Ethylene-mediated stomatal responses to dehydration and rehydration in seed plants.

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 (ROS), subsequent synthesis of nitric oxide (NO) and hydrogen sulfide (H2S), and SLOW ANION CHANNEL-ASSOCIATED 1 (SLAC1) 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 (JA)-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 through an ethylene synthesis dependent way. 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, gaining a better understanding of the mechanisms underlying ethylene-induced stomatal movement in seed plants.

Rapid drought-recovery of gas exchange in Caragana species adapted to low mean annual precipitation.

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.

Polyamines inhibit abscisic acid-induced stomatal closure by scavenging hydrogen peroxide.

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 (H2 O2 ) 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 H2 O2 . 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.

Aging rate, environmental risk and production efficiency of the low-density polyethylene (LDPE) films with contrasting thickness in irrigated region.

Physical thickness of low-density polyethylene (LDPE) films might determine the release rate of phthalic acid esters (PAEs) & structural integrity and affect production efficiency. However, this critical issue is still unclear and little reported. Aging effects were evaluated in LDPE films with the thickness of 0.006, 0.008, 0.010 and 0.015 mm in a maize field of irrigation region. The Scanning electron microscope (SEM) results showed that the proportion of damaged area (Dam) to total area of LDPE films was massively lowered with increasing thickness after aging. The highest and lowest Dam was 32.2% and 3.5% in 0.006 and 0.015 mm films respectively. Also, the variations in peak intensity of asymmetric & symmetrical stretching vibrations (ASVI & SSVI) were detected using Fourier transform infrared spectrum (FTIR), indicating that the declines in peak intensity tended to be slower with thickness. Interestingly, the declines in physical integrity were tightly associated with increasing exhalation rate of PAEs. Average releasing rate of PAEs was 38.2%, 31.4%, 31.5% and 19.7% in LDPE films from 0.006 to 0.015 mm respectively. Critically, thicker film mulching can lead to greater soil water storage at plough layer (SWS-PL) and better thermal status, accordingly harvesting higher economic benefit. Therefore, LDPE film thickening may be a solution to reduce environmental risk but improve production efficiency in arid region.

Stomatal morphology and physiology explain varied sensitivity to abscisic acid across vascular plant lineages.

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.

Evolution of stomatal closure to optimize water-use efficiency in response to dehydration in ferns and seed plants.

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.

Combined high leaf hydraulic safety and efficiency provides drought tolerance in Caragana species adapted to low mean annual precipitation.

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 (Kleaf P50 ) and maximum Kleaf (Kleaf-max ) across species. Species from low-MAP environments exhibited more negative Kleaf P50 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.

Ethylene, not ABA, is closely linked to the recovery of gas exchange after drought in four Caragana species.

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.

A clear trade-off between leaf hydraulic efficiency and safety in an aridland shrub during regrowth.

It has been suggested that a trade-off between hydraulic efficiency and safety is related to drought adaptation across species. However, whether leaf hydraulic efficiency is sacrificed for safety during woody resprout regrowth after crown removal is not well understood. We measured leaf water potential (ψleaf ) at predawn (ψpd ) and midday (ψmid ), leaf maximum hydraulic conductance (Kleaf-max ), ψleaf at induction 50% loss of Kleaf-max (Kleaf P50 ), leaf area-specific whole-plant hydraulic conductance (LSC), leaf vein structure and turgor loss point (πtlp ) in 1- to 13-year-old resprouts of the aridland shrub (Caragana korshinskii). ψpd was similar, ψmid and Kleaf P50 became more negative, and Kleaf-max decreased in resprouts with the increasing age; thus, leaf hydraulic efficiency clearly traded off against safety. The difference between ψmid and Kleaf P50 , leaf hydraulic safety margin, increased gradually with increasing resprout age. More negative ψmid and Kleaf P50 were closely related to decreasing LSC and more negative πtlp , respectively, and the decreasing Kleaf-max arose from the lower minor vein density and the narrower midrib xylem vessels. Our results showed that a clear trade-off between leaf hydraulic efficiency and safety helps C. korshinskii resprouts adapt to increasing water stress as they approach final size.

Dew absorption by leaf trichomes in Caragana korshinskii: An alternative water acquisition strategy for withstanding drought in arid environments.

