Water Potential Components, Stomatal Function, and Liquid Phase Water Transport Resistances of Four Arctic and Alpine Species in Relation to Moisture Stress

Transpiration measurements of two alpine tundra species, Deschampsia caespitosa and Geum rossii, and two arctic tundra species, Dupontia fischeri and Carex aquatilis, were conducted under varying atmospheric and soil moisture stress regimes to determine if the stomatal response to water stress may play a role in the local distributions of these species. Under low soil moisture stress, stomata of the species restricted typically to wet meadow areas, Deschampsia and Dupontia, did not exhibit closure until leaf water potential declined. However, when soil moisture stress was low and atmospheric stress increased, Geum and particularly Carex exhibited partial stomatal closure before leaf water potential dropped, suggesting a direct response of the stomata to the vapor pressure gradient between the leaf and the atmosphere. Lower liquid phase water transport resistance from the soil to the leaves may also reduce the development of leaf moisture stress in Geum. Furthermore, Geum and possibly Carex appeared to undergo less of a loss of leaf turgor when leaf water potential decreased. This response may serve to maintain leaf cell turgor and to abate the reduction in leaf enlargement.     

Water use strategy affects avoidance of ozone stress by stomatal closure in Mediterranean trees—A modelling analysis

Both ozone (O₃) and drought can limit carbon fixation by forest trees. To cope with drought stress, plants have isohydric or anisohydric water use strategies. Ozone enters plant tissues through stomata. Therefore, stomatal closure can be interpreted as avoidance to O₃ stress. Here, we applied an optimization model of stomata involving water, CO₂, and O₃ flux to test whether isohydric and anisohydric strategies may affect avoidance of O₃ stress by stomatal closure in four Mediterranean tree species during drought. The data suggest that stomatal closure represents a response to avoid damage to the photosynthetic mechanisms under elevated O₃ depending on plant water use strategy. Under high-O₃ and well-watered conditions, isohydric species limited O₃ fluxes by stomatal closure, whereas anisohydric species activated a tolerance response and did not actively close stomata. Under both O₃ and drought stress, however, anisohydric species enhanced the capacity of avoidance by closing stomata to cope with the severe oxidative stress. In the late growing season, regardless of the water use strategy, the efficiency of O₃ stress avoidance decreased with leaf ageing. As a result, carbon assimilation rate was decreased by O₃ while stomata did not close enough to limit transpirational water losses.     

Stomatal action directly feeds back on leaf turgor: new insights into the regulation of the plant water status from non‐invasive pressure probe measurements

Uptake of CO₂ by the leaf is associated with loss of water. Control of stomatal aperture by volume changes of guard cell pairs optimizes the efficiency of water use. Under water stress, the protein kinase OPEN STOMATA 1 (OST1) activates the guard‐cell anion release channel SLOW ANION CHANNEL‐ASSOCIATED 1 (SLAC1), and thereby triggers stomatal closure. Plants with mutated OST1 and SLAC1 are defective in guard‐cell turgor regulation. To study the effect of stomatal movement on leaf turgor using intact leaves of Arabidopsis, we used a new pressure probe to monitor transpiration and turgor pressure simultaneously and non‐invasively. This probe permits routine easy access to parameters related to water status and stomatal conductance under physiological conditions using the model plant Arabidopsis thaliana. Long‐term leaf turgor pressure recordings over several weeks showed a drop in turgor during the day and recovery at night. Thus pressure changes directly correlated with the degree of plant transpiration. Leaf turgor of wild‐type plants responded to CO₂, light, humidity, ozone and abscisic acid (ABA) in a guard cell‐specific manner. Pressure probe measurements of mutants lacking OST1 and SLAC1 function indicated impairment in stomatal responses to light and humidity. In contrast to wild‐type plants, leaves from well‐watered ost1 plants exposed to a dry atmosphere wilted after light‐induced stomatal opening. Experiments with open stomata mutants indicated that the hydraulic conductance of leaf stomata is higher than that of the root–shoot continuum. Thus leaf turgor appears to rely to a large extent on the anion channel activity of autonomously regulated stomatal guard cells.     

