Stomatal response to air humidity and its relation to stomatal density in a wide range of warm climate species

The gas exchange of 19 widely different warm climate species was observed at different leaf to air vapour pressure deficits (VPD). In all species stomata tended to close as VPD increased resulting in a decrease in net photosynthesis. The absolute reduction in leaf conductance per unit increase in VPD was greatest in those species which had a large leaf conductance at low VPDs. This would be expected even if stomata of all species were equally sensitive. However the percentage reduction in net photosynthesis (used as a measure of the relative sensitivity of stomata of the different species) was also closely related to the maximal conductance at low VPD. Similarily the relative sensitivity of stomata to changes in VPD was closely related to the weighted stomatal density or crowding index.The hypothesis is presented that stomatal closure at different VPDs is related to peristomatal evaporation coupled with a high resistance between the epidermis and the mesophyll and low resistance between the stomatal apparatus and the epidermal cells. This hypothesis is consistent with the greater relative sensitivity of stomata on leaves with a high crowding index.The results and the hypothesis are discussed in the light of selection, for optimal productivity under differing conditions of relative humidity and soil water availablility, by observation of stomatal density and distribution on the two sides of the leaf.Visiting scientist, plant physiologist and research assitant of the Cassava Program     

Stomatal responses to long-term high vapor pressure deficits mediated most limitation of photosynthesis in tomatoes

Plants grown at high vapor pressure deficit (VPD) usually present decreased photosynthesis, but stomatal and mesophyll limitation to photosynthesis remain poorly quantified. To better understand the regulation of high VPD on photosynthesis and plant growth in tomatoes, we investigated the limitation of stomatal conductance and mesophyll conductance to photosynthesis and relative importance of stomatal morphology and function in stomatal conductance. Both the net photosynthesis rate and total biomass were significantly limited by high VPD. Meanwhile, stomatal conductance and mesophyll conductance were decreased under high VPD. The stomatal conductance limitation was responsible for 60% of the total photosynthetic limitation. Moreover, a reduction in stomatal density and stomatal size occurred under high VPD, which was significantly correlated with the down-regulation of stomatal conductance. The stomatal morphology contributed to more than half the change in stomatal conductance. Nevertheless, stomatal movement was also an important factor in regulating stomatal conductance. The decrease of hydraulic conductance and transpiration rate with no significant difference in relative water content, leaf water potential, and/or osmotic potential suggested passive hydraulic regulation in the feedforward responses of stomata to high VPD.     

The Hydraulic Pathways in Nicotiana glauca (Grah.) and Tradescantia virginiana (L.) Leaves, and Water Potentials in Leaf Epidermes

The hydraulic conductances of leaves of a species which exhibitsstomatal responses to humidity (Nicotiana glauca) are significantlylower than the conductances in a species which does not exhibitsuch responses (Tradescantia virginiana). This difference couldat least partly account for their difference in stomatal responseto humidity. In both species, the hydraulic conductance betweenthe leaf bulk and its epidermis is much lower than the conductancein any other part of the pathway. The apparently conflictingresults, reported in recent literature, on the hydraulic conductancesand water pathways in leaves are reinterpreted, and shown tobe due to misinterpretation of results. The recently publishedcriticisms of a technique used to measure hydraulic conductivityare commented on and refuted. An examination of the factors that influence the water potentialat the sites of evaporation from the inner walls of the epidermisnear stomatal pores showed that the water potential at thesesites is lower than the bulk epidermal water potential. Thewater potential at these sites changes in a complex way as stomatalaperture changes. As it is reduced the ratio of: ‘waterpotential at sites of evaporation on the inner walls of theepidermis near stomatal pores/bulk leaf water potential‘increases. The positive feedback effect of this phenomenon,which tends to keep stomatal water potential constant as thestomata close and therefore enhances closure, and two other‘passive’ positive feedback effects on the waterpotential at sites of evaporation near stomata that have beenreported in the literature are briefly discussed. Nicotiana glauca (Grah.), Tradescantia virginiana (L.), sub-stomatal cavities, peristomatal evaporation, stomata, humidity response, leaf hydraulic conductance, water potential     

