Plant resistance to drought depends on timely stomatal closure

Stomata play a significant role in the Earth's water and carbon cycles, by regulating gaseous exchanges between the plant and the atmosphere. Under drought conditions, stomatal control of transpiration has long been thought to be closely coordinated with the decrease in hydraulic capacity (hydraulic failure due to xylem embolism). We tested this hypothesis by coupling a meta‐analysis of functional traits related to the stomatal response to drought and embolism resistance with simulations from a soil–plant hydraulic model. We report here a previously unreported phenomenon: the existence of an absolute limit by which stomata closure must occur to avoid rapid death in drought conditions. The water potential causing stomatal closure and the xylem pressure at the onset of embolism formation were equal for only a small number of species, and the difference between these two traits (i.e. safety margins) increased continuously with increasing embolism resistance. Our findings demonstrate the need to revise current views about the functional coordination between stomata and hydraulic traits and provide a mechanistic framework for modeling plant mortality under drought conditions.     

Stomatal and nonstomatal regulation of water use in cotton, corn, and sorghum

Stomata of corn (Zea mays L.) and sorghum (Sorghum bicolor L.) responded to changes in leaf water potential during the vegetative growth phase. During reproductive growth, leaf resistances were minimal and stomata were no longer sensitive to bulk leaf water status even when leaf water potentials approached −27 bars. Stomata of corn, cotton (Gossypium hirsutum L.), and sorghum appear to respond to changes in the humidity deficit between the leaf and air and in this manner, regulated transpirational flux to some degree. Distinct differences in water transport efficiency were observed in the three species. Under nonlimiting soil water conditions, sorghum exhibited the greatest efficiency of water transport while under limiting soil moisture conditions, cotton appeared most efficient. Corn was the least efficient with respect to nonstomatal regulation of water use. Differences in drought tolerance among the three species are partially dependent on stomatal regulation of water loss, but efficiency of the water transport system may be more related to drought adaptation. This is particularly important since stomata of all three species did not respond to bulk leaf water status during a large portion of the growing season.     

STOMATA ON THE FRUITS AND SEEDS OF ESCHSCHOLZIA (PAPAVERACEAE)

Stomata are present on the outer and inner fruit walls and seed coats of Eschscholzia californica, E. covillei, E. glyptosperma, E. lemmonii and E. minutiflora. The stomata on the inner fruit wall and seed coat remain constantly open, even under plasmolyzing conditions, whereas those of the outer fruit wall are able to open and close. This allows for gas exchange in these chlorophyllous structures. Fibrous bundle caps in the costal regions of the fruit act as windows allowing light transmission to the photosynthetic seeds within. Preliminary results show that the total photosynthesis by the fruits and seeds of Eschscholzia californica together appears to at least balance respiratory losses, and under favorable conditions might significantly contribute to seed and fruit development.     

Utilizing systems biology to unravel stomatal function and the hierarchies underpinning its control

Stomata control the concomitant exchange of CO₂ and transpiration in land plants. While a constant supply of CO₂ is need to maintain the rate of photosynthesis, the accompanying water losses must be tightly regulated to prevent dehydration and undesired metabolic changes. The factors affecting stomatal movement are directly coupled with the cellular networks of guard cells. Although the guard cell has been used as a model for characterization of signaling pathways, several important questions about its functioning remain elusive. Current modeling approaches describe the stomatal conductance in terms of relatively few easy‐to‐measure variables being unsuitable for in silico design of genetic manipulation strategies. Here, we argue that a system biology approach, combining modeling and high‐throughput experiments, may be used to elucidate the mechanisms underlying stomata control and to determine targets for modulation of stomatal responses to environment. In support of our opinion, we review studies demonstrating how high‐throughput approaches have provided a systems‐view of guard cells. Finally, we emphasize the opportunities and challenges of genome‐scale modeling and large‐scale data integration for in silico manipulation of guard cell functions to improve crop yields, particularly under stress conditions which are of pertinence both to climate change and water use efficiency.     

Stomatal density distribution patterns in leaves of the Jatobá (Hymenaea courbaril L.)

