Review Article | Volume 12, Issue 2, March, 2024

Plant growth regulator-mediated response under abiotic stress: A review

Shahreen Khan Ravinder Singh Harpreet Kaur Ajay Kumar Amit Vashishth Moyad Shahwan Hardeep Singh Tuli   

Open Access   

Published:  Feb 20, 2024

DOI: 10.7324/JABB.2024.157299
Abstract

Plant growth regulators (PGRs) are essential for regulating how plants respond to abiotic stress by activating their proliferation and development pathways. Abiotic stressors that plants encounter during their development cycle are attempted to be managed by growth regulators. Growth hormones are chemical messengers that control a plant’s regular growth and responsiveness to external stimuli. They control tissue development and differentiation, which controls how quickly plants develop. PGRs are necessary for the plant’s reaction to abiotic stress. In addition, hormones in plants allow them to recognize adverse environmental circumstances. The plants’ ability to biosynthesize phytohormones allows them to adapt to their environment. Abscisic acid aids plants in dealing with salt and drought stress, whereas ethylene aids plants in salinity, overwatering, cold, and drought. Plants may recover from mechanical damage and drought stress with the help of jasmonic acid. Research has also offered some tips for maximizing the ability of growth regulators to strengthen crops’ tolerance to abiotic stressors.


Keyword:     Abiotic stress Abscisic acid Ethylene Hormone Jasmonic acid Salinity


Citation:

Khan S, Singh R, Kaur H, Kumar A, Vashishth A, Shahwan M, Tuli HS. Plant growth regulator-mediated response under abiotic stress: A review. J App Biol Biotech. 2024;12(2):13-21. http://doi.org/10.7324/JABB.2024.157299

Copyright: Author(s). This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial-ShareAlike license.

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ARTICLE HIGHLIGHTS

  • PGRs play a vital role in helping plants to tolerate abiotic stress.

  • Research on the use of PGRs to improve crop tolerance to abiotic stress is promising.

  • PGRs can help plants to tolerate abiotic stress by altering their physiology and morphology.

  • PGRs have the potential to be used as sustainable tools to mitigate the effects of climate change on agriculture.


1. INTRODUCTION

Global agricultural output suffers enormous losses due to abiotic stress, which comprises elements such as salt, drought, and severe temperatures [1]. The degree to which plants react to abiotic stressors varies. The stress factors already present have been amplified by several human activities. They are unable to realize their full genetic potential due to all these stressors, which also reduce agricultural output. Abiotic stressors are becoming the main danger to agriculture productivity globally due to increased worries about climate uncertainty [2]. Complex abiotic plant stress conditions arise due to the intricate interplay of stress impacts and responses across various levels of plant structure. To enhance their resilience against stressors, plants possess intrinsic biochemical mechanisms that function at the cellular level [3]. Heavy metals adversely affect plant’s processes such as photosynthesis, respiration, division and elongation of cells, and mineral nutrition. [4,5]. Reactive oxygen species (ROS) is produced as a result of their harmful effects. The structure of cellular processes is harmed by the oxidative stress caused by heavy metal stress [6]. All these toxic effects reduce the growth and yields of plants, in severe cases, may also result in the plant death [4,7]. To maximize crop growth and production under stressful circumstances, knowledge of the processes connected to these mechanisms is essential. The regulating activities of plant growth regulators (PGR) are one of the significant and frequently studied elements of abiotic stress tolerance [8]. PGR is chemicals that have a significant impact on how plants differentiate and grow. They serve as chemical messengers for intracellular communication in addition. Auxins and gibberellins (GAS) primarily promote cell lengthening; cytokinins, which promote cell division, and fruit ripening, among other plant processes. Abscisic acid (ABA) promotes plant senescence and the disintegration of plant parts while also assisting in the preservation of plant water relations [9]. The steroid hormones polyamines and brassinosteroids, in addition to these five naturally occurring PGR types, are also known to have impacts on plant growth regulation and are the focus of extensive investigation [10]. Each of these natural PGRs can affect nearly all aspects of plant growth and development to increase stress resistance, despite being unique in terms of both chemical properties and the way they typically exhibit typical responses [11]. It has been demonstrated that increasing endogenous PGR levels under stressful conditions affect the growth of stress tolerance by aiding plants in several ways. Due to improvements achieved in the exact analysis of PGR employing reliable and effective physiochemical techniques as well as improvements made in molecular and genetic approaches, PGR research has drastically changed in connection to plant adaptation to abiotic challenges. A broad variety of compounds have been synthesized with clearly defined growth regulatory roles, in addition to the naturally occurring PGR. Numerous of them have been demonstrated to have a variety of applications for boosting plant growth, production, and output quality. In addition, significant work has been done in showing how such PGR might potentially improve agricultural plants’ abiotic stress responses. The numerous physiological and pharmacological components of PGR in connection to its participation in abiotic stress are discussed in this review.


2. METHODOLOGY

2.1. Literature Search

Articles from PubMed, Google Scholar, and Scopus were used to compile the data. Plant growth, hormone, stress, biotic stress, abiotic stress, ABA, GAS, jasmonates (JAs), and heavy metal were the search terms utilized. There were many different kinds of pertinent books, summaries, and articles. The electronic databases were searched, and 400 articles were found. After 2000, papers that appeared in credible scientific publications were included. After duplicate entries were eliminated, there were 250 articles left. The authors next checked the titles and abstracts of the remaining papers for data relevance. One hundred and fifty articles were created as a result of this approach. A total of 109 studies were chosen for the investigation after applying the inclusion (including specifically matched articles from our study related to PGR and stress) and exclusion criteria (excluding duplicate articles, personal opinions, book chapters, conference abstracts, full copies not available, and low-quality paper).


3. ABA MECHANISMS AND FUNCTION

Early in the 1960s, researchers established that the phytohormone ABA released seeds from dormancy [12]. Later, research revealed that it may have a part in plant development and plant stress adaptation [13]. ABA is produced in plastids through the methylerythritol phosphate route from carotenoids, a derivative of isopentenyl diphosphate [14-17]. ABA, often referred to as the “stress hormone,” plays a crucial role in regulating multiple processes that enhance a plant’s ability to withstand stress. Stressful conditions are known to impact about 10% of the expression of genes involved in protein-coding [18]. This hormone controls many crucial plant processes, such as seed dormancy, nutrition storage in growing seeds, tolerance to desiccation, and the cessation of embryonic development during seed maturation [18-20]. In addition, it is crucial for the production of various osmolytes and proteins. Depending on the type of disease, ABA may positively or negatively affect the defense response [21-23]. The potential of plants to adapt to drought circumstances is made possible by ABA’s involvement in several plant growth processes and its stress-related responses. In times of drought stress, ABA induces the closure of stomata, which, in turn, reduces water loss by lowering the transpiration rate. Moreover, ABA fosters the elongation of root cells and gradually enhances hydraulic conductivity, facilitating plants in recovering from water scarcity [24]. For instance, ABA affects how seeds develop and spread, how water is regulated in plant bodies through the opening and closing of stomata, and how plants react to environmental stresses [Figure 1]. The concentration of ABA in the leaves rises when there is a water deficit because it is biosynthesized in the roots and transported there by the xylem. The production and redistribution of ABA during the abiotic stress response led to stomatal closure and a decrease in transpiration rate, which restricts cell growth. ABA signals through phosphatases and kinases, which upregulate hormone-responsive transcription factors and fast responses to various stresses [25]. In addition, ABA regulates physiological responses in plants subjected to abiotic challenges by interacting with other phytohormones and mediating physiological responses through its signaling. It is crucial to remember that ABA mobilizes and activates a number of biochemical defenses, such as the biosynthesis of proline, antioxidants, ROS detoxifying enzymes, heat shock proteins, and unsaturated fatty acids, as well as the strengthening of cuticular waxes, which aids plants in partially reversing the negative effects of abiotic stress. Reducing stress causes the ABA level to return to pre-stressed levels and reverses the stress-induced rise [26]. Increased levels of ABA enable plants to reduce water loss through the closure of stomata, while simultaneously enhancing the plant’s water status due to improved root hydraulic conductivity [27]. ABA also helps the root and aboveground sections of the plant communicate through stomatal closure, metabolic adjustments, and gene expression [28].

