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Abscisic acid (ABA) can induce rapid stomatal closure in seed plants, but the action of this hormone on the stomata of fern and lycophyte species remains equivocal. Here, ABA-induced stomatal closure, signaling components, guard cell K+ and Ca2+ fluxes, vacuolar and actin cytoskeleton dynamics, and the permeability coefficient of guard cell protoplasts (Pf) were analyzed in species spanning the diversity of vascular land plants including 11 seed plants, 6 ferns, and 1 lycophyte. We found that all 11 seed plants exhibited ABA-induced stomatal closure, but the fern and lycophyte species did not. ABA-induced hydrogen peroxide elevation was observed in all species, but the signaling pathway downstream of nitric oxide production, including ion channel activation, was only observed in seed plants. In the angiosperm faba bean (Vicia faba), ABA application caused large vacuolar compartments to disaggregate, actin filaments to disintegrate into short fragments and Pf to increase. None of these changes was observed in the guard cells of the fern Matteuccia struthiopteris and lycophyte Selaginella moellendorffii treated with ABA, but a hypertonic osmotic solution did induce stomatal closure in fern and the lycophyte. Our results suggest that there is a major difference in the regulation of stomata between the fern and lycophyte plants and the seed plants. Importantly, these findings have uncovered the physiological and biophysical mechanisms that may have been responsible for the evolution of a stomatal response to ABA in the earliest seed plants., (© American Society of Plant Biologists 2021. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Ethylene, a plant hormone that significantly influences both plant growth and response to stress, plays a well-established role in stress signaling. However, its impact on stomatal opening and closure during dehydration and rehydration remains relatively unexplored and is still debated. Exogenous ethylene has been proven to induce stomatal closure through a series of signaling pathways, including the accumulation of reactive oxygen species, subsequent synthesis of nitric oxide and hydrogen sulfide, and SLOW ANION CHANNEL-ASSOCIATED 1 activation. Thus, it has been suggested that ethylene might function to induce stomatal closure synergistically with abscisic acid (ABA). Furthermore, it has also been shown that increased ethylene can inhibit ABA- and jasmonic acid-induced stomatal closure, thus hindering drought-induced closure during dehydration. Simultaneously, other stresses, such as chilling, ozone pollution, and K+ deficiency, inhibit drought- and ABA-induced stomatal closure in an ethylene synthesis-dependent manner. However, ethylene has been shown to take on an opposing role during rehydration, preventing stomatal opening in the absence of ABA through its own signaling pathway. These findings offer novel insights into the function of ethylene in stomatal regulation during dehydration and rehydration, giving a better understanding of the mechanisms underlying ethylene-induced stomatal movement in seed plants. [ABSTRACT FROM AUTHOR]

While variation in mean annual precipitation (MAP) of the native habitat of a species has been shown to determine the ability of a species to resist a hydraulic decrease during drought, it remains unknown whether these variations in MAP also influence the ability of a species to recover and survive drought. Leaf hydraulic and gas exchange recovery following drought and the underlying mechanisms of these responses in six Caragana species from habitats along a large precipitation gradient were investigated during rehydration in a common garden. The gas exchange of species from arid habitats recovered more rapidly during rehydration after mild, moderate and severe drought stress treatments than species from humid habitats. The recovery of gas exchange was not associated with foliar abscisic acid concentration, but tightly related to the recovery of leaf hydraulic conductance (Kleaf). The recovery of Kleaf was associated with the loss of Kleaf during dehydration under mild and moderate drought stress, and to leaf xylem embolism formation under severe drought stress. Results pointed to the different ability to recover in gas exchange in six Caragana species post‐drought is associated with the MAP of the species in its native habitat. Summary Statement: Mean annual precipitation (MAP) has been shown to shape species' ability to resist a hydraulic dysfunction during drought. Our findings suggest that the ability to recover in gas exchange post‐drought as a strong driver in shaping species' distribution along a MAP gradient. [ABSTRACT FROM AUTHOR]

Stomatal closure is regulated by plant hormones and some small molecules to reduce water loss under stress conditions. Both abscisic acid (ABA) and polyamines alone induce stomatal closure; however, whether the physiological functions of ABA and polyamines are synergistic or antagonistic with respect to inducing stomatal closure is still unknown. Here, stomatal movement in response to ABA and/or polyamines was tested in Vicia faba and Arabidopsis thaliana, and the change in the signaling components under stomatal closure was analyzed. We found that both polyamines and ABA could induce stomatal closure through similar signaling components, including the synthesis of hydrogen peroxide (H2O2) and nitric oxide (NO) and the accumulation of Ca2+. However, polyamines partially inhibited ABA‐induced stomatal closure both in epidermal peels and in planta by activating antioxidant enzymes, including superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT), to eliminate the ABA‐induced increase in H2O2. These results strongly indicate that polyamines inhibit abscisic acid‐induced stomatal closure, suggesting that polyamines could be used as potential plant growth regulators to increase photosynthesis under mild drought stress. [ABSTRACT FROM AUTHOR]