Investigating plant morphological traits can provide insights into plant drought tolerance. To date, many papers have focused on plant hydraulic responses to drought during dehydration, but atmospheric water absorption by trichomes to mitigate drought stress by influencing leaf hydraulics in plant species that inhabit arid environments has been largely ignored. The experiment in this study was designed to assess how dew absorbed by leaf trichomes helps Caragana korshinskii withstand drought. The results showed that under a drought stress and dew (DS & D) treatment, C. korshinskii displayed a strong capacity to absorb dew with trichomes; exhibited slow decreases in leaf water potential (Ψleaf ), leaf hydraulic conductivity (Kleaf ), and gas exchange; experienced 50% Kleaf and gas exchange losses at lower relative soil water content levels than plants treated with drought stress and no dew (DS & ND); and experienced 50% Kleaf loss (Kleaf P50 ) at similar Ψleaf levels as DS & ND plants. Its congener C. sinica, which does not have leaf trichomes, displayed little ability to absorb dew under drought stress and did not show any remarkable improvement in the above parameters under the DS & D treatment. Our results indicated that leaf trichomes are important epidermal dew-uptake structures that assist in partially sustaining the leaf hydraulic assimilation system, mitigate the adverse effects of drought stress and contribute to the distribution of C. korshinskii in arid environments.

GABA: A Key Player in Drought Stress Resistance in Plants.

γ-aminobutyric acid (GABA) is a non-protein amino acid involved in various physiological processes; it aids in the protection of plants against abiotic stresses, such as drought, heavy metals, and salinity. GABA tends to have a protective effect against drought stress in plants by increasing osmolytes and leaf turgor and reducing oxidative damage via antioxidant regulation. Guard cell GABA production is essential, as it may provide the benefits of reducing stomatal opening and transpiration and controlling the release of tonoplast-localized anion transporter, thus resulting in increased water-use efficiency and drought tolerance. We summarized a number of scientific reports on the role and mechanism of GABA-induced drought tolerance in plants. We also discussed existing insights regarding GABA's metabolic and signaling functions used to increase plant tolerance to drought stress.

Ozone Induced Stomatal Regulations, MAPK and Phytohormone Signaling in Plants.

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.

Small ubiquitin-like modifiers E3 ligases in plant stress.

Plants regularly encounter various environmental stresses such as salt, drought, cold, heat, heavy metals and pathogens, leading to changes in their proteome. Of these, a post-translational modification, SUMOylation is particularly significant for its extensive involvement in regulating various plant molecular processes to counteract these external stressors. Small ubiquitin-like modifiers (SUMO) protein modification significantly contributes to various plant functions, encompassing growth, development and response to environmental stresses. The SUMO system has a limited number of ligases even in fully sequenced plant genomes but SUMO E3 ligases are pivotal in recognising substrates during the process of SUMOylation. E3 ligases play pivotal roles in numerous biological and developmental processes in plants, including DNA repair, photomorphogenesis, phytohormone signalling and responses to abiotic and biotic stress. A considerable number of targets for E3 ligases are proteins implicated in reactions to abiotic and biotic stressors. This review sheds light on how plants respond to environmental stresses by focusing on recent findings on the role of SUMO E3 ligases, contributing to a better understanding of how plants react at a molecular level to such stressors.

Ethylene constrains stomatal reopening in Fraxinus chinensis post moderate drought.

Clarifying the mechanisms underlying the recovery of gas exchange following drought is the key to providing insights into plant drought adaptation and habitat distribution. However, the mechanisms are still largely unknown. Targeting processes known to inhibit gas exchange during drought recovery, we measured leaf water potential, the leaf hydraulic conductance, stomatal reopening, abscisic acid (ABA) and the ethylene emission rate (EER) following moderate drought stress in seedlings of the globally pervasive woody tree Fraxinus chinensis. We found strong evidence that the slow stomatal reopening after rehydration is regulated by a slow decrease in EER, rather than changes in leaf hydraulics or foliar ABA levels. This was supported by evidence of rapid gas exchange recovery in plants after treatment with the ethylene antagonist 1-methylcyclopropene. These findings provide evidence to rigorously support ethylene as a key factor constraining stomatal reopening from moderate drought directly, thereby potentially opening new windows for understanding species drought adaptation.

Phytocytokine SCREWs increase plant immunity through actively reopening stomata.

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.

Light: a crucial factor for rhizobium-induced root nodulation.

Wang et al. recently showed that, in soybean (Glycine max), root nodule formation is induced by a light-triggered signal that moves from the upper part of the plant to the roots. This novel signaling process opens a new area of research aimed to optimize the carbon-nitrogen balance in plant-rhizobium symbiosis.