Responses of stomata to environmental factors-experiments with isolated epidermal strips of Polypodium vulgare

Summary Stomatal apertures of isolated and suitably conditioned epidermal strips of Polypodium vulgare are described as the stomata respond to the influences of temperature, air humidity, and water potential at the epidermal inner walls. Water stress as a result of reduced water potential in the substomatal airspace leads to narrower stomatal pores when water potential falls below -8 bar. Water potentials above this threshold value show minor influence. Stomatal responses to such water stress strongly interact with the responses to humidity changes in ambient air and to temperature. The linear dependence of stomatal apertures on the vapor saturation deficit of the air (closing) is shifted to lower values (more closed) by lower leaf bulk water potentials.Stomatal behavior depending on the temperature factor seems to be reversed by higher water stress. Without water stress, rising temperatures between 20 and 28° C are accompanied by further opening of the pores, whereas an increase of temperature within this range leads to narrowing of the stomata under the influence of lower water potentials within the substomatal airspace. It can be demonstrated that stomatal aperture values of Polypodium vulgare depending on temperature always describe optimum curves. With no water stress, closing does not occur before rather high temperatures are reached and above a broad range of maximal opening. Water stress, on the other hand, results in more pronounced narrowing of stomatal pores and shifts the onset to considerably lower temperatures.     

Stomata of the CAM plant Tillandsia recurvata respond directly to humidity

Summary Under controlled conditions, CO₂ exchange of Tillandsia recurvata showed all characteristics of CAM. During the phase of nocturnal CO₂ fixation stomata of the plant responded sensitively to changes in ambient air humidity. Dry air resulted in an increase, moist air in a decrease of diffusion resistance. The evaporative demand of the air affected the level of stomatal resistance during the entire night period. Due to stomatal closure, the total nocturnal water loss of T. recurvata was less at low than at high humidity. It is concluded that stomata respond directly to humidity and not via bulk tissue water conditions of the leaves. Such control of transpiration may optimize water use efficiency for this almost rootless, extreme epiphyte.     

Stomatal response to vapour pressure deficit and the effect of plant water stress

Abstract The dynamic response of stomata to changes in atmospheric humidity was investigated in Fragaria × ananassa Duch., Picea engelmannii Parry, and Pseudotsuga menziesii (Mirb.) Franco; and the effect of water stress on this response was determined in Pseudotsuga menziesii. The plants were rotated through three regimes of ambient temperature and vapour pressure deficit: 35°C–3. 5kPa, 35°C–0. 5 kPa, and 20°C–1. 5kPa. Branch and leaflet conductance were measured with a steady-state porometer, first at ambient vapour pressure deficit and then at one of four treatment conditions achieved by increasing or decreasing vapour pressure within the porometer cuvette. All three species showed similar stomatal response: enhanced conductance at low vapour pressure deficit and depressed conductance at high vapour pressure deficit. Engelmann spruce was more sensitive than Douglas fir and strawberry. Plant water status significantly altered stomatal response to vapour pressure deficit. The relationship of conductance of xylem water potential was linear under ambient conditions but became curvilinear when conductance was measured above and below ambient vapour pressure deficit. Between ?0. 5 MPa and ?2. 0 MPa xylem water potential, the stomata were sensitive to vapour pressure deficit, but below ? 2. 0 MPa, the sensitivity decreased.     

Stomatal sensitivities to changes in leaf water potential, air humidity, CO2 concentration and light intensity, and the effect of abscisic acid on the sensitivities in six temperate deciduous tree species