Acclimation to humidity modifies the link between leaf size and the density of veins and stomata

The coordination of veins and stomata during leaf acclimation to sun and shade can be facilitated by differential epidermal cell expansion so large leaves with low vein and stomatal densities grow in shade, effectively balancing liquid‐ and vapour‐phase conductances. As the difference in vapour pressure between leaf and atmosphere (VPD) determines transpiration at any given stomatal density, we predict that plants grown under high VPD will modify the balance between veins and stomata to accommodate greater maximum transpiration. Thus, we examined the developmental responses of these traits to contrasting VPD in a woody angiosperm (Toona ciliata M. Roem.) and tested whether the relationship between them was altered. High VPD leaves were one‐third the size of low VPD leaves with only marginally greater vein and stomatal density. Transpirational homeostasis was thus maintained by reducing stomatal conductance. VPD acclimation changed leaf size by modifying cell number. Hence, plasticity in vein and stomatal density appears to be generated by plasticity in cell size rather than cell number. Thus, VPD affects cell number and leaf size without changing the relationship between liquid‐ and vapour‐phase conductances. This results in inefficient acclimation to VPD as stomata remain partially closed under high VPD.     

The influence of plant water stress on stomatal control of gas exchange at different levels of atmospheric humidity

Summary Leaves of well-watered and mildly water-stressed seedlings of Betula pendula Roth. and Gmelina aroborea L. were subjected to a range of vapour pressure deficits (VPD) between 10 and 24 kPa. The stomatal conductance of birch seedlings decreased as VPD was increased and at least in mildly-stressed seedlings this response seemed to be closely linked to the water status of the air rather than to the bulk water status of the plant. Mild water stressing enhanced the degree of the stomatal humidity-response and resulted in a significant increase in the efficiency of water use at high VPD. Stomata of Gmelina were apparently insensitive to variation in VPD, but were more sensitive to a decrease in bulk leaf water status than were stomata of birch. Water use efficiency of Gmelina seedlings was comparatively high, even when VPD was high and the stomata were fully open.     

Epidermal transpiration and stomatal responses to humidity: Some hypotheses explored

Abstract Stomatal responses to humidity as affected by both evaporation from the epidermis and the hydraulic conductance of the transpiration stream to evaporation sites on the epidermis are discussed.
Recent estimates of evaporation from the inner walls of the epidermis are too high because the cell wall surfaces were assumed completely wet, and leaves have usually been considered isothermal.
It is suggested that a fall in humidity increases evaporation from the epidermis, and that stomata respond to the consequent fall in water potential. Cuticular transiration is inversely related to stomatal conductance. Thus, evaporation from the epidermis is dependent on the stomatal, boundary layer, and cuticular conductances, and on evaporation from the inner walls of the epidermis. Stomatal responses to humidity will change as the boundary layer conductance changes.
The conductance of the transpiration stream is a determinant of the water potential of the epidermis. Water potentials of adjacent cells will be more similar if flow is symplastic than if it is apoplastic. It is concluded that flow in living tissues is primarily symplastic over long distances, but over shorter distances it is increasingly apoplastic, and that stomatal responses to humidity are mediated by the water potential of the whole epidermis.     