Figure 1: The underlying molecular mechanisms of drought response in plants mediated by abscisic acid.



[Click here to view]

Since this regulatory mechanism is more strongly influenced by soil moisture levels rather than the water content of leaves, it is likely that ABA functions as a chemical signal produced by stressed roots [29]. The responsiveness of stomata to ABA is impacted by multiple factors, such as leaf age, environmental conditions (including the plant’s nutritional status), the ionic composition of the xylem sap, and the water content of the leaves. These variables influence the sensitivity of stomata to ABA and can vary among different plant species and cultivars. The differences in ABA supply to the active area of guard cells may explain the variations in ABA-mediated stomatal responses [30]. Under stressful conditions, the volume of guard cells actively responds to signals to minimize carbon dioxide (CO2) efflux for photosynthesis and reduce transpirational water loss [31]. The increase in ABA within guard cells prompts stomatal closure, leading to a reduction in transpirational water loss. In response to stress, gene expression related to stress protection produces essential proteins for signal transduction mechanisms within cells. The regulation of stress-inducible gene expression occurs through the interaction of transcription factors with specific cis-regulatory regions [32].


4. GAS MECHANISMS AND FUNCTION

One of the most well-known and earliest subclasses of regulatory hormones, the GAs are involved in a variety of plant developmental processes, including seed germination, inter-nodal elongation, forced blooming, and fruit formation [33]. Recent investigations utilized modern biochemical and genetic methods to investigate the genetic foundation, genes encoding the manufacture of GAs, and deactivation enzymes [34]. The GAs group has more than 250 members, although just a few of them play significant physiological roles in plant development [35]. The control of the GAs at the cellular level is intricate. For instance, enzymes from tiny multi-genic families that each play a distinct expression pattern control the phases of the GAs metabolic pathway. However, GA biosynthesis is highly correlated with two gene families that encode GAs: GA20-oxidases (GA20ox) and can accelerate the last steps in the creation of bioactive GAs. Nuclear-localized GID1 is the receptor protein for GA, and binding of GA to the receptor protein facilitates interactions with the repressor protein DELLA [36]. The DELLA protein’s protein-protein interaction domain interacts with the F-Box protein, which, then, calls for the SLY1 complex to be ubiquitinated by the SCF-E3 ligase complex [37]. The DELLA repressor protein is ubiquitinated by the 26S proteasome and is then degraded, helping to bind transcription factors to the GA-inducible genes’ promoter region [38]. Therefore, GA mediates signaling by breaking down repressor proteins, which cause a variety of biochemical reactions [39]. Thus, GA generated in plant tissues, whether from endogenous production or exogenous application, lessens the negative effects of stress, and aids in developing plants’ stress tolerance [40]. For instance, the rice seed yield and tolerance to drought stress were improved when endogenous GA levels were reduced by GA2ox6 ectopic expression under water deficiency conditions. GA’s role in redox equilibrium, plants can adapt to less-than-ideal growth environments. GRAS transcription factors are essential in plant growth and signaling [41]. Liu et al. showed that abiotic stress treatments such as NaC and H2O2 increased the GRAS40 expression in tomatoes [42]. During the vegetative and reproductive stages of tomato development, SlGRAS40 is interrelated with auxin/GA pathways. Salt and drought resistance were also developed through transgenic SlGRAS40 plant expression [43].


5. JAs MECHANISMS AND FUNCTIONS

Methyl jasmonate, jasmonic acid (JA), and jasmonoyl isoleucine are significant members of the family of JAs, which are naturally occurring plant oxylipins [44]. In addition, JAs help plants defend themselves from disease and insect damage. According to recent studies, JAs can alter how plants develop and respond to various abiotic challenges, including those caused by heavy metals, heat, light, drought, salinity, and flooding [45]. JAs generally increase gene expression and control some physiological and biochemical responses to protect plants from the harmful impacts of environmental stressors.

The JAs, which include JA and its derivatives, are hormones produced by plants that control a number of developmental processes, such as the growth of roots, the development of stamens, flowering, and the aging of leaves. In addition, they increase resistance to abiotic stressors such as ozone, ultraviolet rays, hot temperatures, and cold. JAs are well-known stress hormones generated from lipids that regulate how plants respond to biotic and abiotic stresses, including pathogen infections and herbivore attacks [46]. Pathogen infections and insect bites are the two main types of plant biotic stressors. They both seriously impair crop development and production, affecting the economy and human health [47]. JAs can increase antioxidant capacity and the production of various osmoprotectants, such as proline and glycine betaine, helping maintain plant cells’ integrity under various biotic conditions [48]. The photoassimilates can also be remobilized to the vegetative sinks by these growth regulators, enhancing plant growth under adverse conditions. Plants can use direct defense systems, such as creating harmful substances, phytoalexins, and proteinase inhibitors to combat infections in herbivores [49].

JAs’ physiological and biochemical effects on Vicia faba and Artemisia absinthium (leaf senescence) have been documented for the 1st time (growth inhibition) [50,51]. This group of growth regulators affects plant species in a variety of ways, from promoting physiological processes to inhibiting them. Smaller petioles were produced by plants treated with JAs than control plants [52]. Barley, plants treated with JA (2.5 106 and 2.5 104 147 M) had slower seedling development. The growth of the seedlings was unaffected by the lower concentration, though [53]. Jasmonate concentrations above 50 mM accelerate the aging process in leaves by causing chlorophyll damage and the breakdown of chloroplast proteins [54].


6. METHYL JASMONATE BIOSYNTHESIS AND ACCUMULATION

JAs are cyclopentanones that are formed from fatty acids and are members of the family of oxylipins, which are produced by plants through the oxidative metabolism of polyunsaturated fatty acids. JAs are present in almost all higher plants, but their concentrations are higher in flowering and reproductive tissues and significantly lower in mature leaves and roots [55].

The octadecanoid pathway in plants is used to synthesize that JAs Linolenic acid synthesis in plants is facilitated by phospholipase D and deficient anther dehiscence in the lipids of the chloroplast and anther plastid membranes, respectively. JA, which is created from linolenic acid as a precursor, is, then, oxygenated by the enzymes lipoxygenase, allene oxide synthase, and allene oxide cyclase to create 12-oxo-phytodienoic acid (AOC). After that, OPDA is converted into JA and, then, catabolized by JA carboxyl methyltransferase [56].


7. METHYL JASMONATE (EXOGENOUS) IN REDUCING ABIOTIC STRESSORS IN PLANTS

7.1. Salt Stress

One of the biggest issues with agriculture globally is salinity, especially in arid and semiarid areas. Heavy salt stress affects plants in three different ways. First, the osmotic action of salt in the soil decreases water retention and nutrient imbalances, such as competition between Na+ and K+, interfering with the uptake of critical minerals [57-59].

