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.

Soybean-mediated suppression of BjaI/BjaR1 quorum sensing in Bradyrhizobium diazoefficiens impacts symbiotic nitrogen fixation.

The acyl-homoserine lactones (AHLs)-mediated LuxI/LuxR quorum sensing (QS) system orchestrates diverse bacterial behaviors in response to changes in population density. The role of the BjaI/BjaR1 QS system in Bradyrhizobium diazoefficiens USDA 110, which shares homology with LuxI/LuxR, remains elusive during symbiotic interaction with soybean. Here this genetic system in wild-type (WT) bacteria residing inside nodules exhibited significantly reduced activity compared to free-living cells, potentially attributed to soybean-mediated suppression. The deletion mutant strain ΔbjaR1 showed significantly enhanced nodulation induction and nitrogen fixation ability. Nevertheless, its ultimate symbiotic outcome (plant dry weight) in soybeans was compromised. Furthermore, comparative analysis of the transcriptome, proteome, and promoter activity revealed that the inactivation of BjaR1 systematically activated and inhibited genomic modules associated with nodulation and nitrogen metabolism. The former appeared to be linked to a significant decrease in the expression of NodD2, a key cell-density-dependent repressor of nodulation genes, while the latter conferred bacterial growth and nitrogen fixation insensitivity to environmental nitrogen. In addition, BjaR1 exerted a positive influence on the transcription of multiple genes involved in a so-called central intermediate metabolism within the nodule. In conclusion, our findings highlight the crucial role of the BjaI/BjaR1 QS circuit in positively regulating bacterial nitrogen metabolism and emphasize the significance of the soybean-mediated suppression of this genetic system for promoting efficient symbiotic nitrogen fixation by B. diazoefficiens.IMPORTANCEThe present study demonstrates, for the first time, that the BjaI/BjaR1 QS system of Bradyrhizobium diazoefficiens has a significant impact on its nodulation and nitrogen fixation capability in soybean by positively regulating NodD2 expression and bacterial nitrogen metabolism. Moreover, it provides novel insights into the importance of suppressing the activity of this QS circuit by the soybean host plant in establishing an efficient mutual relationship between the two symbiotic partners. This research expands our understanding of legumes' role in modulating symbiotic nitrogen fixation through rhizobial QS-mediated metabolic functioning, thereby deepening our comprehension of symbiotic coevolution theory. In addition, these findings may hold great promise for developing quorum quenching technology in agriculture.

Multiple independent losses of the biosynthetic pathway for two tropane alkaloids in the Solanaceae family.

Hyoscyamine and scopolamine (HS), two valuable tropane alkaloids of significant medicinal importance, are found in multiple distantly related lineages within the Solanaceae family. Here we sequence the genomes of three representative species that produce HS from these lineages, and one species that does not produce HS. Our analysis reveals a shared biosynthetic pathway responsible for HS production in the three HS-producing species. We observe a high level of gene collinearity related to HS synthesis across the family in both types of species. By introducing gain-of-function and loss-of-function mutations at key sites, we confirm the reduced/lost or re-activated functions of critical genes involved in HS synthesis in both types of species, respectively. These findings indicate independent and repeated losses of the HS biosynthesis pathway since its origin in the ancestral lineage. Our results hold promise for potential future applications in the artificial engineering of HS biosynthesis in Solanaceae crops.

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.

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.

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.

Effects of Heavy Metal Stress on Physiology, Hydraulics, and Anatomy of Three Desert Plants in the Jinchang Mining Area, China.

