When does stomata close

To minimize transpiration, movement of gases into or out of a leaf is controlled by the stomata. The stomata are small pores in the leaf epidermis that can be opened or closed. Stomatal opening is highly regulated by multiple mechanisms so as to minimize transpiration. Transpiration is minimized even under conditions of high ambient temperature. Stomata close at high temperature. They do not open in order to cool the leaf.

Stomata are composed of two guard cells. These cells have walls that are thicker on the inner side than on the outer side. This unequal thickening of the paired guard cells causes the stomata to open when they take up water and close when they lose water. A diagram of stomata is shown on page of your text.

The opening and closing of stomata is governed by increases or decreases of solutes in the guard cells, which cause them to take up or lose water, respectively. In general, stomata open by day and close at night. During the day, photosynthesis requires that the leaf mesophyll be exposed to the air to get CO 2. At night, the stomata close to avoid losing water when photosynthesis is not occurring. During the day, stomata close if the leaves experience a lack of water, such as during a drought.

The opening or closing of stomata occur in response to signals from the external environment. Closure of stomata by drought is caused by abscisic acid, a plant hormone that is synthesized in response to drought. Abscisic acid overrides other signals and closes stomata when saving water is more important than photosynthesis.

The gene encoding this transporter is mainly expressed in the guard cells. The ABA mode of action is linked to diurnal stomatal movements. It has been proposed that this link is based on both the molecular connections between ABA and circadian-clock pathways and on ABA biosynthesis and response to light reviewed in Tallman, Although several studies have been carried out linking the diurnal cycle with ABA signaling, there is still a need for further research that would clarify this connection.

It has been confirmed that the elevated ABA levels in the dark phase of the day are responsible for stomatal closure but, on the other hand, the molecular basis of the sensing CO 2 molecules by guard cells is still not well understood.

This part of investigations still needs confirmation through the use of well-established methods. In darkness, stomata are closed. During the night, elevated levels of CO 2 in the leaves were observed due to respiration.

It has been proved that CO 2 has a positive effect on the stomatal closure process. The role of ABA in the diurnal regulation of stomatal movements. As a result of these processes, elevated levels of ABA are present in the guard cells. In the dawn B , the first light promotes ABA catabolism processes and the level of ABA biosynthesis decreases, which leads to a decreased concentration of active ABA in the guard cells.

The accumulation of sugars such as glucose, fructose and sucrose has been reported during the light phase of the day Talbott and Zeiger, In the evening, ABA biosynthesis outweighs the ABA catabolism in the guard cells, which leads to stomatal closure for review, see Tallman, Under drought stress conditions, ABA would reach a concentration high enough to cause ion efflux and an inhibition of sugar uptake by the guard cells in the midday, thus reducing the apertures for the rest of the day.

In order to define the role of ABA in stress response, the action of several components of the pathways mentioned were tested in response to stress. It has been shown that ABA concentrations can increase up to fold in response to drought stress Outlaw, Cheng et al. An immunohistochemical analysis, using antibodies raised against AtNCED3, revealed that protein is accumulated in the leaf vascular parenchyma cells in response to drought stress.

This was not detected in non-stressed conditions. These data indicate that drought-induced ABA biosynthesis occurs primarily in the vascular parenchyma cells and that vascular-derived ABA might trigger stomatal closure via the transport to the guard cells Endo et al.

AtNCED3 expression is upregulated by drought conditions across the species observed and decreases after rehydration. Drought, like the dark part of a diurnal cycle, also promotes the deconjugation of the ABA-glucose ester ABA-GE , which is stored in the vacuoles of leaf cells and also circulates in the plant Xu et al. Although its function is clear and confirmed by advanced molecular analysis, there is still a need to explain the impact of single components, such as kinases, on the regulation of the ion channels or the proton pump e.

On the other hand, the interaction between ABA regulated kinases SnRK2s and S-type anion channels, and the potassium inwardly rectifying channels, described below, has been well established and documented. Kinases are able to regulate the activity of ion channels and the proton pump. SLAC1 encodes the anion-conducting subunit of an S-type anion channel.