Differential response of stomata to air humidity in the parasitic mistletoe (Phthirusa pyrifolia) and its host,mandarin orange (Citrus resitulata)

Measurements of CO₂ and H₂O exchange rate and the calculated leaf conductance of attached leaves were conducted over a range of leaf-to-air vapour pressure difference (VPD) (1.5 to 5.5 kPa) to compare the response of the parasitic mistletoe, Phthirusa pyrifolia, with that of its host, the mandarin orange, Citrus reticulata. Seedlings of the host infected with the parasite were grown in well-watered and adequately fertilized large pots outdoors at the CIAT headquarters, Palmira, Colombia, South America. Observations of leaf anatomy of the parasite and nutrient analysis of young tissues of both the parasite and host were made. The photosynthetic rate of the host decreased linearly with increased VPD, whereas the parasite showed a constant rate. This trend coincided with similar responses in leaf conductance. Due to the insensitivity of the parasite stomata, the transpiration rate increased linearly with VPD as compared with an initial increase and then a decrease in the host transpiration rate. The higher photosynthetic rate and the closure of stomata of the host resulted in high water use efficiency as compared with that of the parasite. The parasite accumulated in its leaves more N, P, K and less Ca and Mg than the host. The significance of the host-parasite differential response to air humidity is discussed in relation to mechanism underlying stomatal sensitivity and in the context of host-parasite association.Visiting Scientist, Coordinator, and Research Assistant of the Cassava Physiology Program.     

The magnitude of the stomatal response to blue light : modulation by atmospheric humidity

The effect of leaf-air vapor pressure difference (VPD) on the magnitude of the stomatal response to blue light was investigated in soybean (Glycine max) by administering blue light pulses (22 seconds by 120 micromoles per square meter per second) at different levels of VPD and temperature. At 20 °C and 25 °C, the magnitude of the integrated conductance response decreased with increasing VPD (0.4 to 2.6 kiloPascals), due to an earlier onset of stomatal closure that terminated the pulse response. In contrast, at 30 °C this magnitude increased with rising VPD (0.9 to 3.5 kiloPascals), due to an increasing maximum excursion of the conductance response despite the accelerated onset of stomatal closure. When the feedforward response of stomata to humidity caused steady state transpiration to decrease with increasing VPD, the magnitude of the pulse-induced conductance response correlated with VPD rather than with transpiration. This suggests that water relations or metabolite movements within epidermal rather than bulk leaf tissue interacted with guard cell photobiological properties in regulating the magnitude of the blue light response. VPD modulation of pulse magnitude could reduce water loss during stomatal responses to transient illumination in natural light environments.     

Nonstomatal inhibition of photosynthesis by water stress. Reduction in photosynthesis at high transpiration rate without stomatal closure in field-grown tomato

Large underestimates of the limitation to photosynthesis imposed by stomata can occur because of an error in the standard method of calculating average substomatal pressures of carbon dioxide when heterogeneity of those pressures occurs across a leaf surface. Most gas exchange data supposedly indicating nonstomatal inhibition of photosynthesis by water stress could have this error. However, if no stomatal closure occurs, any reduction in photosynthesis must be due to nonstomatal inhibition of photosynthesis. Net carbon dioxide exchange rates and conductances to water vapor were measured under field conditions in upper canopy leaves of tomato plants during two summers in Beltsville, Maryland, USA. Comparisons were made near midday at high irradiance between leaflets in air with the ambient water vapor content and in air with a higher water content. The higher water content, which lowered the leaf to air water vapor pressure difference (VPD), was imposed either one half hour or several hours before measurements of gas exchange. In both seasons, and irrespective of the timing of the imposition of different VPDs, net photosynthesis increased 60% after decreasing the VPD from 3 to 1 kPa. There were no differences in leaf conductance between leaves at different VPDs, thus transpiration rates were threefold higher at 3 than at 1 kPa VPD. It is concluded that nonstomatal inhibition of photosynthesis did occur in these leaves at high transpiration rate.     

Photosynthesis, Transpiration, Leaf Temperature, and Stomatal Activity of Cotton Plants under Varying Water Potentials

Cotton plants, Gossypium hirsutum L. were grown in a growth room under incident radiation levels of 65, 35, and 17 Langleys per hour to determine the effects of vapor pressure deficits (VPD's) of 2, 9, and 17 mm Hg at high soil water potential, and the effects of decreasing soil water potential and reirrigation on transpiration, leaf temperature, stomatal activity, photosynthesis, and respiration at a VPD of 9 mm Hg.

