treatment conditions caused a substantial decrease in maximum rate of primary production P_max and the photoinhibition parameter β when compared to cells pre‐treated in the dark. 3. While the data collected in this study were not sufficient to develop a full dynamic model of C. raciborskii productivity, P_max and β were modelled as a function of pre‐treatment irradiance, and these models were applied to predict the rate of primary production as a function of both instantaneous and historical irradiance. The results indicated that while cells with a history of exposure to high irradiance will be the most productive in high irradiance, production rates will be highest overall for dark‐acclimated cells in moderate irradiance. 4. Our results may explain why optically‐deep mixing favours C. raciborskii. If the mixing depth z_m exceeds the euphotic depth z_eu, cells will be dark‐acclimated, which will increase their rate of production when they are circulated through the euphotic zone. These results also predict that production rates will be higher during morning hours than for the same irradiance in the afternoon, which is consistent with other phytoplankton studies. 5. Since the rate of production of C. raciborskii‐dominated systems cannot be described by a single P–I curve, accurate estimates of production rates will require measurements over the daily light cycle.     

Rates of stomatal opening in conifer seedlings in relation to air temperature and daily carbon gain

Experiments were conducted on well watered 1-year-old Douglas fir [Pseudotsuga menziesii (Mirb.) Franco], western hemlock [Tsuga heterophylla (Raf.) Sarg.] and western redcedar (Thuja plicata Donn) seedlings to determine the effects of temperature on whole-plant photosynthetic and stomatal responses to short-term fluctuations in irradiance (Q). Following a step change in Q, time constants (τ, the period over which 63% of the total change occurs) for stomatal conductance (g_s) and assimilation rate (A) decreased linearly with increasing air temperature (T_air). For example, in western redcedar τ_A decreased from 30 ± 4 min at 5 °C to 10 ± 1 min at 25 °C. In all cases, τ_A was within 10–15% of τ_gs. There was considerable variation in τ among individuals within a given species. Differences between species became more pronounced with decreasing temperature. Multiplicative models that included functions for τ accounted for 99% of the diurnal variability in A and g_s for seedlings exposed to varying air temperature, irradiance and vapour pressure deficit. Estimates of daily A were within 2% of those measured. Intermittent cloud cover and understory shading were approximated by exposing seedlings to 3–4 episodes (≥1 h) of shade (200 or 500 μmol m^?2 s^?1) or complete darkness during the day. In such cases, daily A was overestimated by up to 4 and 21%, respectively, if a function for τ was excluded from the models. Our results suggest that there is scope for selecting seedling stock for increased carbon assimilation on the basis of reduced time constants. For example, in western redcedar, a 40% reduction in τ could lead to increases in daily carbon gains of almost 5% depending on the frequency and degree of shading. If these daily gains were translated into increased dry matter production and compounded, seasonal gains would be even larger.     

A critical appraisal of a combined stomatal-photosynthesis model for C3 plants

Gas-exchange measurements on Eucalyptus grandis leaves and data extracted from the literature were used to test a semi-empirical model of stomatal conductance for CO₂ g_Sc=g_o+a₁A/(c_s-I) (1+D_s/D_o)] where A is the assimilation rate; D_s and c_s are the humidity deficit and the CO₂ concentration at the leaf surface, respectively; g₀ is the conductance as A → 0 when leaf irradiance → 0; and D₀ and a₁ are empirical coefficients. This model is a modified version of g_sc=a₁A h_s/c_s first proposed by Ball, Woodrow & Berry (1987, in Progress in Photosynthesis Research, Martinus Mijhoff, Publ., pp. 221–224), in which h_s is relative humidity. Inclusion of the CO₂ compensation point, τ, improved the behaviour of the model at low values of c_s, while a hyperbolic function of D_s for humidity response correctly accounted for the observed hyperbolic and linear variation of g_sc and c_i/c_s as a function of D_s, where C_i is the intercellular CO₂ concentration. In contrast, use of relative humidity as the humidity variable led to predictions of a linear decrease in g_sc and a hyperbolic variation in c_i/c_s as a function of D_s, contrary to data from E. grandis leaves. The revised model also successfully described the response of stomata to variations in A, D_s and c_s for published responses of the leaves of several other species. Coupling of the revised stomatal model with a biochemical model for photosynthesis of C₃ plants synthesizes many of the observed responses of leaves to light, humidity deficit, leaf temperature and CO₂ concentration. Best results are obtained for well-watered plants.     

Stomatal control by both [ABA] in the xylem sap and leaf water status: a test of a model for draughted or ABA-fed field-grown maize

A model of maize stomatal behaviour has been developed, in which stomatal conductance is linked to the concentration of abscisic acid ([ABA]) in the xylem sap, with a sensitivity dependent upon the leaf water potential (Ψ₁). It was tested against two alternative hypotheses, namely that stomatal sensitivity to xylem [ABA] would be linked to the leaf-to-air vapour pressure difference (VPD), or to the flux of ABA into the leaf. Stomatal conductance (g_s) was studied: (1) in field-grown plants whose xylem [ABA] and Ψ₁ depended on soil water status and evaporative demand; (2) in field-grown plants fed with ABA solutions such that xylem [ABA] was artificially raised, thereby decreasing g_s and increasing Ψ₁ and leaf-to-air VPD; and (3) in ABA-fed detached leaves exposed to varying evaporative demands, but with a constant and high Ψ₁. The same relationships between g_s, xylem [ABA] and Ψ₁, showing lower stomatal sensitivity to [ABA] at high Ψ₁, applied whether variations in xylem [ABA] were due to natural increase or to feeding, and whether variations in Ψ₁, were due to changes in evaporative demand or to the increased Ψ₁ observed in ABA-fed plants. Conversely, neither the leaf-to-air VPD nor the ABA flux into the leaf accounted for the observed changes in stomatal sensitivity to xylem [ABA]. The model, using parameters calculated from previous field data and the detached-leaf data, was tested against the observations of both ABA-fed and droughted plants in the field. It accounted with reasonable accuracy for changes in g_s (r² ranging from 0.77 to 0.81). These results support the view that modelling of stomatal behaviour requires consideration of both chemical and hydraulic aspects of root-to-shoot communication.     

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.