To characterise the stomata of six temperate deciduous tree species, sets of stomatal sensitivities to all the most important environmental factors were measured. To compare the importance of abscisic acid (ABA) in the different stomatal responses, the effect of exogenous ABA on all the stomatal sensitivities was determined.Almost all the stomatal sensitivities: the sensitivity to a decrease in leaf water potential, air humidity, CO₂ concentration ([CO₂]) and light intensity, and to an increase in [CO₂] and light intensity were the highest in the slow-growing species, and the lowest in the fast-growing species. Drought increased the sensitivity to the environmental changes that induce a decrease in the stomatal conductance, and decreased the sensitivity to the changes that induce an increase in this conductance. The sensitivities of the slow-growers were most strongly affected by drought and ABA. Therefore the success of the slow-growers in their ecological niches can be based on the highly sensitive and strictly regulated responses of their stomata. The fast-growers had the highest sensitivity to an increase in leaf water potential and this sensitivity was sharply reduced by drought and ABA. Thus, the dominance of the trees in riparian areas can be based on the ability of their stomata to quickly reach high conductance in well-watered conditions and to efficiently decrease this rate during drought.Stomatal sensitivities to the hydraulic environmental factors (water potentials in plant and air) had higher values in well-watered trees and a more pronounced response to drought than the sensitivities to the photosynthetic environmental factors ([CO₂] and light intensity). Thus, the hydraulic factors most likely prevail over the photosynthetic factors in determining stomatal conductance in these species.In response to exogenous ABA, the rates of stomatal closure, following a decrease in air humidity and light intensity, and an increase in [CO₂], were accelerated. Stomatal opening following an increase in air humidity and light intensity and a decrease in [CO₂] was replaced by slow closing. The rate of stomatal opening following an increase in leaf water potential was reduced. As the sensitivities to changes in light were modified less by the ABA than the other stomatal sensitivities, the prediction of stomatal responses on the basis of the sensitivity to light alone should be excluded in stomatal models.     

Stomata response to changes in temperature and humidity in wheat cultivars grown under contrasting climatic conditions

Stomatal response to changes in temperature and humidity was studied in wheat (Triticum aestivum L.) cv. Iren’ cultivated under conditions of high water supply and cv. Kazakhstanskaya 10, which is relatively drought tolerant. Experiments were performed under both laboratory and field conditions. It was demonstrated that stomata of cv. Kazakhstanskaya 10 plants closed rapidly with reducing humidity (the response of the first type), whereas, in cv. Iren’, this response was less expressed and, under conditions of a high water content in soil, stomatal conductance could increase in response to reduced humidity (the response of the second type). At an increased stomatal conductance and transpiration, water content in cv. Iren’ plants was maintained due to the increase in hydraulic conductance and water inflow from the roots. A possible role of the first-type response (rapid stomata closure) for growth maintenance under drought and of the second-type response (a parallel increase in the stomatal and hydraulic conductance) for providing of rapid growth and high productivity under sufficient water supply is discussed. A possibility to use the type of stomata behavior for cultivar assessment is considered.     

The trafficking protein SYP121 of Arabidopsis connects programmed stomatal closure and K⁺ channel activity with vegetative growth

The vesicle‐trafficking protein SYP121 (SYR1/PEN1) was originally identified in association with ion channel control at the plasma membrane of stomatal guard cells, although stomata of the Arabidopsis syp121 loss‐of‐function mutant close normally in ABA and high Ca²⁺. We have now uncovered a set of stomatal phenotypes in the syp121 mutant that reduce CO₂ assimilation, slow vegetative growth and increase water use efficiency in the whole plant, conditional upon high light intensities and low relative humidity. Stomatal opening and the rise in stomatal transpiration of the mutant was delayed in the light and following Ca²⁺‐evoked closure, consistent with a constitutive form of so‐called programmed stomatal closure. Delayed reopening was observed in the syp121, but not in the syp122 mutant lacking the homologous gene product; the delay was rescued by complementation with wild‐type SYP121 and was phenocopied in wild‐type plants in the presence of the vesicle‐trafficking inhibitor Brefeldin A. K⁺ channel current that normally mediates K⁺ uptake for stomatal opening was suppressed in the syp121 mutant and, following closure, its recovery was slowed compared to guard cells of wild‐type plants. Evoked stomatal closure was accompanied by internalisation of GFP‐tagged KAT1 K⁺ channels in both wild‐type and syp121 mutant guard cells, but their subsequently recycling was slowed in the mutant. Our findings indicate that SYP121 facilitates stomatal reopening and they suggest that K⁺ channel traffic and recycling to the plasma membrane underpins the stress memory phenomenon of programmed closure in stomata. Additionally, they underline the significance of vesicle traffic for whole‐plant water use and biomass production, tying SYP121 function to guard cell membrane transport and stomatal control.     