Stomata are leaf structures that are essential for regulating gas exchange and water balance in terrestrial plants. Accurately quantifying stomatal characteristics is consequently of great importance for understanding the physiological processes of plants under different environmental conditions. The objective of this study was to investigate the spatial distribution pattern of stomata on leaflet surfaces, and the possible mechanisms that influence this pattern, particularly leaf expansion. To achieve this, we used geostatistical tools combined with an analysis of biometric relationships of leaves from Hymenaea courbaril L. Our analysis indicates that stomata show a clear spatial structure in this species: average values of foliar expansion rates (ERs) were different on right and left-hand sides of the primary venation of each leaflet and there was a close relationship between the spatial pattern of stomatal density and leaf expansion rate. Such differences in lateral expansion may therefore be partially responsible for the heterogeneous distribution of stomata documented here and in other studies.     

Some Effects of Abscisic Acid and Water Stress on Stomata of Vicia faba L.

Vicia faba seedlings grown under a plastic tent in the laboratorywere either watered well throughout their growth period or weresubjected to a water stress treatment for several days priorto an experimental treatment. The effects of a further waterstress treatment or an application of an aqueous solution ofabscisic acid (ABA) on the stomata of these plants were determined.Stomata of previously water-stressed plants proved to be moresensitive than stomata of well watered plants to ABA appliedthrough the petiole via the transpiration stream and sprayedonto leaf surfaces. Stomata of previously water-stressed plantsclosed more rapidly and to a greater degree than stomata ofwell watered plants. The hormone had only a small effect whenapplied directly to epidermal fragments removed from both groupsof plants. Stomata of plants which had received a water stresspretreatment were less sensitive to a subsequent period of waterstress than were stomata of previously well watered plants.It is proposed that stomatal adaptation to water stress maybe related to changes in the hormonal balance of the plant.     

Evidence for the Uptake of Large Anions through Stomatal Pores

Experiments were conducted with leek (Allium porrum L.) leaves to investigate whether aqueous solutions are able to penetrate stomata. Epidermal strips were used for the determination of transport rates. Stomata were opened by fusicoccin or closed by darkness or abscisic acid. A droplet containing the anionic fluorescent dye, uranine, was placed on the physiologically outer side of the epidermis and allowed to dry. With open stomata 30 times more uranine penetrated through the epidermal strips than with closed stomata (comparison of medians). In another experiment droplets of uranine solution were placed on leaf segments and epidermal strips were removed after drying of the droplets. Penetration of uranine through stomata was detectable under the microscope both with epidermal strips from the transport experiments and with strips obtained after application on leaf segments. As maximum uptake rates occurred during the drying process, it is concluded that penetration took place via water films. These results show that the physical restrictions preventing stomatal penetration of static droplets are not decisive for drying droplets and that stomatal uptake of dissolved ionic substances occurs under natural conditions, i.e. without surfactants or applied pressure.     

The blue light-specific response of Vicia faba stomata acclimates to growth environment

Stomata in epidermal strips from growth chamber-grown Vicia faba leaves opened less in response to white light than did stomata from greenhouse-grown leaves. Chlorophyll-mediated, red light-stimulated opening was similar in stomata from the two growth conditions, but stomata from the growth chamber environment had a severely reduced response to blue light. Transfer of plants between the two growth conditions resulted in an acclimation of the stomatal blue light response. Stomata lost blue light sensitivity within 1 d of transfer to growth chamber conditions and gained sensitivity to blue light over an 8 d period after transfer to a greenhouse. Short-term transfer experiments confirmed that the rapid loss of blue light sensitivity was an acclimation response, requiring between 12 and 20 h exposure to growth chamber conditions. The acclimation of the stomatal response to blue light was inversely related to a previously reported acclimation response in which stomata change between high CO2 sensitivity under growth chamber conditions and low CO2 sensitivity under greenhouse conditions. The time courses of the blue light and CO2 acclimation responses were virtually identical, suggesting the possibility of a common acclimation mechanism.     

The ultrastructure of micropropagated and greenhouse rose plant stomata

Stomata of leaves from in vitro grown rose plantlets remain opened in the dark. The ultrastructure of their guard cells was studied after a 7 h light and a 7 h dark period, and compared to that of functional stomata from plants which have been acclimatized to greenhouse conditions. Qualitative and quantitative observations concerning the shape of the guard cells, mitochondria, plastids and starch grains, demonstrated the similarity in guard cell ultrastructure. The peculiarity of guard cell ultrastructure of in vitro cultured plants was the inability to close in the dark; vacuolar area was 40% of the whole guard cell area during both light and dark period whereas, in guard cells from greenhouse plants, the vacuolar area was 40% of the whole guard cell area during the light and only 25% during the dark period. These results indicate that stomata from in vitro plants are duly developed and possess an ultrastructure suitable for a typical functioning. The inability to close in the dark results from atypical water relation.     