7.2. Drought Stress

A key environmental stressor that can trigger anxiety is drought stress. ROS affects several organelles that control plant growth and development, lower plant output, and speed up lipid oxidation, nucleic acid breakdown, and chlorophyll loss. These occurrences usually result in oxidative damage and hasten the senescence of plants [60]. Plants can use various defense strategies in response to drought stress by changing their cellular metabolism and increasing the activity of oxygen-scavenging enzymes.

This means that MeJA effectively improves plants’ capacity to endure drought by altering a variety of biochemical and physiological traits, such as increasing the activity of oxidative enzymes and organic osmoprotectants [61] [Figure 2].

Figure 2: A layout of the different potentials of jasmonic acid and its derivatives.



[Click here to view]

7.3. Micronutrients Toxicity

Although boron is a crucial mineral for the healthy growth of higher plants, there is a very little window between its lack and toxicity [62]. High levels of boron in the soil can increase the production of ROS, cause lipid peroxidation and H2O2 to build up in plant cells and tissues, and reduce plant development and crop output [63,64].

7.4. Chilling Stress

Chilling stress is a post-harvest concern that has positive economic implications. Physiological abnormalities brought on by chilling damage symptoms might interfere with normal cell metabolism, break down unsaturated fatty acids, and induce peroxidation in plant tissues with membrane lipidosis. MeJA has successfully treat tomatoes chilling damage [65], and loquat [66]. It causes the activation of some defensive substances (such as heat shock proteins and phenolic compounds) to shield these crops’ cell membranes against chilling damage. By modulating arginine metabolism and increasing the production of heat-shock proteins in tomato, the exogenous administration of MeJA also reduces the effects of chilling injury [67].

7.5. Biochemical Parameters of Plants Modulated by Heavy Metals Stress

Oxidative stress is brought on by heavy metal concentrations above a certain threshold, which harms plant tissues and changes their morphophysiological characteristics [68,69]. Copper and cadmium reduced cell extension, diminished potassium leaf uptake, and inhibited soil-borne root and shoot growth. They also prevented photosynthesis in Phaseolus coccineus L. due to its extraordinary capacity to respond to environmental metal toxicity through a system involving signaling molecules that might strengthen stress reaction responses [70]. The best signaling chemicals have been identified as Jas [71]. MeJA administered exogenously so significantly lessens plant heavy metal damage by increasing enzyme activity and secondary metabolites. Heavy metal toxicity leads to the generation of ROS in plants such as non-radical molecules (singlet oxygen [O2], and hydrogen peroxide [H2O2]) and free radicals (hydroxyl radical, OH; superoxide anion, O2•-; alkoxy radical, RO; and hydroperoxyl radical, HO2) [3] [Table 1]. Production of ROS is a part of normal metabolic processes occurring in cellular organelles such as peroxisomes, chloroplasts, and mitochondria, but their overaccumulation under heavy metal toxicity leads to damage to cellular molecules such as carbohydrates, lipids, deoxyribonucleic acid, and proteins [72,73]. Cu and Fe undergo auto-oxidation, resulting in ROS generation mediated by a fenton-type reaction [74]. In a study by Georgiadou et al. (2018), Zn and Cu toxicity in Ocimum basilicum caused a reduction in overall antioxidant capacity and concentration of total proteins. Cu and Zn toxicity also caused oxidative or nitrosative stress, as observed in increased concentrations of H2O2 and NO in the Ocimum basilicum plants [75]. The effect of three heavy metals at 0, 500, 1000, and 2000/mg/kg-1 concentrations was explored on bamboo (Indocalamus latifolius) plants grown in a greenhouse [76]. It was revealed that the concentrations (1000 and 2000/mg/kg-1) induced a reduction in a number of indices related to photosynthesis, such as intercellular CO2 concentration (μmol CO2 mol−1), photosynthetic rate (μmol CO2 m−2/s−1), and conductance of H2O (mol H2O m−2/s−1). A decline was also observed in net assimilation and transpiration. A corresponding decline was also observed in morphological indices such as the reduced number of emerged plants and reduced shoot length [76]. Cd stress adversely affected the chlorophyll synthesis and modulated the photosystem I (PS I) and photosystem II (PS II) in tobacco leaves [5]. Downregulation of ROS scavenging and reduced expression of proteins linked with ferredoxin-dependent nitrogen metabolism caused blockage of photosynthetic electron transport. It severely inhibited both the photosystems (PS I and PS II). Downward trend in photosynthetic light-response parameters and photosynthetic gas exchange parameters with increasing concentrations of Cd (5, 20, 50, and 100 mg/kg) in Sassafras tzumu seedlings indicates a significant negative impact of cadmium contamination on the photosynthetic efficiency and overall physiological health of these seedlings [4]. Young plants of Virola surinamensis were evaluated for responses to different Cd concentrations (30, 45, and 60/mg/l−1). Cd stress caused a reduction in stomatal conductance, leaf water potential, and transpiration. In addition, the reduced maximum photochemical efficiency and electron transport rate were accompanied by a decline in photosynthesis [77].

Table 1: Effect of heavy metals on plant growth metrics.

S. No.PlantHeavy metalMetal dosesParameter affectedReferences
1.Artemisia annuaAl0.01, 0.10, 1.00 and 10 mMShoot and root length; dry weight; and dry leaf yield[86]
2.Brassica junceaCu3–5 ppmShoot and root length[87]
4.Brassica juncea and Brassica napusCd10 and 30 mg kg−1Plant height, weight, and total leaf area[80]
5.Brassica napusCd10 µMRoot and shoot dry and fresh weight[88]
6.Cenchrus ciliarisCd30 and 60 mg l−1Shoot and root fresh weight[89]
7.Cicer arietinumCd150 µMShoot and root length and weight[90]
8.Cicer arietinumCd25, 75, 150 µg mL−1Root length; number of leaves and branches[91]
9.Cicer arietinumAs5 mg/kg−1Shoot and root dry weight[92]
10.Citrus aurantiumCr200 ppmShoot and root length[93]
12.Cynodon dactylonCd30 and 60 mg l−1Shoot and root fresh weight[89]
13.Eichhornia crassipes (Mart.)Pb100, 200, 400, 600, 800 and 1000 mg l−1Plant height, dry weight, shoot length, and root length[94]
20.Ipomoea aquaticaAs20 and 40 mg kg−1Plant height and biomass[95]
21.Lactuca sativaCd1, 10 and 50 µMPlant dry weight[96]
22.Lactuca sativaCr5, 10 and 20 mg l−1Root and leaf length; and fresh seedling weight[97]
23.Laguncularia racemosaCr0.05 and 0.50 mg l−1Stem length; root and stem dry weight; and total dry weight[98]
24.Lolium perenneAs100 mg/kg−1Plant dry weight[99]
25.Oryza sativa cultivars Indica, Ediget, X-Jigna, and FuratPb1.2 mMPlant height, tillers number/plant, panicles number/plant, and spikelets number/panicle[81]
28.Pisum sativumCd1 and 6 mMShoot fresh biomass; and shoot dry biomass[100]
29.Pteris creticaAs20 and 100 mg kg−1Shoot and root dry biomass[101]
30.Raphanus sativusPb2.5 mMPlant height; plant weight; and plant dry weight[102]
31.Sassafras tzumuCd5, 20, 50 and 100 mg/kg−1Plant height; root, branch and leaf biomass[4]
33.Sorghum bicolorCr2 and 4 ppmShoot and root length; fresh leaf weight; and leaf dry weight[103]
34.Spinacia oleraceaPb1.5, 3 and 15 mMPlant fresh and dry weight[104]
35.Spinacia oleraceaAs20 and 100 mg/kg−1Shoot and root dry biomass[101]
39.Vigna radiataCr120 µMShoot and root length; shoot and root fresh weight; shoot and root dry weight[105]
42.Zea maysCd10 and 20 mg l−1Shoot and root dry weight[106]
43.Zea maysCd400 and 800 μmol l−1Root length; plant height; plant weight; and plant dry weight[107,108]
44.Zea maysAs3, 10 and 30 mg kg−1Shoot and root dry weight[108]

