Epigenetic Modifications of Hormonal Signaling Pathways in Plant Drought Response and Tolerance for Sustainable Food Security.

Drought significantly challenges global food security, necessitating a comprehensive understanding of plant molecular responses for effective mitigation strategies. Epigenetic modifications, such as DNA methylation and histone modifications, are key in regulating genes and hormones essential for drought response. While microRNAs (miRNAs) primarily regulate gene expression post-transcriptionally, they can also interact with epigenetic pathways as potential effectors that influence chromatin remodeling. Although the role of miRNAs in epigenetic memory is still being explored, understanding their contribution to drought response requires examining these indirect effects on epigenetic modifications. A key aspect of this exploration is epigenetic memory in drought-adapted plants, offering insights into the transgenerational inheritance of adaptive traits. Understanding the mechanisms that govern the maintenance and erasure of these epigenetic imprints provides nuanced insights into how plants balance stability and flexibility in their epigenomes. A major focus is on the dynamic interaction between hormonal pathways-such as those for abscisic acid (ABA), ethylene, jasmonates, and salicylic acid (SA)-and epigenetic mechanisms. This interplay is crucial for fine-tuning gene expression during drought stress, leading to physiological and morphological adaptations that enhance plant drought resilience. This review also highlights the transformative potential of advanced technologies, such as bisulfite sequencing and CRISPR-Cas9, in providing comprehensive insights into plant responses to water deficit conditions. These technologies pave the way for developing drought-tolerant crops, which is vital for sustainable agriculture.

Molecular Mechanisms of CBL-CIPK Signaling Pathway in Plant Abiotic Stress Tolerance and Hormone Crosstalk.

Abiotic stressors, including drought, salt, cold, and heat, profoundly impact plant growth and development, forcing elaborate cellular responses for adaptation and resilience. Among the crucial orchestrators of these responses is the CBL-CIPK pathway, comprising calcineurin B-like proteins (CBLs) and CBL-interacting protein kinases (CIPKs). While CIPKs act as serine/threonine protein kinases, transmitting calcium signals, CBLs function as calcium sensors, influencing the plant's response to abiotic stress. This review explores the intricate interactions between the CBL-CIPK pathway and plant hormones such as ABA, auxin, ethylene, and jasmonic acid (JA). It highlights their role in fine-tuning stress responses for optimal survival and acclimatization. Building on previous studies that demonstrated the enhanced stress tolerance achieved by upregulating CBL and CIPK genes, we explore the regulatory mechanisms involving post-translational modifications and protein-protein interactions. Despite significant contributions from prior research, gaps persist in understanding the nuanced interplay between the CBL-CIPK system and plant hormone signaling under diverse abiotic stress conditions. In contrast to broader perspectives, our review focuses on the interaction of the pathway with crucial plant hormones and its implications for genetic engineering interventions to enhance crop stress resilience. This specialized perspective aims to contribute novel insights to advance our understanding of the potential of the CBL-CIPK pathway to mitigate crops' abiotic stress.

Sodium nitroprusside modulates oxidative and nitrosative processes in Lycopersicum esculentum L. under drought stress.