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.     

Stomatal Features in the Leaf of Aganosma dichotoma (Roth) K. Schum

Paracytic and anisocytic types of mature stomata are found inthe leaf of Aganosma dichotoma. Stomata with one guard cell,stomata with degenerated guard cells, and contiguous stomataare common. Stomata with arrested pore development are alsofound in certain cases. A single guard cell without any porehas not been designated as a stoma with one guard cell in thepresent investigation. Ontogeny of contiguous stomata have beentraced. Subsidiary cells are, morphologically, just like theircontiguous guard cells. Subsidiary cells may retain their shapeand contents even when their contiguous stoma becomes mature,or may change their shape and lose their contents. They mayor may not divide. Subsidiary cells form a whorl of more thantwo subsidiary cells around a stoma by their divisions. Degenerationof guard cell(s)— their contents and nuclei—havebeen traced. In certain cases guard cells divide forming morethan two guard cells associated to a single pore. Cytoplasmicconnections are found between two guard cells of nearby stomata,and between a guard cell and an epidermal cell. Near the wound,the epidermal cells over the veins become meristermatic givingrise to new epidermal cells but no meristemoid.     

Stomatal Opening Mechanism of CAM Plants

Stomata usually open when leaves are transferred from darkness to light. However, reverse-phase stomatal opening in succulent plants has been known. CAM plants such as cacti and Opuntia ficus–indica achieve their high water use efficiency by opening their stomata during the cool, desert nights and closing them during the hot, dry days. Signal transduction pathway for stomatal opening by blue light photoreceptors including phototropins and the carotenoid pigment zeaxanthin has been suggested. Blue light regulated signal transduction pathway on stomatal opening could not be applied to CAM plants, but the most possible theory for a nocturnal response of stomata in CAM plants is photoperiodic circadian rhythm.     

水分亏缺对冬小麦净光合速率影响程度研究

水分亏缺对冬小麦净光合速率影响程度研究王慧（西北大学城市与资源学系,西安７１００６９）ＥｆｆｅｃｔｏｆＷａｔｅｒＤｅｆｉｃｉｔｏｎＮｅｔＰｈｏｔｏｓｙｎｔｈｅｓｉｓＲａｔｅｏｆＷｉｎｔｅｒＷｈｅａｔ．ＷａｎｇＨｕｉ（ＤｅｐａｔｍｅｎｔｏｆＵｒｂａｎ...     

Secret Message at the Plant Surface

In general, stomata open during the day and close at night. This behavior has a crucial importance because it maximizes the update of CO₂ for photosynthesis and minimizes the water loss. Blue light is one of the environmental factors that regulates this process. Certainly, when either entire plants or epidermal strips adapted to the dark are exposed to blue light, the stomata open widely their pores. But, what does happen if we illuminate individual stomata instead of peels or entire plants? In the inaugural issue of PLoS ONE, we have answered this question by irradiating individual stomata with a laser attached to a confocal microscope. Our study not only demonstrates that the stomata function independently from the behavior of their neighbors, and illuminates the implication of the blue light receptors PHOTOTROPIN1 and PHOTOTROPIN2 in such response. It also gives clues about the physiological relevancy of this behavior.Key WordS: Stomata, phototropins, autonomous function, blue light, cellular irradiation     

Functional Stomata in a Variegated Leaf Chimera of Pelargonium zonale L. without Guard Cell Chloroplasts

Fluorescence microscopy indicated that chlorophyll was absentfrom epidermal and guard cells overlying all white areas andgreen areas (of certain leaves) in variegated leaves of Pelargoniumzonale, cv. Chelsea Gem. Stomata with chlorophyll-free guardcells, in general, responded normally to light and CO₂ as gaugedby direct measurements of stomatal aperture and by transpirationalwater loss studies, although stomata from white regions of variegatedleaves were more reluctant to open than stomata from green regionsof the leaves. Thus, functional stomata without guard cell chloroplastshave been discovered in another genus, namely Pelargonium, besidesthat originally discovered in Paphiopedilum. When stomata withchlorophyll-free guard cells opened, K⁺ accumulated in the guardcells. This indicates that chloroplasts are not essential forthe normal functioning of stomata and that the energy sourcefor driving stomatal movements can come from sources other thanphotophosphorylation. Key words: Guard cell chloroplasts, Leaf chimera, Pelargonium, Stomata     