Transpiration was positively correlated with radiation level, air VPD and soil water potential. Reirrigation following stress led to slow recovery, which may be related to root damage occurring during stress. Leaf water potential decreased with, but not as fast as, soil water potential.

Leaf temperature was usually positively correlated with light intensity and negatively correlated with transpiration, air VPD, and soil water. At high soil water, leaf temperatures ranged from a fraction of 1 to a few degrees above ambient, except at medium and low light and a VPD of 19 mm Hg when they were slightly below ambient, probably because of increased transpirational cooling. During low soil water leaf temperatures as high as 3.4° above ambient were recorded. Reirrigation reduced leaf temperature before appreciably increasing transpiration. The upper leaf surface tended to be warmer than the lower at the beginning of the day and when soil water was adequate; otherwise there was little difference or the lower surface was warmer. This pattern seemed to reflect transpiration cooling and leaf position effects.

Although stomata were more numerous in the lower than the upper epidermis, most of the time a greater percentage of the upper were open. With sufficient soil water present, stomata opened with light and closed with darkness. Fewer stomata opened under low than high light intensity and under even moderate, as compared with high soil water. It required several days following reirrigation for stomata to regain original activity levels.

Apparent photosynthesis of cotton leaves occasionally oscillated with variable amplitude and frequency. When soil water was adequate, photosynthesis was nearly proportional to light intensity, with some indication of higher rates at higher VPD's. As soil water decreased, photosynthesis first increased and then markedly decreased. Following reirrigation, photosynthesis rapidly recovered.

Respiration was slowed moderately by decreasing soil water but increased before watering. Respiration slowed with increasing leaf age only on leaves that were previously under high light intensity.

     

Soil water deficits decrease the internal conductance to CO2 transfer but atmospheric water deficits do not

The internal conductance to CO₂ supply from substomatal cavitiesto sites of carboxylation poses a large limitation to photosynthesis.It is known that internal conductance is decreased by soil waterdeficits, but it is not known if it is affected by atmosphericwater deficits (i.e. leaf to air vapour pressure deficit, VPD).The aim of this paper was to examine the responses of internalconductance to atmospheric and soil water deficits in seedlingsof the evergreen perennial Eucalyptus regnans F. Muell and theherbaceous plants Solanum lycopersicum (formerly Lycopersiconesculentum) Mill. and Phaseolus vulgaris L. Internal conductancewas estimated with the variable J method from concurrent measurementsof gas exchange and fluorescence. In all three species steady-statestomatal conductance decreased by 30% as VPD increased from1 kPa to 2 kPa. In no species was internal conductance affectedby VPD despite large effects on stomatal conductance. In contrast,soil water deficits decreased stomatal conductance and internalconductance of all three species. Decreases in stomatal andinternal conductance under water deficit were proportional,but this proportionality differed among species, and thus therelationship between stomatal and internal conductance differedamong species. These findings indicate that soil water deficitsaffect internal conductance while atmospheric water deficitsdo not. The reasons for this distinction are unknown but areconsistent with soil and atmospheric water deficits having differingeffects on leaf physiology and/or root–shoot communication. Key words: Carbon dioxide, drought, internal conductance, mesophyll conductance, photosynthesis, stomatal conductance, transfer conductance, vapour pressure deficit, water deficit Received 11 October 2007; Revised 9 November 2007 Accepted 15 November 2007     

The Effect of Air Humidity on Photosynthesis of Ryegrass and White Clover at Three Temperatures