Summary: Plants control water‐use efficiency (WUE) by regulating water loss and CO2 diffusion through stomata. Variation in stomatal control has been reported among lineages of vascular plants, thus giving rise to the possibility that different lineages may show distinct WUE dynamics in response to water stress.Here, we compared the response of gas exchange to decreasing leaf water potential among four ferns and nine seed plant species exposed to a gradually intensifying water deficit. The data collected were combined with those from 339 phylogenetically diverse species obtained from previous studies.In well‐watered angiosperms, the maximum stomatal conductance was high and greater than that required for maximum WUE, but drought stress caused a rapid reduction in stomatal conductance and an increase in WUE in response to elevated concentrations of abscisic acid. However, in ferns, stomata did not open beyond the optimum point corresponding to maximum WUE and actually exhibited a steady WUE in response to dehydration. Thus, seed plants showed improved photosynthetic WUE under water stress.The ability of seed plants to increase WUE could provide them with an advantage over ferns under drought conditions, thereby presumably increasing their fitness under selection pressure by drought. [ABSTRACT FROM AUTHOR]

Drought is a cyclical phenomenon in natural environments. During dehydration, stomatal closure is mainly regulated by abscisic acid (ABA) dynamics that limit transpiration in seed plants, but following rehydration, the mechanism of gas exchange recovery is still not clear. In this study, leaf water potential (ψleaf), stomatal conductance (gs), leaf hydraulic conductance (Kleaf), foliar ABA level, ethylene emission rate in response to dehydration and rehydration were investigated in four Caragana species with isohydric (Caragana spinosa and C. pruinosa) and anisohydric (C. intermedia and C. microphylla) traits. Two isohydric species with ABA‐induced stomatal closure exhibited more sensitive gs and Kleaf to decreasing ψleaf than two anisohydric species which exhibited a switch from ABA to water potential‐driven stomatal closure during dehydration. Following rehydration, the recovery of gas exchange was not associated with a decrease in ABA level but was strongly limited by the degradation of the ethylene emission rate in all species. Furthermore, two anisohydric species with low drought‐induced ethylene production exhibited more rapid recovery in gas exchange upon rehydration. Our results indicated that ethylene is a key factor regulating the drought‐recovery ability in terms of gas exchange, which may shape species adaptation to drought and potential species distribution. Drought is a cyclical phenomenon in natural environments. During dehydration, stomatal closure is mainly regulated by abscisic acid (ABA) dynamics that limit transpiration in seed plants, but the mechanisms of gas exchange recovery are still not clear following rehydration. The results in this study indicate that ethylene is a key factor regulating the drought‐recovery ability in terms of gas exchange, which may shape species adaptation to drought and potential species distribution. [ABSTRACT FROM AUTHOR]

Summary: Clarifying the coordination of leaf hydraulic traits with gas exchange across closely‐related species adapted to varying rainfall can provide insights into plant habitat distribution and drought adaptation.The leaf hydraulic conductance (Kleaf), stomatal conductance (gs), net assimilation (A), vein embolism and abscisic acid (ABA) concentration during dehydration were quantified, as well as pressure–volume curve traits and vein anatomy in 10 Caragana species adapted to a range of mean annual precipitation (MAP) conditions and growing in a common garden.We found a positive correlation between Ψleaf at 50% loss of Kleaf (KleafP50) and maximum Kleaf (Kleaf‐max) across species. Species from low‐MAP environments exhibited more negative KleafP50 and turgor loss point, and higher Kleaf‐max and leaf‐specific capacity at full turgor, along with higher vein density and midrib xylem per leaf area, and a higher ratio of Kleaf‐max : maximum gs. Tighter stomatal control mediated by higher ABA accumulation during dehydration in these species resulted in an increase in hydraulic safety and intrinsic water use efficiency (WUEi) during drought.Our results suggest that high hydraulic safety and efficiency combined with greater stomatal sensitivity triggered by ABA production and leading to greater WUEi provides drought tolerance in Caragana species adapted to low‐MAP environments. [ABSTRACT FROM AUTHOR]