KEY MESSAGE: Sodium nitroprusside mediates drought stress responses in tomatoes by modulating nitrosative and oxidative pathways, highlighting the interplay between nitric oxide, hydrogen sulfide, and antioxidant systems for enhanced drought tolerance. While nitric oxide (NO), a signalling molecule, enhances plant tolerance to abiotic stresses, its precise contribution to improving tomato tolerance to drought stress (DS) through modulating oxide-nitrosative processes is not yet fully understood. We aimed to examine the interaction of NO and nitrosative signaling, revealing how sodium nitroprusside (SNP) could mitigate the effects of DS on tomatoes. DS-seedlings endured 12% polyethylene glycol (PEG) in a 10% nutrient solution (NS) for 2 days, then transitioned to half-strength NS for 10 days alongside control plants. DS reduced total plant dry weight, chlorophyll a and b, Fv/Fm, leaf water potential (ΨI), and relative water content, but improved hydrogen peroxide (H2O2), proline, and NO content. The SNP reduced the DS-induced H2O2 generation by reducing thiol (-SH) and the carbonyl (-CO) groups. SNP increased not only NO but also the activity of L-cysteine desulfhydrase (L-DES), leading to the generation of H2S. Decreases in S-nitrosoglutathione reductase (GSNOR) and NADPH oxidase (NOX) suggest a potential regulatory mechanism in which S-nitrosylation [formation of S-nitrosothiol (SNO)] may influence protein function and signaling pathways during DS. Moreover, SNP improved ascorbate (AsA) and glutathione (GSH) and reduced oxidized glutathione (GSSG) levels in tomato plants under drought. Furthermore, the interaction of NO and H2S, mediated by L-DES activity, may serve as a vital cross-talk mechanism impacting plant responses to DS. Understanding these signaling interactions is crucial for developing innovative drought-tolerance strategies in crops.

Microbial consortia-mediated arsenic bioremediation in agricultural soils: Current status, challenges, and solutions.

Arsenic poisoning in agricultural soil is caused by both natural and man-made processes, and it poses a major risk to crop production and human health. Soil quality, agricultural production, runoff, ingestion, leaching, and absorption by plants are all influenced by these processes. Microbial consortia have become a feasible bioremediation technique in response to the urgent need for appropriate remediation solutions. These diverse microbial populations collaborate to combat arsenic poisoning in soil by facilitating mechanisms including oxidation-reduction, methylation-demethylation, volatilization, immobilization, and arsenic mobilization. The current state, problems, and remedies for employing microbial consortia in arsenic bioremediation in agricultural soils are examined in this review. Among the elements affecting their success include diversity, activity, community organization, and environmental conditions. Also, we emphasize the sensitivity and accuracy limits of existing assessment techniques. While earlier reviews have addressed a variety of arsenic remediation options, this study stands out by concentrating on microbial consortia as a viable strategy for arsenic removal and presents performance evaluation and technical problems. This work gives vital insights for tackling the major issue of arsenic pollution in agricultural soils by explaining the potential methods and components involved in microbial consortium-mediated arsenic bioremediation.

Mitigating salt toxicity and overcoming phosphate deficiency alone and in combination in pepper (Capsicum annuum L.) plants through supplementation of hydrogen sulfide.

While it is widely recognized that hydrogen sulfide (H2S) promotes plant stress tolerance, the precise processes through which H2S modulates this process remains unclear. The processes by which H2S promotes phosphorus deficiency (PD) and salinity stress (SS) tolerance, simulated individually or together, were examined in this study. The adverse impacts on plant biomass, total chlorophyll and chlorophyll fluorescence were more pronounced with joint occurrence of PD and SS than with individual application. Malondialdehyde (MDA), hydrogen peroxide (H2O2), and electrolyte leakage (EL) levels in plant leaves were higher in plants exposed to joint stresses than in plants grown under an individual stress. When plants were exposed to a single stress as opposed to both stressors, sodium hydrosulfide (NaHS) treatment more efficiently decreased EL, MDA, and H2O2 concentrations. Superoxide dismutase, peroxidase, glutathione reductase and ascorbate peroxidase activities were increased by SS alone or in conjunction with PD, whereas catalase activity decreased significantly. The favorable impact of NaHS on all the evaluated attributes was reversed by supplementation with 0.2 mM hypotaurine (HT), a H2S scavenger. Overall, the unfavorable effects caused to NaHS-supplied plants by a single stress were less severe compared with those caused by the combined administration of both stressors.

Epigenetic and Hormonal Modulation in Plant-Plant Growth-Promoting Microorganism Symbiosis for Drought-Resilient Agriculture.