Increasing leaf-air vapour pressure deficit (VPD) decreasedthe stomatal conductance and the photosynthetic rate of leavesof ryegrass (Lolium perenne L.) and white clover (Trifolhimrepens L.) at light saturation and at lower irradiance. In ryegrassboth conductance and photosynthesis, and in clover photosynthesis,decreased less with increasing VPD in low irradiance than theydid at an irradiance which saturated photosynthesis. In ryegrass,relative to their values at 10 mb, photosynthesis and conductancedecreased less with increasing VPD at 25 °C than at 20 or16·5 °C. In white clover, relative conductance (butnot photosynthesis) was less reduced at 25 than at 16·5°C Measurements of VPD of air in the leaf canopy of a field-growncrop are combined with the observed responses of photosynthesisto VPD and temperature in a model. This shows that high VPDis likely to depress photosynthesis significantly and that,during a typical day, the rate of light saturated photosynthesismay remain fairly steady, because the depression of photosynthesisdue to rising VPD is offset by the stimulation due to risingtemperature Perennial ryegrass, Lolium perenne L., White clover, Trifolhim repens L., photosynthesis, leaf conductance, water vapour pressure deficit, temperature     

冬小麦叶片气孔导度模型水分响应函数的参数化

植物气孔导度模型的水分响应函数用来模拟水分胁迫对气孔导度的影响过程, 是模拟缺水环境下植物与大气间水、碳交换过程的关键算法。水分响应函数包括空气湿度响应函数和土壤湿度(或植物水势)响应函数, 该研究基于田间实验观测, 分析了冬小麦(Triticum aestivum)叶片气孔导度对不同空气饱和差和不同土壤体积含水量或叶水势的响应规律。一个土壤水分梯度的田间处理在中国科学院禹城综合试验站实施, 不同水分胁迫下的冬小麦叶片气体交换过程和气孔导度以及其他的温湿度数据被观测, 同时观测了土壤含水量和叶水势。实验数据表明, 冬小麦叶片气孔导度对空气饱和差的响应呈现双曲线规律, 变化趋势显示大约1 kPa空气饱和差是一个有用的阈值, 在小于1 kPa时, 冬小麦气孔导度对空气饱和差变化反应敏感, 而大于1 kPa后则反应缓慢; 分析土壤体积含水量与中午叶片气孔导度的关系发现, 中午叶片气孔导度随土壤含水量增加大致呈现线性增加趋势, 但在平均土壤体积含水量大于大约25%以后, 气孔导度不再明显增加, 而是维持在较高导度值上下波动; 冬小麦中午叶片水势与相应的气孔导度之间, 随着叶水势的增加, 气孔导度呈现增加趋势。根据冬小麦气孔导度对空气湿度、土壤湿度和叶水势的响应规律, 研究分别采用双曲线和幂指数形式拟合了水汽响应函数, 用三段线性方程拟合了土壤湿度响应函数和植物水势响应函数, 得到的参数可以为模型模拟冬小麦的各类水、热、碳交换过程采用。     

The effect of vapor pressure on stomatal control of gas exchange in Douglas fir (Pseudotsuga menziesii) saplings

Summary Increasing leaf to air vapor pressure deficit (VPD) caused reductions in stomatal conductance of both current year and previous season needles of Pseudotsuga menziesii saplings. The stomata of current year needles were found to be more responsive to changes in VPD than those of previous season needles. The reductions in stomatal conductance of current year needles were not associated with decreases in xylem pressure potential. In fact, the reductions in stomatal conductance of current year needles were sometimes sufficient to reduce transpiration and thus raise xylem pressure potential even though VPD was increasing. There was a decline in stomatal responsiveness to VPD in current year needles between early and late summer. Pressure-volume curves determined for different age needles at different times of the year suggested that differences and changes in stomatal responsiveness to VPD may have been caused in part by differences and changes in needle water potential components. Hexane washes of current year needles during the late summer succeeded in partially restoring their VPD sensitivity, suggesting that changes in the water permeability of the external cuticle during needle maturation may also have played a role in causing the summer decline in VPD responsiveness.In both current and previous year needles VPD-induced changes in stomatal conductance had a greater relative effect on transpiration (q _w) than on net photosynthesis (Ph_N). In maturing needles the ratio of the sensitivities of transpiration and net photosynthesis to changes in stomatal conductance, (q _w/g _s)/Ph_N/g _s), remained nearly constant as VPD was varied. This provides experimental support for a recent hypothesis that stomata respond to environmental fluctuations in such a manner as to maintain the above ratio constant, which optimizes CO₂ uptake with respect to water loss.     