Increased SLAC1 activity causes an efflux of anions which results in depolarization of the membrane as a consequence of phosphorylation by SnRK. Mori et al. ABA regulation of stomatal closure during drought stress. Together, these events lead to a decrease in the turgor of the guard cells and to stomatal closure under drought conditions. The sequence of events, which is explained in detail in the main text and presented in green in the figure, is the core of the reactions that are induced or inhibited by different proteins that are activated by ABA.

Blue arrows indicate activation, while red blunt ended lines indicate inhibition. Selected genes involved in the regulation of stomatal movement under stress. Mutants in OST1 showed a wilty phenotype in water deficit conditions because of the impairment of stomatal closure and ROS production Mustilli et al. Exogenously applied NO donors triggered stomatal closure, whereas the application of an NO scavenger inhibited ABA-induced stomatal closure Neill et al. There is some evidence that both H 2 O 2 and NO actions in the guard cells require calcium.

Jasmonates are lipid-derived phytohormones that are involved in the regulation of vegetative and reproductive growth and the defense response against abiotic stress Katsir et al. JA biosynthesis is induced by stress conditions Wasternack, and many genes related to JA signaling are regulated by drought stress Huang et al.

The positive role of JA in the regulation of stomatal closure was observed in many studies Gehring et al. Similar to the ABA signaling pathway, JA signaling has been under intense investigation, particularly in relation to stress response. With the progress in research, many new components and their roles in JA-mediated stress response will be identified. Although the interaction between ABA and JA signaling pathways in stomata function has been established, there is still a need for further investigation and identification of the nodes linking these two signaling pathways, such as CPK6, which is described below.

Me-JA regulated stomatal closure during drought stress. Munemasa et al. In coi1 coronatine insensitive 1 and cpk6 mutants, the activation of S-type anion channels was disrupted Munemasa et al. Geiger et al. Hormonal crosstalk in the regulation of stomatal closure and opening during water stress.

The regulation of stomatal opening and closure is not only regulated by ABA, whose role is dominant, but also by other phytohormones. Jasmonates JA and brassinosteroids BR induce stomatal closure and inhibit stomatal opening under drought conditions, whereas the role of other hormones is ambiguous.

Cytokinins CK and auxins AUX in low physiological concentrations promote stomatal opening while in high concentrations, they are able to inhibit this process. The role of ethylene ET is the most curious. It can stimulate the closing and opening of the stomata. The details are described in the text. Suhita et al. This suggests that jasmonate-induced changes in stomatal movements require endogenous ABA.

In order to clarify this hypothesis, Hossain et al. In the wild-type, 0. Ethylene is a gaseous phytohormone that is involved in the regulation of numerous plant processes such as seed germination, root-hair growth, leaf and flower senescence and abscission, fruit ripening, nodulation, and plant responses to stresses Bleecker and Kende, It has been observed that ethylene can influence stomatal response via crosstalk with ABA; however, reports on its effect have been contradictory.

Ethylene has been linked to the promotion of both stomatal closure Pallas and Kays, and stomatal opening Madhavan et al. These contradictory effects need to be verified. One possible reason could be related to the methods used for stomatal observation that use detached leaves. Experiments with detached leaves do not always reflect the real response to stress or other applied factors in plants. Tanaka et al. This was clear evidence that ethylene repressed ABA action in stomatal closure.

In a drought stressed eto1 ethylene overproducer 1 mutant, stomata closed more slowly and were less sensitive to ABA than in the drought-treated wild type Tanaka et al. In order to elucidate the interaction between ethylene and ABA during stomatal response, epidermal peels from the wild-type and eto1 were treated with ABA, ethylene, and both phytohormones.

When ethylene was applied to the ABA-pretreated wild-type epidermal peels, an inhibition of stomatal closure was observed Tanaka et al. Desikan et al. There have been some studies that revealed both increased and decreased ethylene production in response to drought stress. However, most of them described experiments with detached leaves, which may not reflect the response of intact plants under drought conditions Morgan et al.