Plant growth-promoting microorganisms (PGPMs) have emerged as valuable allies for enhancing plant growth, health, and productivity across diverse environmental conditions. However, the complex molecular mechanisms governing plant-PGPM symbiosis under the climatic hazard of drought, which is critically challenging global food security, remain largely unknown. This comprehensive review explores the involved molecular interactions that underpin plant-PGPM partnerships during drought stress, thereby offering insights into hormonal regulation and epigenetic modulation. This review explores the challenges and prospects associated with optimizing and deploying PGPMs to promote sustainable agriculture in the face of drought stress. In summary, it offers strategic recommendations to propel research efforts and facilitate the practical implementation of PGPMs, thereby enhancing their efficacy in mitigating drought-detrimental effects in agricultural soils.

Synergistic mitigation of nickel toxicity in pepper (Capsicum annuum) by nitric oxide and thiourea via regulation of nitrogen metabolism and subcellular nickel distribution.

Nickel (Ni) contamination hinders plant growth and yield. Nitric oxide (NO) and thiourea (Thi) aid plant recovery from heavy metal damage, but their combined effects on pepper (Capsicum annuum ) plant tolerance to Ni stress need more study. Sodium nitroprusside (0.1mM, SNP) and 400mgL-1 Thi, alone and combined, were studied for their impact on pepper growth under Ni toxicity. Ni stress reduces chlorophyll, PSII efficiency and leaf water and sugar content. However, SNP and Thi alleviate these effects by increasing leaf water, proline and sugar content. It also increased the activities of superoxide dismutase, catalase, ascorbate peroxidase and peroxidase. Nickel stress lowered nitrogen assimilation enzymes (nitrate reductase, nitrite reductase, glutamine synthetase, glutamate synthase and glutamate dehydrogenase) and protein content, but increased nitrate, ammonium and amino acid content. SNP and Thi enhanced nitrogen assimilation, increased protein content and improved pepper plant growth and physiological functions during Ni stress. The combined treatment reduced Ni accumulation, increased Ni in leaf cell walls and potentially in root vacuoles, and decreased Ni concentration in cell organelles. It effectively mitigated Ni toxicity to vital organelles, surpassing the effects of SNP or Thi use alone. This study provides valuable insights for addressing heavy metal contamination in agricultural soils and offers potential strategies for sustainable and eco-friendly farming practices.

5-Aminolevulinic Acid Induces Chromium [Cr(VI)] Tolerance in Tomatoes by Alleviating Oxidative Damage and Protecting Photosystem II: A Mechanistic Approach.

Chromium [Cr(VI)] pollution is a major environmental risk, reducing crop yields. 5-Aminolevunic acid (5-ALA) considerably improves plant abiotic stress tolerance by inducing hydrogen peroxide (H2O2) and nitric oxide (NO) signalling. Our investigation aimed to uncover the mechanism of tomato tolerance to Cr(VI) toxicity through the foliar application of 5-ALA for three days, fifteen days before Cr treatment. Chromium alone decreased plant biomass and photosynthetic pigments, but increased oxidative stress markers, i.e., H2O2 and lipid peroxidation (as MDA equivalent). Electrolyte leakage (EL), NO, nitrate reductase (NR), phytochelatins (PCs), glutathione (GSH), and enzymatic and non-enzymatic antioxidants were also increased. Foliar application of 5-ALA before Cr treatment improved plant growth and photosynthetic pigments, diminished H2O2, MDA content, and EL, and resulted in additional enhancements of enzymatic and non-enzymatic antioxidants, NR activity, and NO synthesis. In Cr-treated tomato seedlings, 5-ALA enhanced GSH and PCs, which modulated Cr sequestration to make it nontoxic. 5-ALA-induced Cr tolerance was further enhanced by sodium nitroprusside (SNP), a NO donor. When sodium tungstate (ST), a NR inhibitor, was supplied together with 5-ALA to Cr-treated plants, it eliminated the beneficial effects of 5-ALA by decreasing NR activity and NO synthesis, while the addition of SNP inverted the adverse effects of ST. We conclude that the mechanism by which 5-ALA induced Cr tolerance in tomato seedlings is mediated by NR-generated NO. Thus, NR and NO are twin players, reducing Cr toxicity in tomato plants via antioxidant signalling cascades.