Selective oviposition by a leaf miner in response to temporal variation in abscission

Summary Responses to humidity of net photosynthesis and leaf conductance of single attached leaves were examined in populations of herbs from wet soil sites in Beltsville, Maryland and Davis, California, USA. Plants were grown in controlled environments under three conditions which differed in the magnitude of the day-night temperature difference and in daytime air saturation deficit. No population differences in response were found in Abutilon theophrasti. In Amaranthus hybridus stomatal conductance and net photosynthesis were more reduced by increasing leaf to air water vapor pressure difference (VPD) in the population from Beltsville, but only for the growth condition with a constant 25°C temperature. In Chenopodium album, stomatal conductance was more sensitive to VPD in the population from Davis, but only for the growth condition with 28/22°C day/night temperatures. Population differences in the sensitivity to VPD of leaf conductance were associated with differences in leaf area to root weight ratio. The relative reduction of net photosynthesis as VPD increased was greater than, equal to, or less than the relative decrease in substomatal carbon dioxide partial pressure. The pattern depended on species, and on growth condition. From these results one can not conclude that environmental humidity has been a strong selective force in determining sensitivity to humidity of stomatal conductance.     

Stomatal response to humidity in sugarcane and soybean: effect of vapour pressure difference on the kinetics of the blue light response

Abstract. The effect of atmospheric humidity on the kinetics of stomatal responses was quantified in gas exchange experiments using sugarcane ( Saccharum spp. hybrid) and soybean ( Glycine max ). Pulses of blue light were used to elicit pulses of stomatal conductance that were mediated by the specific blue light response of guard cells. Kinetic parameters of the conductance response were more closely related to leaf-air vapour pressure difference (VPD) than to relative humidity or transpiration. Increasing VPD significantly accelerated stomatal opening in both sugarcane and soybean, despite an approximately five-fold faster response in sugarcane. In contrast, the kinetics of stomatal recovery (closure) following the pulse were similar in the two species. Acceleration of opening by high VPD was observed even under conditions where soybean exhibited a feedforward response of decreasing transpiration (E) with increasing evaporative demand (VPD). This result suggests that epidermal, rather than bulk leaf, water status mediates the VPD effect on stomatal kinetics. The data are consistent with the hypothesis that increased cpidermal water loss at high VPD decreases the backpressure exerted by neighbouring cells on guard cells. allowing more rapid stomatal opening per unit of guard cell metabolic response to blue light.     

The responses of stomata and leaf gas exchange to vapour pressure deficits and soil water content