Brassinosteroids (BRs) are well recognized for their defensive role in plants under abiotic stress conditions, but 28-homobrassinolide (HBR)-induced tolerance to drought stress has not been reported in soybean (Glycine max L.). The present study investigated the effect of HBR on soybean seedlings under drought stress. Drought stress suppressed growth and photosynthetic systems while increased the proline, glycine betaine (GB), anthocyanin, total phenolic (TP), and total flavonoid (TF) levels in soybean seedlings. HBR restricted reactive oxygen species (ROS) accumulation and decreased the hydrogen peroxide (H2O2) and malondialdehyde (MDA) content by triggering the antioxidant systems. HBR acts as a shield in soybean, protecting the plant against the harmful effects of methylglyoxal (MG) effects by upregulating the enzymes glyoxalase I, (Gly I;15%) and glyoxalase II (Gly II;29.1%) compared to the levels in drought stressed seedlings. Overall, HBR improved drought tolerance in soybean seedlings by modulating osmolytes, the AsA–GSH cycle, and enzyme activities. [ABSTRACT FROM AUTHOR]

Ozone (O3) is a gaseous environmental pollutant that can enter leaves through stomatal pores and cause damage to foliage. It can induce oxidative stress through the generation of reactive oxygen species (ROS) like hydrogen peroxide (H2O2) that can actively participate in stomatal closing or opening in plants. A number of phytohormones, including abscisic acid (ABA), ethylene (ET), salicylic acid (SA), and jasmonic acid (JA) are involved in stomatal regulation in plants. The effects of ozone on these phytohormones' ability to regulate the guard cells of stomata have been little studied, however, and the goal of this paper is to explore and understand the effects of ozone on stomatal regulation through guard cell signaling by phytohormones. In this review, we updated the existing knowledge by considering several physiological mechanisms related to stomatal regulation after response to ozone. The collected information should deepen our understanding of the molecular pathways associated with response to ozone stress, in particular, how it influences stomatal regulation, mitogen-activated protein kinase (MAPK) activity, and phytohormone signaling. After summarizing the findings and noting the gaps in the literature, we present some ideas for future research on ozone stress in plants [ABSTRACT FROM AUTHOR]

In recent years, research on spermine (Spm) has turned up a lot of new information about this essential polyamine, especially as it is able to counteract damage from abiotic stresses. Spm has been shown to protect plants from a variety of environmental insults, but whether it can prevent the adverse effects of drought has not yet been reported. Drought stress increases endogenous Spm in plants and exogenous application of Spm improves the plants' ability to tolerate drought stress. Spm's role in enhancing antioxidant defense mechanisms, glyoxalase systems, methylglyoxal (MG) detoxification, and creating tolerance for drought-induced oxidative stress is well documented in plants. However, the influences of enzyme activity and osmoregulation on Spm biosynthesis and metabolism are variable. Spm interacts with other molecules like nitric oxide (NO) and phytohormones such as abscisic acid, salicylic acid, brassinosteroids, and ethylene, to coordinate the reactions necessary for developing drought tolerance. This review focuses on the role of Spm in plants under severe drought stress. We have proposed models to explain how Spm interacts with existing defense mechanisms in plants to improve drought tolerance. [ABSTRACT FROM AUTHOR]

Plants secreted phytocytokine SMALL PHYTOCYTOKINES REGULATING DEFENSE AND WATER LOSS (SCREWs) and its receptor PLANT SCREW UNRESPONSIVE RECEPTOR (NUT) to counter abscisic acid (ABA)- and pathogen-induced stomatal closure (Liu et al.). This novel signaling process provides plants with a new strategy to increase immunity through disrupting an aqueous habitat for pathogen colonization. [ABSTRACT FROM AUTHOR]

• ABA is involved in stomatal closure in ferns, gymnosperms and angiosperms. • P type and R type ABA responses occur during dehydration and rehydration in gymnosperms as well as angiosperms. • Increasing and decreasing behaviour of foliar ABA content in angiosperms during dehydration. The variation in stomatal movement across vascular plants in response to abscisic acid (ABA) has received great attention in recent years. This review is an attempt to better understand the role of ABA in the stomatal movement of seed and seedless plants under dehydration and rehydration. Seed plants, i.e., gymnosperms and angiosperms, have two divergent ABA responses, the peaking type (P-type) and rising type (R-type), to induce stomatal closure under sustained drought stress. However, in the case of ferns and lycophytes, stomata of almost all species exhibit insensitive (I-type) behaviour to ABA. Consequently, seed plants have evolved an optimized water use efficiency to improve their succession in terrestrial ecosystems. During rehydration, the recovery of gas exchange is constrained by ABA accumulation under drought in R-type plants and constrained by hydraulics in P- and I-type plants. Thus, future studies should investigate the mechanisms underlying the divergence in stomata in response to ABA, focusing on P-type vs. R-type ABA responses in seed plants, the competition of seed plants, and the interaction between ABA and hydraulic pathways during rehydration. [ABSTRACT FROM AUTHOR]