Environmental regulation of stomatal response in the <Emphasis Type="Italic">Arabidopsis</Emphasis> Cvi-0 ecotype

The Arabidopsis Cape Verde Islands (Cvi-0) ecotype is known to differ from other ecotypes with respect to environmental stress responses. We analyzed the stomatal behavior of Cvi-0 plants, in response to environmental signals. We investigated the responses of stomatal conductance and aperture to high [CO₂] in the Cvi-0 and Col-0 ecotypes. Cvi-0 showed constitutively higher stomatal conductance and more stomatal opening than Col-0. Cvi-0 stomata opened in response to light, but the response was slow. Under low humidity, stomatal opening was increased in Cvi-0 compared to Col-0. We then assessed whether low humidity affects endogenous ABA levels in Cvi-0. In response to low humidity, Cvi-0 had much higher ABA levels than Col-0. However, epidermal peels experiments showed that Cvi-0 stomata were insensitive to ABA. Measurements of organic and inorganic ions in Cvi-0 guard cell protoplasts indicated an over-accumulation of osmoregulatory anions (malate and Cl⁻). This irregular anion homeostasis in the guard cells may explain the constitutive stomatal opening phenotypes of the Cvi-0 ecotype, which lacks high [CO₂]-induced and low humidity-induced stomatal closure.     

Ecophysiological studies of Sonoran Desert plants

Summary The gas exchange and water relations of two Sonoran Desert plants are compared during contrasting periods of water and heat stress. Photosynthesis of Acacia greggii, a winter deciduous shrub, and Cercidium microphyllum, a chlorophyllous stemmed tree, show a moderate correlation with dawn plant water potential. For both species a relationship between stomatal conductance and dawn plant water potential was not apparent, although A. greggii demonstrated a greater overall stomatal conductance. This affected a greater daytime decrease in plant water potential at all levels of water stress and suggests A. greggii is less sensitive to water stress. Our results suggest the lower limit for gross photosynthesis occurs when dawn plant water potentials are less than -44 and -31 bars for the shrub and tree species, respectively. During periods of extreme water and heat stress the photosynthetic capacity of both species is regulated more by mesophyll than stomatal conductance. However, partial stomatal closure causes plant water potential to increase during the day and exceed dawn values. During periods of minimal water and heat stress the daily course of photosynthesis parallels the change in stomatal conductance and irradiance. Maximum gross photosynthesis rates are nearly three-fold higher than the rates observed during periods of stress, with those of A. greggii generally greater than the rates observed in plants of C. microphyllum.     

Stomatal behaviour and leaf water potential of chilled and water-stressed Solanum melongena, as influenced by growth history

Abstract Leaf diffusion resistance and leaf water potential of intact Solanum melongena plants were measured during a period of chilling at 6 °C. Two pretreatments, consisting of a period of water stress or a foliar spraying of abscisic acid (ABA), were imposed upon the plants prior to chilling. The control plants did not receive a pretreatment. In addition to intact plant studies, stomatal responses to water loss and exogenous abscisic acid were investigated using excised leaves, and the influence of the pretreatment observed. Chilled, control plants wilted slowly and maintained open stomata despite a decline in leaf water potential to –2.2 MPa after 2 d of chilling. In contrast plants that had been water stressed or had been sprayed with abscisic acid, prior to chilling, did not wilt and maintained a higher leaf water potential and a greater leaf diffusion resistance. In plants that had not received a pretreatment, abscisic acid caused stomatal closure at 35 °C, but at 6°C it did not influence stomatal aperture. The two pretreatments greatly increased stomatal sensitivity to both exogenous ABA and water stress, at both temperatures. Stomatal response to water loss from excised leaves was greatly reduced at 6°C. These results are discussed in relation to low temperature effects on stomata and the influence of preconditioning upon plant water relations.     