Summary The responses of photosynthesis, transpiration and leaf conductance to changes in vapour pressure deficit were followed in well-watered plants of the herbaceous species, Helianthus annuus, Helianthus nuttallii, Pisum sativum and Vigna unguiculata, and in the woody species having either sclerophyllous leaves, Arbutus unedo, Nerium oleander and Pistacia vera, or mesomorphic leaves, Corylus avellana, Gossypium hirsutum and Prunus dulcis. When the vapour pressure deficit of the air around a single leaf in a cuvette was varied from 10 to 30 Pa kPa^-1 in 5 Pa kPa^-1 steps, while holding the remainder of the plant at a vapour presure deficit of 10 Pa kPa^-1, the leaf conductance and net photosynthetic rate of the leaf decreased in all species. The rate of transpiration increased initially with increase in vapour pressure deficit in all species, but in several species a maximum transpiration rate was observed at 20 to 25 Pa kPa^-1. Concurrent measurements of the leaf water potential by in situ psychrometry showed that an increase in the vapour pressure deficit decreased the leaf water potential in all species. The decrease was greatest in woody species, and least in herbaceous species. When the vapour pressure deficit around the remainder of the plant was increased while the leaf in the cuvette was exposed to a low and constant vapour pressure deficit, similar responses in both degree and magnitude in the rates of transpiration and leaf conductance were observed in the remainder of the plant as those occurring when the vapour pressure deficit around the single leaf was varied. Increasing the external vapour pressure deficit lowered the water potential of the leaf in the cuvette in the woody species and induced a decrease in leaf conductance in some, but not all, speies. The decrease in leaf conductance with decreasing water potential was greater in the woody species when the vapour pressure deficit was increased than when it remained low and constant, indicating that changing the leaf-to-air vapour pressure difference had a direct effect on the stomata in these species. The low hydraulic resistance and maintenance of a high leaf water potential precluded such an analysis in the herbaceous species. We conclude that at least in the woody species studied, an increase in the vapour pressure deficit around a leaf will decrease leaf gas exchange through a direct effect on the leaf epidermis and sometimes additionally through a lowering of the mesophyll water potential.     

The Effect of Drought and Vapour Pressure Deficit on Gas Exchange of Young Kiwifruit (Actinidia deliciosavar.deliciosa) Vines

To evaluate the effect of drought and vapour pressure deficit (VPD) on stomatal behaviour and gas exchange parameters, young kiwifruit vines (Actinidia deliciosavar.deliciosacv. Hayward) were exposed to alternating periods of drought and drought-relief over two growing seasons. Vines were grown either in the field or in containers. Stomatal conductance of fully-expanded leaves rapidly decreased as pre-dawn leaf water potential was reduced below a threshold value of -0.3MPa. Stomatal conductance reached minimum values of 10–20mmol m^-2s^-1. Transpiration rate was similarly sensitive to changes in leaf water status, whereas more severe drought levels were necessary to affect photosynthesis significantly. Net daily carbon gains were estimated at 4.7 and 2.7gm^-2for irrigated and droughted vines, respectively. Gas exchange parameters recovered to values of irrigated vines within a few hours after relief of stress. Rate of recovery depended on the level of stress reached during the previous drought period. There was a steady decline in stomatal conductance when VPD was increased from 0.8 to 2.5kPa in both irrigated and droughted vines. The VPD at which stomatal conductance reached 50% of maximum values was 2.1–2.2kPa for both treatments. We conclude that stomata were highly sensitive to changes in soil water status and that midday depression of photosynthesis measured in kiwifruit vines was related to water deficits arising in the leaf because of both transpirational losses and to the direct effect of increasing VPD.     

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.     

Stomatal sensitivity to carbon dioxide and humidity: a comparison of two c(3) and two c(4) grass species

The sensitivity of stomatal conductance to changes of CO₂ concentration and leaf-air vapor pressure difference (VPD) was compared between two C₃ and two C₄ grass species. There was no evidence that stomata of the C₄ species were more sensitive to CO₂ than stomata of the C₃ species. The sensitivity of stomatal conductance to CO₂ change was linearly proportional to the magnitude of stomatal conductance, as determined by the VPD, the same slope fitting the data for all four species. Similarly, the sensitivity of stomatal conductance to VPD was linearly proportional to the magnitude of stomatal conductance. At small VPD, the ratio of intercellular to ambient CO₂ concentration, C_i/C_a, was similar in all species (0.8-0.9) but declined with increasing VPD, so that, at large VPD, C_i/C_a was 0.7 and 0.5 (approximately) in C₃ and C₄ species, respectively. Transpiration efficiency (net CO₂ assimilation rate/transpiration rate) was larger in the C₄ species than in the C₃ species at current atmospheric CO₂ concentrations, but the relative increase due to high CO₂ was larger in the C₃ than in the C₄ species.