ABA activated SnRK2 kinases: an emerging role in plant growth and physiology.

Members of the SNF1-related protein kinase 2 (SnRK2) family are plant-specific serine or threonine kinases that play a pivotal role in the response of plants to abiotic stresses. Members of this plant-specific kinase family have included a critical regulator (SnRK2) of abscisic acid (ABA) response in plants. Plant organ development is governed substantially by the interaction of the SnRK2 and the phytohormone abscisic acid (ABA). Recent research has revealed a synergistic link between SnRK2 and ABA signaling in a plant's response to stress such as drought and shoot growth. SnRK2 kinases play a dual role in the control of SnRK1 and the development of a plant. The dual role of SnRK2 kinases promotes plant growth under optimal conditions and in the absence of ABA while inhibiting the growth of plants in response to ABA. In this review, we have uncovered the roles of ABA-activated SnRK2 kinases in plants, as well as their physiological mechanisms.

Bacillus amyloliquefaciens GB03 augmented tall fescue growth by regulating phytohormone and nutrient homeostasis under nitrogen deficiency.

Nitrogen is an important nutrient for plant growth and development. Soil microorganisms have been used to curb the imbalance between the limited content of natural environmental nitrogen and the pollution caused by increasing nitrogen fertilizer use in ecologically fragile areas. Bacillus amyloliquefaciens GB03 has been shown to confer growth promotion and abiotic stress tolerance in Arabidopsis thaliana. This study provided a new insight into the role of the plant growth-promoting rhizobacterium B. amyloliquefaciens GB03 as an initiator of defense against nitrogen deficiency in non-leguminous grass tall fescue (Festuca arundinacea). Two-week-old seedlings of tall fescue were grown with or without GB03 for 4 weeks under total nitrogen (3.75 mM NO3 -) or low nitrogen (0.25 mM NO3 -) treatment. Growth parameters, chlorophyll content, endogenous total nitrogen, total phosphorus content, and phytohormone content, including those of auxin indole-3-acetic acid, cytokinin, gibberellic acid, and abscisic acid, were determined at the time of harvest. Tall fescue grown in GB03-inoculated soil was more robust than the non-inoculated controls with respect to plant height, root length, plant biomass, chlorophyll concentration, and nutrient (total nitrogen and total phosphorus) contents under total nitrogen treatment. GB03 increased indole acetic acid content by 24.7%, whereas decreased cytokinin and abscisic acid contents by 28.4% and 26.9%, respectively, under a total nitrogen level. Remarkably, GB03 increased indole acetic acid content by more than 80% and inhibited abscisic acid production by nearly 70% under a low nitrogen level. These results showed, for the first time, that GB03 played a crucial role in mediating NO3 -dependent regulation of tall fescue growth and development, especially revealing the mechanism of soil bacteria improve resistance to nitrogen deficiency stress in non-nitrogen-fixing species.

Divergent stem hydraulic strategies of Caragana korshinskii resprouts following a disturbance.

Resprouting plants are distributed in many vegetation communities worldwide. With increasing resprout age post-severe-disturbance, new stems grow rapidly at their early age, and decrease in their growth with gradually decreasing water status thereafter. However, there is little knowledge about how stem hydraulic strategies and anatomical traits vary post-disturbance. In this study, the stem water potential (Ψstem), maximum stem hydraulic conductivity (Kstem-max), water potential at 50% loss of hydraulic conductivity (Kstem  P50) and anatomical traits of Caragana korshinkii resprouts were measured during a 1- to 13-year post-disturbance period. We found that the Kstem-max decreased with resprout age from 1-year-old resprouts (84.2 mol m-1 s-1 MPa-1) to 13-year-old resprouts (54.2 mol m-1 s-1 MPa-1) as a result of decreases in the aperture fraction (Fap) and the sum of aperture area on per unit intervessel wall area (Aap). The Kstem  P50 of the resprouts decreased from 1-year-old resprouts (-1.8 MPa) to 13-year-old resprouts (-2.9 MPa) as a result of increases in vessel implosion resistance (t/b)2, wood density (WD), vessel grouping index (GI) and decreases in Fap and Aap. These shifts in hydraulic structure and function resulted in an age-based divergence in hydraulic strategies i.e., a change from an acquisitive strategy to a conservative strategy, with increasing resprout age post-disturbance.

Spermine: Its Emerging Role in Regulating Drought Stress Responses in Plants.

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.