Generally, elevated ABA concentrations limit the production of ethylene; and therefore a dramatic increase of ABA concentration during water stress probably causes a reduction in the production of ethylene Sharp, The physiological mechanism of ethylene inhibition of the ABA-mediated stomatal closure may be related to the function of ethylene as a factor that ensures a minimum carbon dioxide supply for photosynthesis by keeping stomata half-opened under the stress conditions Leung and Giraudat, ; Tanaka et al.

Auxins and cytokinins are major phytohormones that are involved in processes related to plant growth and development such as cell division, growth and organogenesis, vascular differentiation, lateral root initiation as well as gravi- and phototropism Berleth and Sachs, The impact of cytokinins on stomatal movements is also ambiguous.

It has been shown that an increased cytokinin concentration in xylem sap promotes stomatal opening and decreases sensitivity to ABA. Generally, exogenous cytokinins and auxins can inhibit ABA-induced stomatal closure in diverse species Stoll et al. Brassinosteroids BR are polyhydroxylated steroidal phytohormones that are involved in seed germination, stem elongation, vascular differentiation, and fruit ripening Clouse and Sasse, ; Steber and McCourt, ; Symons et al. Together, these results suggest that there is an interaction between BR and ABA in drought response that is related to stomatal closure.

Many factors that are responsible for the regulation of stomatal movements have been already identified, such as components of ABA and other phytohormone signaling pathways. However, further analyses of the networks of protein interactions, the co-expression of genes, metabolic factors, etc.

Taking into account that phytohormone pathways are still under intensive investigations and there are still many gaps to be elucidated, many of the already established interactions may be changed as further progress in research is achieved.

There are ambiguous reports in regards to the role of some phytohormones, such as ethylene, auxins, or cytokinins, in the regulation of stomatal movement that need to be clarified. In addition, the interaction between the diurnal cycle and ABA pathway should be further investigated in order to achieve a full understanding of this process. There are some points that should be highlighted as a possible cause of the ambiguous reports related to the action of the regulators of stomatal movements.

The first of these is the technique that is used to observe the stomata. Most analyses of stomata under stress are based on stomatal aperture observations.

Some studies rely on stomata replicas from plants treated with stress and control, and observed under the light microscopy. This method is simple and inexpensive but generates problems due to the type of material used for the replicas. The accuracy and precision in the determination of stomatal aperture width is limited by the resolution of the standard light microscope. In contrast, scanning microscopy SEM offers high resolution images of stomata but requires expensive equipment and is not suitable for collecting large numbers of probes Lawson et al.

As long as a proper technique that is not controversial in regards to its influence on stomatal response is not applied, all aperture measurements will be under discussion. Another crucial problem is that most reports describe experiments with detached leaves, which may not reflect the response of intact plants under drought conditions Morgan et al.

Franks and Farquhar addressed the problem of data integration in stomatal research. They pointed out the lack of the integration of mechanical and quantitative physical information about guard cells and adjacent cells in model of stomatal function. Such integration of data should allow gas-exchange regulation to be better described and predicted.

As long as guard cells are considered as a model without their surroundings, the results obtained may not be relevant. Another problem noted by Franks and Farquhar is that research on the impact of various environmental factors on the stomatal regulation and stomatal density should be performed on and compared among several species, not only one.

This would allow a full picture of a broad morphological and evolutionary spectrum of possibilities of stomata development, density, and movement regulation in response to stresses to be obtained. Summarizing, there are still many questions about the techniques used for evaluating the stomatal response to stress. Further development of proper methods will bring us closer to a fuller and more relevant understanding of stomatal action.

The great progress in molecular biology studies enable insights into the signaling pathways, identification of new components, and interactions between them to be gained. The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. Further information about the project can be found at www. National Center for Biotechnology Information , U.

Journal List Front Plant Sci v. Front Plant Sci. Published online May Author information Article notes Copyright and License information Disclaimer. Received Jan 15; Accepted Apr This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in other forums, provided the original authors and source are credited and subject to any copyright notices concerning any third-party graphics etc.