Decoupling between stomatal conductance and photosynthesis occurs under extreme heat in broadleaf tree species regardless of water access

High air temperatures increase atmospheric vapor pressure deficit (VPD) and the severity of drought, threatening forests worldwide. Plants regulate stomata to maximize carbon gain and minimize water loss, resulting in a close coupling between net photosynthesis (A_net) and stomatal conductance (g_s). However, evidence for decoupling of g_s from A_net under extreme heat has been found. Such a response both enhances survival of leaves during heat events but also quickly depletes available water. To understand the prevalence and significance of this decoupling, we measured leaf gas exchange in 26 tree and shrub species growing in the glasshouse or at an urban site in Sydney, Australia on hot days (maximum T_air > 40°C). We hypothesized that on hot days plants with ample water access would exhibit reduced A_net and use transpirational cooling leading to stomatal decoupling, whereas plants with limited water access would rely on other mechanisms to avoid lethal temperatures. Instead, evidence for stomatal decoupling was found regardless of plant water access. Transpiration of well-watered plants was 23% higher than model predictions during heatwaves, which effectively cooled leaves below air temperature. For hotter, droughted plants, the increase in transpiration during heatwaves was even more pronounced—g_s was 77% higher than model predictions. Stomatal decoupling was found for most broadleaf evergreen and broadleaf deciduous species at the urban site, including some wilted trees with limited water access. Decoupling may simply be a passive consequence of the physical effects of high temperature on plant leaves through increased cuticular conductance of water vapor, or stomatal decoupling may be an adaptive response that is actively regulated by stomatal opening under high temperatures. This temperature response is not yet included in any land surface model, suggesting that model predictions of evapotranspiration may be underpredicted at high temperature and high VPD.     

Hydraulic control of stomatal conductance in Douglas fir [Pseudotsuga menziesii (Mirb.) Franco] and alder [Alnus rubra (Bong)] seedlings

Experiments were conducted on 1-year-old Douglas fir [Pseudotsuga menziesii (Mirb.) Franco] and 2- to 3-month-old alder [Alnus rubra (Bong)] seedlings growing in drying soils to determine the relative influence of root and leaf water status on stomatal conductance (g_c). The water status of shoots was manipulated independently of that of the roots using a pressure chamber that enclosed the root system. Pressurizing the chamber increases the turgor of cells in the shoot but not in the roots. Seedling shoots were enclosed in a whole-plant cuvette and transpiration and net photosynthesis rates measured continuously. In both species, stomatal closure in response to soil drying was progressively reversed with increasing pressurization. Responses occurred within minutes of pressurization and measurements almost immediately returned to pre-pressurization levels when the pressure was released. Even in wet soils there was a significant increase in g_c with pressurization. In Douglas fir, the stomatal response to pressurization was the same for seedlings grown in dry soils for up to 120 d as for those subjected to drought stress over 40 to 60 d. The stomatal conductance of both Douglas fir and alder seedlings was less sensitive to root chamber pressure at higher vapour pressure deficits (D), and stomatal closure in response to increasing D from 1.04 to 2.06 kPa was only partially reversed by pressurization. Our results are in contrast to those of other studies on herbaceous species, even though we followed the same experimental approach. They suggest that it is not always appropriate to invoke a ‘feedforward’ model of short-term stomatal response to soil drying, whereby chemical messengers from the roots bring about stomatal closure.     

Aquaporin regulation in roots controls plant hydraulic conductance,stomatal conductance,and leaf water potential in Pinus radiata under water stress

Stomatal regulation is crucial for forest species performance and survival on drought‐prone sites. We investigated the regulation of root and shoot hydraulics in three Pinus radiata clones exposed to drought stress and its coordination with stomatal conductance (g_s) and leaf water potential (Ψ_leaf). All clones experienced a substantial decrease in root‐specific root hydraulic conductance (K_root‐r) in response to the water stress, but leaf‐specific shoot hydraulic conductance (K_shoot‐l) did not change in any of the clones. The reduction in K_root‐r caused a decrease in leaf‐specific whole‐plant hydraulic conductance (K_plant‐l). Among clones, the larger the decrease in K_plant‐l, the more stomata closed in response to drought. Rewatering resulted in a quick recovery of K_root‐r and g_s. Our results demonstrated that the reduction in K_plant‐l, attributed to a down regulation of aquaporin activity in roots, was linked to the isohydric stomatal behaviour, resulting in a nearly constant Ψ_leaf as water stress started. We concluded that higher K_plant‐l is associated with water stress resistance by sustaining a less negative Ψ_leaf and delaying stomatal closure.     