7.5.1. Effect on plant growth parameters

Chen et al. [74] investigated the toxicity of copper in rice seedlings the length of the roots gradually decreased as the CuSO4 concentration rose from 20 to 50 M. The shoot length, however, was unaffected [78]. Bimetal combinations of heavy metals Cd, Zn, Cu, and Pb treatment were given to Brassica juncea plants [79]. Combinations of PbCd, CuZn, and CuCd caused the greatest inhibition of B. juncea seedlings biomass growth. The inhibitory effect was weakest in the seedlings treated with ZnCd combination. The metal combinations also caused necrotic spots on the plant leaves and also inhibited the leaf surface growth. B. juncea and B. napus plants had shown a reduction in fresh weight, plant height and total leaf area when grown under the stress of Cd metal (10 and 30 mg/kg) [80]. Pisum sativum plants grown in polluted and non-polluted soils of South Cairo and Giza provinces of Egypt were compared [7]. Analysis of soil of polluted farms revealed the presence of heavy metals such as Pb, Zn, As, Fe, V, Co, and Cd, the concentrations of which were higher than the tolerable limits. It was found that the fresh weight (1654 kg/acre) and dry weight (126 kg/acre) of the plants collected from the polluted farms reduced significantly as compared to the plants collected from the non-polluted farms (5223 kg/acre and 404 kg/acre, respectively). Fruit production also decreased by 85.2% in the plants grown in polluted soil [7]. Rice cultivars – Indica, Ediget, X-Jigna, and Furat were grown in soil supplied with 0.6 mM and 1.2 mM solution of Pb. The soil was treated with the Pb solution 1 month before the transplantation of rice seedlings. Agronomic traits of the cultivars, such as height of the plants, tillers number/plant, panicles number/plant, and spikelets number/panicle, reduced significantly under the effect of 1.2 mM concentration of Pb [81]. Cu stress’ effects on the morphology of Cannabis sativa plants have been investigated. Cu treatment to the plants decreased their dry weights and shoot and root lengths [82]. Three cereal crops, Zea mays L., Triticum aestivum L., and Sorghum bicolor L., were examined to determine how detrimental amounts of Cu affected them [83]. After 4 weeks of Cu treatment, it was discovered that all growth features, including the fresh weight of shoots and roots and the dried weight of roots, had decreased (75 mol/l−1). The length and intensity of the Cu treatment were positively correlated with the decline. All of the growth indices for Phragmites australis and Salix purpurea, including total fresh weight, shoot length, and number of roots, significantly decreased [84]. S. tzumu seedlings grown under Cd stress (5, 20, 50, and 100/mg/kg−1) showed a decrease in root, branch, and leaf biomass as well as reduced plant height [4]. Reduction in the number and diameter of xylem vessels was recorded in Aechmea blanchetiana plant due to Pb toxicity [85]. Xylem vessels are the main conducting elements in angiospermic plants and are responsible for the conduction of water and mineral elements from roots to all other plant parts. Reduction in their number and diameter can significantly lower the supply of water and minerals to plant parts, reducing their overall growth. Table 1 displays how different heavy metals affect the factors of plant growth.


8. CONCLUSIONS AND FUTURE PERSPECTIVES

Plant hormones play a crucial role in the growth and development of plants under abiotic stressors. The stress circumstances change the quantities of these compounds that aid in plant adaptation by affecting how well stomata function, plant water balance, and antioxidant status are maintained. In response to stress, endogenous PGR levels in plants either rise or fall, depending on the cultivar, the length of the stress, and the severity of the stress. Although stressed plants always exhibited a rise in ABA and a reduction in cytokinins, different abiotic stress stimuli have distinct impacts on GAS, auxin, and polyamine levels. Recent years have seen significant advancements in our knowledge of the mechanisms controlling the biosynthesis, metabolism, and signaling functions of naturally occurring PGR. In addition, PGR commonly modifies gene expression by initiating or inhibiting the degradation of transcriptional regulators. Progress in understanding the molecular underpinnings of hormonal physiology has resulted in the discovery of genes involved in manufacturing various PGR and genes that code for their receptors. In addition, knowledge of stress-related gene modifications has been essential in identifying the function of PGR in plants’ response to abiotic stressors. Efforts must be made to use their advantages as a different tool for managing abiotic stressors. Furthermore, it is widely known that polyamines and brassinosteroids have a role in regulatory plant development and stress tolerance. However, additional investigation is required to determine the mechanism behind their stress-protective functions, particularly in light of their interactions and interrelationships with other PGR and stress-responsive genes.


9. AUTHORS’ CONTRIBUTIONS

According to the guidelines and criteria established by the International Committee of Medical Journal Editors, all authors fulfilled the necessary qualifications to be considered as authors. They collectively agreed to submit the article to the current journal, provided final approval of the version to be published, participated in the conception and design of the study, collected and analyzed data, critically revised the article for important intellectual content, and accepted accountability for all aspects of the work.


10. FUNDING

There is no funding to report.


11. CONFLICTS OF INTEREST

The authors report no financial or any other conflicts of interest in this work.


12. ETHICAL APPROVALS

There are no human or animal subjects in this study’s experiments.


13. DATA AVAILABILITY

In this review article, all the data that were produced and examined are included in the study.


14. PUBLISHER’S NOTE

Regarding jurisdictional assertions made in published institutional affiliation, this journal maintains its impartiality.

REFERENCES

1.  Fahad S, Bajwa AA, Nazir U, Anjum SA, Farooq A, Zohaib A, et al. Crop production under drought and heat stress:Plant responses and management options. Front Plant Sci 2017;8:1147. [https://doi.org/10.3389/fpls.2017.01147]

2.  Teshome DT, Zharare GE, Naidoo S. The threat of the combined effect of biotic and abiotic stress factors in forestry under a changing climate. Front Plant Sci 2020;11:601009. [https://doi.org/10.3389/fpls.2020.601009]

3.  Hasanuzzaman M, Nahar K, Alam M, Roychowdhury R, Fujita, M. Physiological, biochemical, and molecular mechanisms of heat stress tolerance in plants. Int J Mol Sci 2013;14:9643-84. [https://doi.org/10.3390/ijms14059643]

4.  Zhao H, Guan J, Liang Q, Zhang X, Hu H, Zhang J. Effects of cadmium stress on growth and physiological characteristics of sassafras seedlings. Sci Rep 2021;11:9913. [https://doi.org/10.1038/s41598-021-89322-0]