Salicylic acid interacts with other plant growth regulators and signal molecules in response to stressful environments in plants.

Salicylic acid (SA) is one of the potential plant growth regulators (PGRs) that regulate plant growth and development by triggering many physiological and metabolic processes. It is also known to be a crucial component of plant defense mechanisms against environmental stimuli. In stressed plants, it is documented that it can effectively modulate a myriad of metabolic processes including strengthening of oxidative defense system by directly or indirectly limiting the buildup of reactive nitrogen and oxygen radicals. Although it is well recognized that it performs a crucial role in plant tolerance to various stresses, it is not fully elucidated that whether low or high concentrations of this PGR is effective to achieve optimal growth of plants under stressful environments. It is also not fully understood that to what extent and in what manner it cross-talks with other potential growth regulators and signalling molecules within the plant body. Thus, this critical review discusses how far SA mediates crosstalk with other key PGRs and molecular components of signalling pathways mechanisms, particularly in plants exposed to environmental cues. Moreover, the function of SA exogenously applied in regulation of growth and development as well as reinforcement of oxidative defense system of plants under abiotic stresses is explicitly elucidated.

Nitric oxide and hydrogen sulfide work together to improve tolerance to salinity stress in wheat plants by upraising the AsA-GSH cycle.

The participation of nitric oxide (NO) in wheat plant tolerance to salinity stress (SS) brought about by hydrogen sulphide (H2S) via modifying the ascorbate-glutathione (AsA-GSH) cycle was studied. The SS-plants received either 0.2 mM sodium hydrosulfide (NaHS; H2S donor), or NaHS plus 0.1 mM sodium nitroprusside (SNP; a NO donor) through the nutrient solution. Salinity stress decreased plant growth, leaf water status, leaf K+, and glyoxalase II (gly II), while it elevated proline content, leaf Na+ content, oxidative stress, methylglyoxal (MG), glyoxalase I (gly I), the superoxide dismutase, catalase and peroxidase activities, contents of endogenous NO and H2S. The NaHS supplementation elevated plant development, decreased leaf Na+ content and oxidative stress, and altered leaf water status, leaf K+ and involved enzymes in AsA-GSH, H2S and NO levels. The SNP supplementation boosted the positive impact of NaHS on these traits in the SS-plants. Moreover, 0.1 mM cPTIO, scavenger of NO, countered the beneficial effect of NaHS by lowering NO levels. SNP and NaHS + cPTIO together restored the beneficial effects of NaHS by increasing NO content, implying that NO may have been a major factor in SS tolerance in wheat plants induced by H2S via activating enzymes connected to the AsA-GSH cycle.

The involvement of hydrogen sulphide in melatonin-induced tolerance to arsenic toxicity in pepper (Capsicum annuum L.) plants by regulating sequestration and subcellular distribution of arsenic, and antioxidant defense system.

Melatonin (MT) and hydrogen sulphide (H2S) are recognised as vital biomolecules actively taking part in plant defence systems as free radical scavengers and antioxidants against a myriad of biotic and abiotic stressors. However, it has been yet unknown in plants subjected to arsenic (As) toxicity whether or not H2S interacts with MT to regulate endogenous antioxidant defence system. Prior to beginning As stress (As-S) treatments, MT (0.10 mM) was applied externally to plants daily for three days. AsS was then started for two weeks with As(V) (0.1 mM as Na2HAsO4·7H2O). The treatment of As reduced plant biomass (24.4%) and chlorophyll a (51.7%), chlorophyll b (25.9%), while it increased subcellular As in roots and leaves, levels of glutathione (GSH), hydrogen peroxide (H2O2), malondialdehyde (MDA), methylglyoxal (MG), H2S and phytochelatins (PCs) in pepper plants. In As-stressed pepper plants, the application of MT increased plant biomass (16.3%), chlorophyll a (52.7%), chlorophyll b (28.2%), antioxidant enzymes' activities, and H2S accumulation, while it lowered the concentrations of MDA and H2O2. In As-treated plants, GSH and phytochelatins (PCs) were increased by MT by regulating As sequestration to make it harmless. The addition of MT increased As accumulation in the vacuoles of roots and caused the soluble fraction of As in vacuoles to become less toxic to vital organelles. MT-induced tolerance to As stress was further enhanced using NaHS, a source of H2S. Hypotaurine (0.1 mM HT), a H2S scavenger, was applied to the control and As-stressed plants together with MT and MT + NaHS to determine whether H2S was implicated in MT-induced increased As-S tolerance. By reducing H2S generation in pepper plants, HT counteracted the beneficial effects of MT, whereas the addition of NaHS to MT + HT restored the negative effects of HT, proving that H2S is necessary for the pepper plants As-stress tolerance caused by MT.