This article has been cited by other articles in PMC. Abstract Two highly specialized cells, the guard cells that surround the stomatal pore, are able to integrate environmental and endogenous signals in order to control the stomatal aperture and thereby the gas exchange.

Keywords: stomata, guard cells, phytohormones, abiotic stress, ABA, jasmonic acid, crosstalk. Introduction Stomata are specialized epidermal structures that are essential for plant survival and productivity.

Open or Close the Gate — The Role of ABA, Ion Channels, and Diurnal Cycle in Stomatal Movements Regulation The regulatory role of ion channels localized in the guard cell membrane in the opening and closing stomata The guard cell turgor is dynamically adjusted to environmental conditions and hormonal signals in order to facilitate the proper gas exchange and prevent excessive water loss.

Open in a separate window. Figure 1. Abscisic acid — how the proper level of the main regulator of stomatal movements is achieved in plants Abscisic acid has been postulated as a main regulator of stomatal movements but its proper functioning depends on the appropriate level of biologically active ABA within the plant cells. Figure 2. Regulation of stomatal movements during the diurnal cycle — the role of ABA The ABA mode of action is linked to diurnal stomatal movements.

Figure 3. ABA on the way to reaching the guard cells under drought stress conditions Under drought stress conditions, ABA would reach a concentration high enough to cause ion efflux and an inhibition of sugar uptake by the guard cells in the midday, thus reducing the apertures for the rest of the day.

ABA triggers changes in ion homeostasis in the guard cells, which leads to stomatal closure under stress The ABA signaling network that leads to stomatal closure under stress is activated by the perception ABA.

Figure 4. Table 1 Selected genes involved in the regulation of stomatal movement under stress. Negative regulator of stomatal closure promoted by ABA abi2 Improper stomatal regulation leading to increased transpiration Pei et al. CPK3 is expressed in both guard cells and mesophyll cells.

Mutant phenotypes were observed in meristem organization and response to abscisic acid and drought era1 ABA hypersensitive and showed enhanced ABA activation of S-type channels Pei et al.

The protein contains one AP2 domain. Phosphorylated by PKS3 in vitro. Transcript increases under conditions that promote stomatal opening white and blue light and decreases under conditions that trigger stomatal closure ABA, desiccation, darkness with the exception of elevated CO 2.

Expressed exclusively in the guard cells of all tissues. It is required for light-induced opening of stomata myb60 Reduced stomatal aperture which helps to limit water loss during a drought Cominelli et al. Expressed in guard cells, plays a role in the regulation of stomatal pore size myb61 Larger stomatal pores than the wild-type Liang et al. Expression is upregulated in response to ABA and drought nfya5 Hypersensitive to drought because their stomata are more open than the wild-type Li et al.

The protein is expressed in guard cells and functions in stomatal opening nrt1. This closure prevents water from escaping through open pores. The opening and closing of stomata are regulated by factors such as light, plant carbon dioxide levels, and changes in environmental conditions. Humidity is an example of an environmental condition that regulates the opening or closing of stomata. When humidity conditions are optimal, stomata are open.

Should humidity levels in the air around plant leaves decrease due to increased temperatures or windy conditions, more water vapor would diffuse from the plant into the air.

Under such conditions, plants must close their stomata to prevent excess water loss. Stomata open and close as a result of diffusion. Under hot and dry conditions, when water loss due to evaporation is high, stomata must close to prevent dehydration. This causes water in the enlarged guard cells to move osmotically from an area of low solute concentration guard cells to an area of high solute concentration surrounding cells.

The loss of water in the guard cells causes them to shrink. This shrinkage closes the stomatal pore. When conditions change such that stomata need to open, potassium ions are actively pumped back into the guard cells from the surrounding cells. Water moves osmotically into guard cells causing them to swell and curve.

This enlarging of the guard cells open the pores. The plant takes in carbon dioxide to be used in photosynthesis through open stomata. Oxygen and water vapor are also released back into the air through open stomata.

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