5.  Zhang H, Xu Z, Guo K, Huo Y, He G, Sun H, et al. Toxic effects of heavy metal Cd and Zn on chlorophyll, carotenoid metabolism and photosynthetic function in tobacco leaves revealed by physiological and proteomics analysis. Ecotoxicol Environ Saf 2020;202:110856. [https://doi.org/10.1016/j.ecoenv.2020.110856]

6.  Rizvi A, Zaidi A, Ameen F, Ahmed B, AlKahtani MD, Khan MS. Heavy metal induced stress on wheat:Phytotoxicity and microbiological management. RSC Adv 2020;10:38379-403. [https://doi.org/10.1039/D0RA05610C]

7.  Galal TM, Hassan LM, Ahmed DA, Alamri SA, Alrumman SA, Eid EM. Heavy metals uptake by the global economic crop (Pisum sativum L.) grown in contaminated soils and its associated health risks. PLoS One 2021;16:e0252229. [https://doi.org/10.1371/journal.pone.0252229]

8.  El Sabagh A, Mbarki S, Hossain A, Iqbal MA, Islam MS, Raza A, et al. Potential role of plant growth regulators in administering crucial processes against abiotic stresses. Front Agron 2021;3:1-28. [https://doi.org/10.3389/fagro.2021.648694]

9.  Iqbal N, Khan NA, Ferrante A, Trivellini A, Francini A, Khan MI. Ethylene role in plant growth, development and senescence:Interaction with other phytohormones. Front Plant Sci 2017;8:475. [https://doi.org/10.3389/fpls.2017.00475]

10.  Bishop GJ, Koncz C. Brassinosteroids and plant steroid hormone signaling. Plant Cell 2002;14:S97-110. [https://doi.org/10.1105/tpc.001461]

11.  Backer R, Rokem JS, Ilangumaran G, Lamont J, Praslickova D, Ricci E, et al. Plant growth-promoting rhizobacteria:Context, mechanisms of action, and roadmap to commercialization of biostimulants for sustainable agriculture. Front Plant Sci 2018;9:1473. [https://doi.org/10.3389/fpls.2018.01473]

12.  Cornforth JW, Milborrow BV, Ryback G, Wareing PF. Chemistry and physiology of 'dormins'in sycamore:Identity of sycamore 'dormin'with abscisin II. Nature 1965;205:1269-70. [https://doi.org/10.1038/2051269b0]

13.  Guschina IA, Harwood JL, Smith M, Beckett RP. Abscisic acid modifies the changes in lipids brought about by water stress in the moss Atrichum androgynum. New Phytologist 2002;156:255-64. [https://doi.org/10.1046/j.1469-8137.2002.00517.x]

14.  Taylor IB, Burbidge A, Thompson AJ. Control of abscisic acid synthesis. J Exp Bot 2000;51:1563-74. [https://doi.org/10.1093/jexbot/51.350.1563]

15.  Takino J, Kozaki T, Sato Y, Liu C, Ozaki T, Minami A, et al. Unveiling biosynthesis of the phytohormone abscisic acid in fungi:Unprecedented mechanism of core scaffold formation catalyzed by an unusual sesquiterpene synthase. J Am Chem Soc 2018;140:12392-5. [https://doi.org/10.1021/jacs.8b08925]

16.  Lievens L, Pollier J, Goossens A, Beyaert R, Staal J. Abscisic acid as pathogen effector and immune regulator. Front Plant Sci 2017;8:587. [https://doi.org/10.3389/fpls.2017.00587]

17.  Rasool N. Plant hormones:Role in alleviating biotic stress. In:Plant Hormones-Recent Advances, New Perspectives and Applications. London:IntechOpen;2022. [https://doi.org/10.5772/intechopen.102689]

18.  Verma V, Ravindran P, Kumar PP. Plant hormone-mediated regulation of stress responses. BMC Plant Biol 2016;16:86. [https://doi.org/10.1186/s12870-016-0771-y]

19.  Rock CD, Quatrano RS. The role of hormones during seed development. In:Plant Hormones. Netherlands, Dordrecht:Springer;1995. 671-97. [https://doi.org/10.1007/978-94-011-0473-9_31]

20.  Miransari M, Smith DL. Plant hormones and seed germination. Environ Exp Bot 2014;99:110-21. [https://doi.org/10.1016/j.envexpbot.2013.11.005]

21.  Ghosh UK, Islam MN, Siddiqui MN, Khan MA. Understanding the roles of osmolytes for acclimatizing plants to changing environment:A review of potential mechanism. Plant Signal Behav 2021;16:1913306. [https://doi.org/10.1080/15592324.2021.1913306]

22.  Rabbani G, Choi I. Roles of osmolytes in protein folding and aggregation in cells and their biotechnological applications. Int J Biol Macromol 2018;109:483-91. [https://doi.org/10.1016/j.ijbiomac.2017.12.100]

23.  Sano N, Marion-Poll A. ABA metabolism and homeostasis in seed dormancy and germination. Int J Mol Sci 2021;22:5069. [https://doi.org/10.3390/ijms22105069]

24.  Daszkowska-Golec A. The role of abscisic acid in drought stress:How ABA helps plants to cope with drought stress. In:Drought Stress Tolerance in Plants. Vol. 2. Cham:Springer International Publishing;2016. 123-51. [https://doi.org/10.1007/978-3-319-32423-4_5]

25.  Kuromori T, Seo M, Shinozaki K. ABA transport and plant water stress responses. Trends Plant Sci 2018;23:513-22. [https://doi.org/10.1016/j.tplants.2018.04.001]

26.  Nakashima K, Yamaguchi-Shinozaki K. ABA signaling in stress-response and seed development. Plant Cell Rep 2013;32:959-70. [https://doi.org/10.1007/s00299-013-1418-1]

27.  Pirasteh-Anosheh H, Saed-Moucheshi A, Pakniyat H, Pessarakli M. Stomatal responses to drought stress. In:Water Stress and Crop Plants. Chichester, UK:John Wiley and Sons Ltd;2016. 24-40. [https://doi.org/10.1002/9781119054450.ch3]

28.  Hsu PK, Dubeaux G, Takahashi Y, Schroeder JI. Signaling mechanisms in abscisic acid-mediated stomatal closure. Plant J 2021;105:307-21. [https://doi.org/10.1111/tpj.15067]

29.  Sehler R, Li J, Reager J, Ye H. Investigating relationship between soil moisture and precipitation globally using remote sensing observations. J Contemp Water Res Educ 2019;168:106-18. [https://doi.org/10.1111/j.1936-704X.2019.03324.x]

30.  Pantin F, Monnet F, Jannaud D, Costa JM, Renaud J, Muller B, et al. The dual effect of abscisic acid on stomata. New Phytol 2013;197:65-72. [https://doi.org/10.1111/nph.12013]

31.  Xu Z, Jiang Y, Zhou G. Response and adaptation of photosynthesis, respiration, and antioxidant systems to elevated CO2 with environmental stress in plants. Front Plant Sci 2015;6:701. [https://doi.org/10.3389/fpls.2015.00701]

32.  Bharath P, Gahir S, Raghavendra AS. Abscisic acid-induced stomatal closure:An important component of plant defense against abiotic and biotic stress. Front Plant Sci 2021;12:615114. [https://doi.org/10.3389/fpls.2021.615114]

33.  El Sabagh A, Islam MS, Hossain A, Iqbal MA, Mubeen M, Waleed M, et al. Phytohormones as growth regulators during abiotic stress tolerance in plants. Front Agron 2022;4:1-16. [https://doi.org/10.3389/fagro.2022.765068]