Hydroreactivity of stomata in kale leaves of different insertion level as determined by analysis of transpiration curves

Values of the water saturation deficit (WSD) for hydroactive stomatal movements of kale leaves were estimated using the method of transpiration curve analysis. Stomata of young leaves started closing at WSD values of 5 to 6 per cent and were completely closed at 18 to 20 per cent WSD. During maturation and ageing of leaves these WSD values increased to 12.5 and 18 to 23 per cent respectively. Thus the stomatal reaction is more sensitive to changes in WSD in adult leaves than in young ones. After maturation is attained both values decrease. In apparently withering leaves the individual phases of transpiration curves can barely be distinguished, probably for the reason that even under optimal conditions their stomata remain half-closed and at high WSD values an incomplete closing of the aperture occurs. The injured cuticle of withering leaves affects the shape of the transpiration curve as well.     

Root to Shoot Communication in Maize Plants of the Effects of Soil Drying

Seedlings of Zea mays L. (John Innes hybrid) were grown withroots divided between two containers such that part of the rootsystem could reduce the water potential of the soil in its immediatevicinity while the rest of the root system was well suppliedwith water. When compared to plants rooted in two pots of moistsoil, drying of part of the root system resulted in partialclosure of stomata, even though leaf water potential, turgorand abscisic acid (ABA) content remained unaffected. When leafpieces were removed from the two groups of plants and incubatedunder conditions favourable for stomatal opening, stomata ofthe ‘half-watered’ plants still showed restrictedapertures. Incubation in kinetin (10 mmol m^–3) or zeatin(100 mmol m^–3) reversed the closure of stomata stimulatedby soil drying. These results suggest that a continuous supplyof cytokinin from roots may be necessary to sustain maximalstomatal opening and an interruption of this supply due to soildrying may act as an indicator of inhibited root activity, resultingin restricted stomatal opening and thereby restricted wateruse. Key words: Zea mays L., Soil drying, Stomata, Roots     

Circadian Rhythms of Leaf and Stomatal Movements in Gymnosperm Species

It is generally accepted that various physiological, morphological and gene expression phenomena are under the control of a circadian clock, and that this time keeping mechanism is universally present. Although such endogenously regulated phenomena have first been documented in plants more than 250 years ago and much work has been accumulated particularly in the past 70 years, it was not obvious from the literature whether such time keeping mechanisms exist in gymnosperms. Two prominent parameters were investigated in several gymnosperm species which have been demonstrated to be under the control of a circadian clock in many plants: (i) leaf movement and (ii) stomata movement. In young plants of Pinus sylvestris, Picea abies, Taxus baccata, Araucaria angustifolia, Araucaria heterophylla and Ginkgo biloba leaf oscillations could be recorded for about 5 days. However, compared to angiosperm plants, the amplitude was small. The period length under free running conditions (constant temperature and continuous light) was characteristic for the species. Stomatal movement was observed in Ginkgo biloba leaves by electron microscopy. Stomata were open at noon and closed at midnight under normal day/night conditions (LD) as well as under constant light conditions (LL), indicating that stomatal aperture is under circadian control in the gymnosperm Ginkgo biloba. Online recordings of stomata conductance however, exhibited diurnal but not circadian oscillations of net CO₂-exchange in G. biloba leaves. Our results show that a circadian clock controls leaf and stomatal movements in gymnosperm species indicating that endogenous time keeping mechanisms are present.     

Gas exchange of a desert shrub (Zygophyllum dumosum Boiss.) under different soil moisture regimes during summer drought

The effects of soil water potential on photosynthesis and transpiration of whole Zygophyllum dumosum Boiss. shrubs were examined with a field IRGA system during a rainless summer. Daily photosynthesis and transpiration activities were not notably different on a unit phyllode area basis among shrubs at naturally differing soil water potentials. Irrigation of shrubs caused phyllodes to increase significantly in water content and new leaflets to appear. Leaflets had three times as many stomata per unit area (23000 stomata cm^-2) as phyllodes (7100 stomata cm^-2) but photosynthesis and transpiration rates were not measurably different between irrigated and non-irrigated shrubs on a unit area basis. This finding suggests that sufficient soil moisture will lead to increased carbon uptake of the entire shrub simply because the total area of photosynthesizing tissue increases. Gas exchange rates appear to be controlled solely by atmospheric conditions under the stresses of summer.