The physiological mechanisms and phytoremediation effects of three kinds of native quinoa in a desert mining area were studied. We used two different types of local soils (native soil and tailing soil) to analyze the changes in the heavy metal content, leaf physiology, photosynthetic parameters, stem hydraulics, and anatomical characteristics of potted quinoa. The results show that the chlorophyll content, photosynthetic rate, stomatal conductance, and transpiration rate of Kochia scoparia were decreased, but intercellular CO2 concentration (Ci) was increased under heavy metal stress, and the net photosynthetic rate (Pn) was decreased due to non-stomatal limitation. The gas exchange of Chenopodium glaucum and Atriplex centralasiatica showed a decrease in Pn, stomatal conductance (Gs), and transpiration rate (E) due to stomatal limitation. The three species showed a similar change in heavy metal content; they all showed elevated hydraulic parameters, decreased vessel density, and significantly thickened vessel walls under heavy metal stress. Physiological indicators such as proline content and activity of superoxide dismutase (SOD) and peroxidase (POD) increased, but the content of malondialdehyde (MDA) and glutathione (GSH), as well as catalase (CAT) activity, decreased in these three plants. Therefore, it can be concluded that these three species of quinoa, possibly the most dominant 30 desert plants in the region, showed a good adaptability and accumulation capacity under the pressure of heavy metal stress, and these plants can be good candidates for tailings remediation in the Jinchang desert mining area.

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.

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.

Nitrogen deposition magnifies destabilizing effects of plant functional group loss.

Terrestrial ecosystems are under threat by the co-occurring biodiversity loss and nitrogen (N) deposition. Awareness is growing that the stabilizing effects of plant diversity on productivity depend on environmental context, but it remains unknown about how the loss of plant functional groups and N deposition interactively influence species richness and community stability. Here we carried out an eight-year experiment of plant functional groups removal and N addition experiment in subalpine meadow. We found that the removal of plant functional groups and N addition interactively affected averaged plant species richness and community stability. Without N addition, the absence of forbs, but not other functional groups, significantly decreased average species richness and community stability through decreasing species asynchrony (i.e., asynchronous dynamics among species under fluctuating conditions). Under N addition, the absence of forbs, grasses and legumes all led to significant declines in average species richness, causing a decrease in community stability by decreasing species asynchrony, among which the absence of forbs had the greatest negative effects on community stability. Moreover, N addition reinforced the destabilizing effects caused by the loss of functional groups. Our findings show that the diverse forbs maintain plant community stability through asynchronous dynamics among species, especially under N deposition scenario. Therefore, we suggest that conservation and restoration of plant communities and their stability would benefit from a functional-group specific strategy by considering the largely ignored forb species, while helps guide conservation management efforts to reduce temporal variability for ecosystem service in the face of uncertain species extinction and N deposition scenarios.

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-Mediated Tolerance to Selenium Toxicity in Wheat (Triticum aestivum L.) Depends on Endogenous Nitric Oxide Synthesis.

Excess selenium (Se) causes toxicity, and nitric oxide (NO)'s function in spermine (Spm)-induced tolerance to Se stress is unknown. Using wheat plants exposed to 1 mM sodium selenate-alone or in combination with either 1 mM Spm, 0.1 mM NO donor sodium nitroprusside (SNP) or 0.1 mM NO scavenger cPTIO-the potential beneficial effects of these compounds to palliate Se-induced stress were evaluated at physiological, biochemical and molecular levels. Se-treated plants accumulated Se in their roots (92%) and leaves (95%) more than control plants. Furthermore, Se diminished plant growth, photosynthetic traits and the relative water content and increased the levels of malondialdehyde, H2O2, osmolyte and endogenous NO. Exogenous Spm significantly decreased the levels of malondialdehyde by 28%, H2O2 by 37% and electrolyte leakage by 42%. Combined Spm/NO treatment reduced the Se content and triggered plant growth, photosynthetic traits, antioxidant enzymes and glyoxalase systems. Spm/NO also upregulated MTP1, MTPC3 and HSP70 and downregulated TaPCS1 and NRAMP1 (metal stress-related genes involved in selenium uptake, translocation and detoxification). However, the positive effects of Spm on Se-stressed plants were eliminated by the NO scavenger. Accordingly, data support the notion that Spm palliates selenium-induced oxidative stress since the induced NO elicits antioxidant defence upregulation but downregulates Se uptake and translocation. These findings pave the way for potential biotechnological approaches to supporting sustainable wheat crop production in selenium-contaminated areas.

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.

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.

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.