34.  Hedden P. The current status of research on gibberellin biosynthesis. Plant Cell Physiol 2020;61:1832-49. [https://doi.org/10.1093/pcp/pcaa092]

35.  Wang Y, Zhao J, Lu W, Deng D. Gibberellin in plant height control:Old player, new story. Plant Cell Rep 2017;36:391-8. [https://doi.org/10.1007/s00299-017-2104-5]

36.  Salazar-Cerezo S, Martínez-Montiel N, García-Sánchez J, Pérez-y-Terrón R, Martínez-Contreras R.D. Gibberellin biosynthesis and metabolism:A convergent route for plants, fungi and bacteria. Microbiol Res 2018;208:85-98. [https://doi.org/10.1016/j.micres.2018.01.010]

37.  Yoshida H, Hirano K, Sato T, Mitsuda N, Nomoto M, Maeo K, et al. DELLA protein functions as a transcriptional activator through the DNA binding of the Indeterminate Domain family proteins. Proc Natl Acad Sci 2014;111:7861-6. [https://doi.org/10.1073/pnas.1321669111]

38.  Coll-Martínez B, Crosas B. How the 26S proteasome degrades ubiquitinated proteins in the cell. Biomolecules 2019;9:395. [https://doi.org/10.3390/biom9090395]

39.  Silverstone AL, Jung HS, Dill A, Kawaide H, Kamiya Y, Sun TP. Repressing a repressor:Gibberellin-induced rapid reduction of the RGA protein in Arabidopsis. Plant Cell 2001;13:1555-66. [https://doi.org/10.2307/3871386 https://doi.org/10.1105/tpc.13.7.1555 https://doi.org/10.1105/TPC.010047]

40.  Asif R, Yasmin R, Mustafa M, Ambreen A, Mazhar M, Rehman A, et al. Phytohormones as plant growth regulators and safe protectors against biotic and abiotic stress. In:Plant Hormones-Recent Advances, New Perspectives and Applications. London:IntechOpen;2022. [https://doi.org/10.5772/intechopen.102832]

41.  Devireddy AR, Tschaplinski TJ, Tuskan GA, Muchero W, Chen JG. Role of reactive oxygen species and hormones in plant responses to temperature changes. Int J Mol Sci 2021;22:8843. [https://doi.org/10.3390/ijms22168843]

42.  Liu Y, Huang W, Xian Z, Hu N, Lin D, Ren H, et al. Overexpression of SlGRAS40 in tomato enhances tolerance to abiotic stresses and influences auxin and gibberellin signaling. Front Plant Sci 2017;8:1659. [https://doi.org/10.3389/fpls.2017.01659]

43.  Huang W, Peng S, Xian Z, Lin D, Hu G, Yang L, et al. Overexpression of a tomato MiR171 target gene SlGRAS24 impacts multiple agronomical traits via regulating gibberellin and auxin homeostasis. Plant Biotechnol J 2017;15:472-88. [https://doi.org/10.1111/pbi.12646]

44.  Farhangi-Abriz S, Ghassemi-Golezani K. Jasmonates:Mechanisms and functions in abiotic stress tolerance of plants. Biocatal Agric Biotechnol 2019;20:101210. [https://doi.org/10.1016/j.bcab.2019.101210]

45.  Siddiqi KS, Husen A. Plant response to jasmonates:Current developments and their role in changing environment. Bull Natl Res Cent 2019;43:153. [https://doi.org/10.1186/s42269-019-0195-6]

46.  Goossens J, Fernández-Calvo P, Schweizer F, Goossens A. Jasmonates:Signal transduction components and their roles in environmental stress responses. Plant Mol Biol 2016;91:673-89. [https://doi.org/10.1007/s11103-016-0480-9]

47.  Wani AB, Chadar H, Wani AH, Singh S, Upadhyay N. Salicylic acid to decrease plant stress. Environ Chem Lett 2017;15:101-23. [https://doi.org/10.1007/s10311-016-0584-0]

48.  Santino A, Taurino M, De Domenico S, Bonsegna S, Poltronieri P, Pastor V, et al. Jasmonate signaling in plant development and defense response to multiple (a)biotic stresses. Plant Cell Rep 2013;32:1085-98. [https://doi.org/10.1007/s00299-013-1441-2]

49.  Hu Z, Shen Y, Shen F, Luo Y, Su X. Evidence for the signaling role of methyl jasmonate, methyl salicylate and benzothiazole between poplar (Populus simonii×P. pyramidalis 'opera 8277') cuttings. Trees 2009;23:1003-11. [https://doi.org/10.1007/s00468-009-0342-z]

50.  Dathe W, Ronsch H, Preiss A, Schade W, Sembdner G, Schreiber K. Endogenous plant hormones of the broad bean, Vicia faba L. (-)-jasmonic acid, a plant growth inhibitor in pericarp. Planta 1981;153:530-5. [https://doi.org/10.1007/BF00385537]

51.  Ueda J, Kato J. Isolation and identification of a senescence-promoting substance from wormwood (Artemisia absinthium L.). Plant Physiol 1980;66:246-9. [https://doi.org/10.1104/pp.66.2.246]

52.  Cipollini D. Interactive effects of lateral shading and jasmonic acid on morphology, phenology, seed production, and defense traits in Arabidopsis thaliana. Int J Plant Sci 2005;166:955-9. [https://doi.org/10.1086/432896]

53.  Popova LP, Vaklinova SG. Effect of jasmonic acid on the synthesis of ribulose-1,5-bisphosphate carboxylase-oxygenase in barley leaves. J Plant Physiol 1988;133:210-5. [https://doi.org/10.1016/S0176-1617(88)80139-1]

54.  Weidhase RA, Kramell HM, Lehmann J, Liebisch HW, Lerbs W, Parthier B. Methyljasmonate-induced changes in the polypeptide pattern of senescing barley leaf segments. Plant Sci 1987;51:177-86. [https://doi.org/10.1016/0168-9452(87)90191-9]

55.  Dar TA, Uddin M, Khan MM, Hakeem KR, Jaleel H. Jasmonates counter plant stress:A review. Environ Exp Bot 2015;115:49-57. [https://doi.org/10.1016/j.envexpbot.2015.02.010]

56.  Song SK, Choi Y, Moon YH, Kim SG, Choi YD, Lee JS. Systemic induction of a Phytolacca insularis antiviral protein gene by mechanical wounding, jasmonic acid, and abscisic acid. Plant Mol Biol 2000;43:439-50. [https://doi.org/10.1023/A:1006444322626]

57.  Ahmadi FI, Karimi K, Struik PC. Effect of exogenous application of methyl jasmonate on physiological and biochemical characteristics of Brassica napus L. Cv. talaye under salinity stress. South Afr J Bot 2018;115:5-11. [https://doi.org/10.1016/j.sajb.2017.11.018]

58.  Salimi F, Shekari F, Hamzei J. Methyl jasmonate improves salinity resistance in german chamomile (Matricaria chamomilla L.) by increasing activity of antioxidant enzymes. Acta Physiol Plant 2016;38:1. [https://doi.org/10.1007/s11738-015-2023-4]

59.  Munns R. Genes and salt tolerance:Bringing them together. New Phytologist 2005;167:645-63. [https://doi.org/10.1111/j.1469-8137.2005.01487.x]