Short-term and long-term effects of plant water deficits on stomatal response to humidity in Corylus avellana L.

Short-term (hours) changes in plant water status were induced in hazel (Corylus avellana L.) by changing the evaporative demand on a major portion of the shoot while maintaining a branch in a constant environment. Stomatal conductance of leaves on the branch was influenced little by these short-term changes in water status even with changes in leaf water potential as great as 8 bars. Long-term (days) changes in plant water status were imposed by soil drying cycles. Stomatal conductance progessively decreased with increases in long-term water stress. Stomata still responded to humidity with long-term water stress but the range of the conductance response decreased. Threshold responses of stomata to leaf water potential were not observed with either short-term or long-term changes in plant water status even when leaves wilted. It is suggested that concurrent measurements of plant water status may not be sufficient for explaining stomatal and other plant responses to drought.     

低空气湿度下气孔运动的调控

植物气孔应答气候环境因子的机制,一直是植物抗逆生理研究领域的热点课题。空气湿度是主要的气候环境因子之一,人们很早就发现降低空气湿度能导致气孔开度变小,但到目前为止,对调控这一生物学过程的机制还知之甚少。本文概述近年来在植物气孔应答低空气湿度信号的运动规律、水被动反馈假说、水主动反馈假说、水力调节与代谢调节的关系等方面的进展,并基于进化学的理论,提出了水力调节和代谢调节的反向消长模式,以期解释大量物种气孔对低空气湿度的不同应答反应。     

Stomatal responses to changes in humidity in plants growing in the desert

Summary The stomata of plants growing in the Negev Desert, namely the stomata of the mesomorphic leaves of Prunus armeniaca, the xeromorphic stems of Hammada scoparia, and the succulent leaves of Zygophyllum dumosum, respond to changes in air humidity. Under dry air conditions diffusion resistance increases. Under moist air conditions diffusion resistance decreases. When the stomata close at low air humidity the water content of the apricot leaves increases. The stomata open at high air humidity in spite of a decrease in leaf water content. This excludes a reaction via the water potential in the leaf tissue and proves that the stomatal aperture has a direct response to the evaporative conditions in the atmosphere. In all species the response to air humidity is maintained over a period of many hours also when the soil is considerably dry. The response is higher in plants with poor water supply then in well watered plants. Thus for field conditions and for morphologically different types of photosynthesizing organs the results confirm former experiments carried out with isolated epidermal strips.     

Ecophysiological studies of Sonoran Desert plants

Summary The gas exchange of two Sonoran Desert plants was measured near optimum soil water conditions occurring in the summer and winter. Photosynthesis and stomatal conductance rates of a drought-deciduous shrub, Ambrosia deltoidea, and an evergreen non-riparian tree, Olneya tesota, are mainly affected by plant water potential. During such periods the diurnal gas exchange patterns are characterized by maximum rates of photosynthesis and stomatal conductance occurring early in the day, which decrease progressively thereafter. The effect of plant water potential on gas exchange is both direct and indirect. Decreasing plant water potential indirectly affects ¹⁴CO₂ assimilation by closure of stomata, and the effect is similar in both species. However, the direct effects of decreasing plant water potential are dissimilar in the two species. Plants of the shrub species have a higher potential maximum photosynthesis, but are more sensitive to plant water stress than are plants of the tree species. Both species respond to favorable growth conditions in the summer and winter, and have the potential for rapid carbon input into the Sonoran Desert ecosystem.This research was supported by National Science Foundation Grant BMS 74-02671-A04, through the U.S./I.B.P. Desert Biome