The participation of nitric oxide in hydrogen sulphide-mediated chromium tolerance in pepper (Capsicum annuum L) plants by modulating subcellular distribution of chromium and the ascorbate-glutathione cycle.

The promising response of chromium-stressed (Cr(VI)-S) plants to hydrogen sulphide (H2S) has been observed, but the participation of nitric oxide (NO) synthesis in H2S-induced Cr(VI)-S tolerance in plants remains to be elucidated. It was aimed to assess the participation of NO in H2S-mediated Cr(VI)-S tolerance by modulating subcellular distribution of Cr and the ascorbate-glutathione (AsA-GSH) cycle in the pepper seedlings. Two weeks following germination, plants were exposed to control (no Cr) or Cr(VI)-S (50 µM K2Cr2O7) for further two weeks. The Cr(VI)-S-plants grown in nutrient solution were supplied with 200 µM sodium hydrosulphide (NaHS, donor of H2S), or NaHS plus 100 µM sodium nitroprusside (SNP, a donor of NO). Chromium stress suppressed plant growth and leaf water status, while elevated proline content, oxidative stress, and the activities of AsA-GSH related enzymes, as well as endogenous H2S and NO contents. The supplementation of NaHS increased Cr accumulation at root cell walls and vacuoles of leaves as soluble fraction to reduce its toxicity. Furthermore it limited oxidative stress, improved plant growth, modulated leaf water status, and the AsA-GSH cycle-associated enzymes' activities, as well as it further improved H2S and NO contents. The positive effect of NaHS was found to be augmented on those parameters in the CrS-plants by the SNP supplementation. However, 0.1 mM cPTIO, the scavenger of NO, inverted the prominent effect of NaHS by decreasing NO content. The supplementation of SNP along with NaHS + cPTIO reinstalled the positive effect of NaHS by restoring NO content, which suggested that NO might have a potential role in H2S-induced tolerance to Cr(VI)-S in pepper plants by stepping up the AsA-GSH cycle.

Combined application of asparagine and thiourea improves tolerance to lead stress in wheat by modulating AsA-GSH cycle, lead detoxification and nitrogen metabolism.

Lead (Pb), like other heavy metals, is not essentially required for optimal plant growth; however, plants uptake it from the soil, which poses an adverse effect on growth and yield. Asparagine (Asp) and thiourea (Thi) are known to assuage the negative impacts of heavy metal pollution on plant growth; however, combined application of Asp and Thi has rarely been tested to discern if it could improve wheat yield under Pb stress. Thus, this experimentation tested the role of individual and combined applications of Asp (40 mM) and Thi (400 mg/L) in improving wheat growth under lead (Pb as PbCl2, 0.1 mM) stress. Lead stress significantly reduced plant growth, chlorophyll contents and photosystem system II (PSII) efficiency, whereas it increased Pb accumulation in the leaves and roots, leaf proline contents, phytochelatins, and oxidative stress related attributes. The sole or combined application of Asp and Thi increased the vital antioxidant biomolecules/enzymes, including reduced glutathione (GSH), ascorbic acid (AsA), ascorbate peroxsidase (APX), catalase (CAT), superoxide dismutase (SOD), glutathione S-transferase (GST), dehydroascorbate reductase (DHAR), and glutathione reductase (GR). Furthermore, the sole or the combined application of Asp and Thi modulated nitrogen metabolism by stimulating the activities of nitrate and nitrite reductase, glutamate synthase (GOGAT) and glutamine synthetase (GS). Asp and Thi together led to improve plant growth and vital physiological processes, but lowered down Pb accumulation compared to those by their sole application. The results suggest that Asp and Thi synergistically can improve wheat growth under Pb-toxicity.