60.  Ma C, Wang ZQ, Zhang LT, Sun MM, Lin TB. Photosynthetic responses of wheat (Triticum aestivum L.) to combined effects of drought and exogenous methyl jasmonate. Photosynthetica 2014;52:377-5. [https://doi.org/10.1007/s11099-014-0041-x]

61.  Qiu Z, Guo J, Zhu A, Zhang L, Zhang M. Exogenous jasmonic acid can enhance tolerance of wheat seedlings to salt stress. Ecotoxicol Environ Saf 2014;104:202-8. [https://doi.org/10.1016/j.ecoenv.2014.03.014]

62.  Yau SK, Ryan J. Boron toxicity tolerance in crops:A viable alternative to soil amelioration. Crop Sci 2008;48:854. [https://doi.org/10.2135/cropsci2007.10.0539]

63.  Karabal E, Yücel M, Öktem HA. Antioxidant responses of tolerant and sensitive barley cultivars to boron toxicity. Plant Sci 2003;164:925-33. [https://doi.org/10.1016/S0168-9452(03)00067-0]

64.  Papadakis IE, Dimassi KN, Bosabalidis AM, Therios IN, Patakas A, Giannakoula A. Effects of B excess on some physiological and anatomical parameters of 'navelina'orange plants grafted on two rootstocks. Environ Exp Bot 2004;51:247-57. [https://doi.org/10.1016/j.envexpbot.2003.11.004]

65.  Zhang X, Sheng J, Li F, Meng D, Shen L. Methyl jasmonate alters arginine catabolism and improves postharvest chilling tolerance in cherry tomato fruit. Postharvest Biol Technol 2012;64:160-7. [https://doi.org/10.1016/j.postharvbio.2011.07.006]

66.  Jin P, Duan Y, Wang L, Wang J, Zheng Y. Reducing chilling injury of loquat fruit by combined treatment with hot air and methyl jasmonate. Food Bioproc Tech 2014;7:2259-66. [https://doi.org/10.1007/s11947-013-1232-3]

67.  Ding CK, Wang CY, Gross KC, Smith DL. Reduction of chilling injury and transcript accumulation of heat shock proteins in tomato fruit by methyl jasmonate and methyl salicylate. Plant Sci 2001;161:1153-9. [https://doi.org/10.1016/S0168-9452(01)00521-0]

68.  Dhankhar R, Solanki, R. Effect of copper and zinc toxicity on physiological and biochemical parameters in Vigna mungo (L.) hepper. Int J Pharma Bio Sci 2011;2:553-65.

69.  Maksymiec W, Krupa Z. Effects of methyl jasmonate and excess copper on root and leaf growth. Biol Plant 2007;51:322-6. [https://doi.org/10.1007/s10535-007-0062-4]

70.  Hanaka A, Maksymiec W, Bednarek W. The effect of methyl jasmonate on selected physiological parameters of copper-treated Phaseolus coccineus Plants. Plant Growth Regul 2015;77:167-77. [https://doi.org/10.1007/s10725-015-0048-8]

71.  Turner JG, Ellis C, Devoto A. The jasmonate signal pathway. Plant Cell 2002;14:S153-64. [https://doi.org/10.1105/tpc.000679]

72.  Berni R, Luyckx M, Xu X, Legay S, Sergeant K, Hausman JF, et al. Reactive oxygen species and heavy metal stress in plants:Impact on the cell wall and secondary metabolism. Environ Exp Bot 2019;161:98-106. [https://doi.org/10.1016/j.envexpbot.2018.10.017]

73.  Raja V, Majeed U, Kang H, Andrabi KI, John R. Abiotic stress:Interplay between ROS, hormones and MAPKs. Environ Exp Bot 2017;137:142-57. [https://doi.org/10.1016/j.envexpbot.2017.02.010]

74.  Sachdev S, Ansari SA, Ansari MI, Fujita M, Hasanuzzaman M. Abiotic stress and reactive oxygen species:Generation, signaling, and defense mechanisms. Antioxidants 2021;10:277. [https://doi.org/10.3390/antiox10020277]

75.  Georgiadou EC, Kowalska E, Patla K, Kulbat K, Smoli?ska B, Leszczy?ska J, et al. Influence of heavy metals (Ni, Cu, and Zn) on nitro-oxidative stress responses, proteome regulation and allergen production in basil (Ocimum basilicum L.) plants. Front Plant Sci 2018;9:862. [https://doi.org/10.3389/fpls.2018.00862]

76.  Emamverdian A, Ding Y, Mokhberdoran F, Xie Y. Growth responses and photosynthetic indices of bamboo plant (Indocalamus latifolius) under heavy metal stress. Sci World J 2018;2018:1219364. [https://doi.org/10.1155/2018/1219364]

77.  da Rocha Nina Junior A, Furtunato Maia JM, Vitor Martins SC, Gonçalves JF. Photochemical efficiency and oxidative metabolism of tree species during acclimation to high and low irradiance. Plants (Basel) 2020;9:1047. [https://doi.org/10.3390/plants9081047]

78.  Chen LM, Lin CC, Kao CH. Copper toxicity in rice seedlings:Changes in antioxidative enzyme activities, H2O2 level, and cell wall peroxidase activity in roots. Bot Bull Acad Sin 2000;41:99-103.

79.  Ma?ecka A, Konkolewska A, Han?A, Ciszewska L, Staszak AM, Jarmuszkiewicz W, et al. Activation of antioxidative and detoxificative systems in Brassica juncea L. Plants against the toxicity of heavy metals. Sci Rep 2021;11:22345. [https://doi.org/10.1038/s41598-021-01827-w]

80.  Du Y, Zhang D, Zhou D, Liu L, Wu J, Chen H, et al. The growth of plants and indigenous bacterial community were significantly affected by cadmium contamination in soil-plant system. AMB Exp 2021;11:103. [https://doi.org/10.1186/s13568-021-01264-y]

81.  Khan M, Rolly NK, Al Azzawi TN, Imran M, Mun BG, Lee IJ, et al. Lead (Pb)-induced oxidative stress alters the morphological and physio-biochemical properties of rice (Oryza sativa L.). Agronomy 2021;11:409. [https://doi.org/10.3390/agronomy11030409]

82.  Elisa B, Marsano F, Cavaletto M, Berta G. Copper stress in Cannabis sativa roots:Morphological and pro-teomic analysis. Caryologia 2007;60:96-101. [https://doi.org/10.1080/00087114.2007.10589553]

83.  Metwali EM, Gowayed SM, Al-Maghrabi OA, Mosleh YY. Evaluation of toxic effect of copper and cadmium on growth, physiological traits and protein profile of wheat (Triticum aestivium L.), Maize (Zea mays L.) and Sorghum (Sorghum bicolor L.). World Appl Sci J 2013;21:301-14.

84.  Hakmaoui A, Ater M, Bóka K, Barón M. Copper and cadmium tolerance, uptake and effect on chloroplast ultrastructure. Studies on Salix purpurea and Phragmites australis. Zeitschr Naturforsch C 2007;62:417-26. [https://doi.org/10.1515/znc-2007-5-616]

85.  Martins JP, Conde LT, Falqueto AR, Gontijo AB. Selenium biofortified Aechmea blanchetiana (Bromeliaceae) can resist lead-induced toxicity during in vitro culture. Acta Physiol Plant 2021;43:149. [https://doi.org/10.1007/s11738-021-03323-0]

86.  Aftab T, Masroor M, Khan A, Naeem M. Effects of aluminium exposures on growth, photosynthetic efficiency, lipid peroxidation, antioxidant enzymes and artemisinin content of Artemisia annua L. J Phytol 2010;28:23-37.