Methyl Jasmonate and Sodium Nitroprusside Jointly Alleviate Cadmium Toxicity in Wheat (Triticum aestivum L.) Plants by Modifying Nitrogen Metabolism, Cadmium Detoxification, and AsA-GSH Cycle.

The principal intent of the investigation was to examine the influence of joint application of methyl jasmonate (MeJA, 10 µM) and a nitric oxide-donor sodium nitroprusside (SNP, 100 µM) to wheat plants grown under cadmium (Cd as CdCl2, 100 µM) stress. Cd stress suppressed plant growth, chlorophylls (Chl), and PSII maximum efficiency (F v /F m ), but it elevated leaf and root Cd, and contents of leaf proline, phytochelatins, malondialdehyde, and hydrogen peroxide, as well as the activity of lipoxygenase. MeJA and SNP applied jointly or singly improved the concentrations of key antioxidant biomolecules, e.g., reduced glutathione and ascorbic acid and the activities of the key oxidative defense system enzymes such as catalase, superoxide dismutase, dehydroascorbate reductase, glutathione S-transferase, and glutathione reductase. Exogenously applied MeJA and SNP jointly or singly also improved nitrogen metabolism by activating the activities of glutamine synthetase, glutamate synthase, and nitrate and nitrite reductases. Compared with individual application of MeJA or SNP, the combined application of both showed better effect in terms of improving plant growth and key metabolic processes and reducing tissue Cd content, suggesting a putative interactive role of both compounds in alleviating Cd toxicity in wheat plants. MAIN FINDINGS: The main findings are that exogenous application of methyl jasmonate and nitric oxide-donor sodium nitroprusside alleviated the cadmium (Cd)-induced adverse effects on growth of wheat plants grown under Cd by modulating key physiological processes and up-regulating enzymatic antioxidants and the ascorbic acid-glutathione cycle-related enzymes.

Melatonin-mediated nitric oxide improves tolerance to cadmium toxicity by reducing oxidative stress in wheat plants.

Two independent trials were conducted to examine the involvement of nitric oxide (NO) in MT-mediated tolerance to Cd toxicity in wheat plants. Cadmium toxicity considerably led to a decrease in plant growth, total chlorophyll, PSII maximum efficiency (Fv/Fm), leaf water potential, potassium (K+) and calcium (Ca2+). Simultaneously, it caused an increase in levels of leaf malondialdehyde (MDA), hydrogen peroxide (H2O2), electron leakage (EL), cadmium (Cd) and nitric oxide (NO) compared to those in control plants. Both MT (50 or 100 µM) treatments increased plant growth attributes and leaf Ca2+ and K+ in the leaves, but reduced MDA, H2O2 as well as leaf Cd content compared to those in Cd-stressed plants. A further experiment was designed to understand whether or not NO played a role in alleviation of Cd stress in wheat seedlings by melotonin using a scavenger of NO, 2-(4-carboxyphenyl)-4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide potassium salt (cPTIO) combined with the MT treatments. Melatonin-enhanced tolerance to Cd stress was completely reversed by the supply of cPTIO, which in turn considerably reduced the levels of endogenous NO. The results evidently showed that MT enhanced tolerance of wheat seedlings to Cd toxicity by triggering the endogenous NO. This was reinforced by the rise in the levels of MDA and H2O2, and decrease in the activities of superoxide dismutase (SOD; EC 1.15.1.1), catalase (CAT; EC. 1.11.1.6) and peroxidase (POD; EC. 1.11.1.7). The cPTO supply along with that of MT caused growth inhibition and a considerable increase in leaf Cd. So, both MT and NO together enhanced Cd tolerance in wheat.