87.  Sen A, Shukla KK, Singh S, Tejovathi G. Impact of heavy metals on root and shoot length of Indian mustard:An initial approach for phytoremediation. Sci Secure J Biotechnol 2014;2:48-55.

88.  Jung HI, Lee TG, Lee J, Chae MJ, Lee EJ, Kim MS, et al. Foliar-applied glutathione mitigates cadmium-induced oxidative stress by modulating antioxidant-scavenging, redox-regulating, and hormone-balancing systems in Brassica napus. Front Plant Sci 2021;12:700413. [https://doi.org/10.3389/fpls.2021.700413]

89.  Mukhtar N, Hameed M, Ashraf M, Ahmed R. Modifications in stomatal structure and function in Cenchrus ciliaris L. and Cynodon dactylon (L.) Pers. in response to cadmium stre. Pak J Bot 2013;45:351-7.

90.  Ahmed SM, Alsamman AM, Jighly A, Mubarak MH, Al-Shamaa K, Istanbuli T, et al. Genome-wide association analysis of chickpea germplasms differing for salinity tolerance based on DArTseq markers. PLoS One 2021;16:e0260709. [https://doi.org/10.1371/journal.pone.0260709]

91.  Baweja P, Kumar D, Maheshwari V, Singla M, Ritu, Parul, et al. Effect of cadmium stress on growth and development of Cicer arietinum (Fabaceae). Environ We Int J Sci Tech 2020;15:49-59.

92.  Malik DK, Sindhu SS. Production of indole acetic acid by Pseudomonas Sp.:Effect of coinoculation with Mesorhizobium Sp. cicer on nodulation and plant growth of chickpea (Cicer arietinum). Physiol Mol Biol Plants 2011;17:25-32. [https://doi.org/10.1007/s12298-010-0041-7]

93.  Shiyab S. Morphophysiological effects of chromium in sour orange (Citrus aurantium L.). Hort Sci 2019;54:829-34. [https://doi.org/10.21273/HORTSCI13809-18]

94.  Malar S, Vikram SS, Favas PJ, Perumal V. Lead heavy metal toxicity induced changes on growth and antioxidative enzymes level in water hyacinths [Eichhornia crassipes(Mart.)]. Bot Stud 2016;55:54. [https://doi.org/10.1186/s40529-014-0054-6]

95.  Joardar JC, Afrin N, Halder M. Arsenic stress on photosynthesis and growth in ipomoea aquatica. Plant Sci Today 2019;6:420. [https://doi.org/10.14719/pst.2019.6.4.589]

96.  Dias MC, Monteiro C, Moutinho-Pereira J, Correia C, Gonçalves B, Santos C. Cadmium toxicity affects photosynthesis and plant growth at different levels. Acta Physiol Plant 2013;35:1281-9. [https://doi.org/10.1007/s11738-012-1167-8]

97.  Ma T, Zhou W, Chen L, Wu L, Christie P, Liu W. Toxicity of phthalate esters to lettuce (Lactuca sativa) and the soil microbial community under different soil conditions. PLoS One 2018;13:e0208111. [https://doi.org/10.1371/journal.pone.0208111]

98.  Rocha A, Canal E, Campostrini E, Reis F, Cuzzuol G. Influence of chromium in Laguncularia racemosa (L). gaertn f. physiology. Braz J Plant Physiol 2009;21:87-94. [https://doi.org/10.1590/S1677-04202009000200001]

99.  Li J, Zhao Q, Xue B, Wu H, Song G, Zhang X. Arsenic and nutrient absorption characteristics and antioxidant response in different leaves of two ryegrass (Lolium perenne) species under arsenic stress. PLoS One 2019;14:e0225373. [https://doi.org/10.1371/journal.pone.0225373]

100.  Januskaitiene I. The effect of cadmium on several photosynthetic parameters of pea (Pisum sativum L.) at two growth stages. Zemdirbyste Agric 2012;99:71-6.

101.  ZemanováV, PavlíkováD, Hnili?ka F, Pavlík M. Arsenic toxicity-induced physiological and metabolic changes in the shoots of Pteris cretica and Spinacia oleracea. Plants (Basel) 2021;10:2009. [https://doi.org/10.3390/plants10102009]

102.  Anuradha S, Rao S. Amelioration of lead toxicity in radish (Raphanus sativus L) plants by brassinolide. J Appl Biol Sci 2011;5:43-8.

103.  Kumar MV, Ramya V, Govindaraj M, Kumar CV, Maheshwaramma S, Gokenpally S, et al. Harnessing sorghum landraces to breed high-yielding, grain mold-tolerant cultivars with high protein for drought-prone environments. Front Plant Sci 2021;12:659874. [https://doi.org/10.3389/fpls.2021.659874]

104.  Lamhamdi M, El Galiou O, Bakrim A, Nóvoa-Muñoz JC, Arias-Estévez M, Aarab A, et al. Effect of lead stress on mineral content and growth of wheat (Triticum aestivum) and spinach (Spinacia oleracea) seedlings. Saudi J Biol Sci 2013;20:29-36. [https://doi.org/10.1016/j.sjbs.2012.09.001]

105.  Singh V, Bell M. Genotypic variability in architectural development of mungbean (Vigna radiata L.) root systems and physiological relationships with shoot growth dynamics. Front Plant Sci 2021;12:725915. [https://doi.org/10.3389/fpls.2021.725915]

106.  Li C, Xiong Y, Huang Q, Xu X, Huang G. Impact of irrigation and fertilization regimes on greenhouse gas emissions from soil of mulching cultivated maize (Zea Mays L.) field in the upper reaches of yellow river, China. J Clean Prod 2020;259:120873. [https://doi.org/10.1016/j.jclepro.2020.120873]

107.  Ling T, Gao Q, Du H, Zhao Q, Ren J. Growing, physiological responses and Cd uptake of corn (Zea Mays L.) under different Cd supply. Chem Speciat Bioavail 2017;29:216-21. [https://doi.org/10.1080/09542299.2017.1400924]

108.  Várallyay S, Bódi É, Garousi F, Veres S, Kovács B. Effect of arsenic on dry weight and relative chlorophyll content in greening maize and sunflower tissues. J Microbiol Biotechnol Food Sci 2015;4:167-9. [https://doi.org/10.15414/jmbfs.2015.4.special3.167-169]

Reference

1. Fahad S, Bajwa AA, Nazir U, Anjum SA, Farooq A, Zohaib A, et al. Crop production under drought and heat stress: Plant responses and management options. Front Plant Sci 2017;8:1147. https://doi.org/10.3389/fpls.2017.01147

2. Teshome DT, Zharare GE, Naidoo S. The threat of the combined effect of biotic and abiotic stress factors in forestry under a changing climate. Front Plant Sci 2020;11:601009. https://doi.org/10.3389/fpls.2020.601009

3. Hasanuzzaman M, Nahar K, Alam M, Roychowdhury R, Fujita, M. Physiological, biochemical, and molecular mechanisms of heat stress tolerance in plants. Int J Mol Sci 2013;14:9643-84. https://doi.org/10.3390/ijms14059643

4. Zhao H, Guan J, Liang Q, Zhang X, Hu H, Zhang J. Effects of cadmium stress on growth and physiological characteristics of sassafras seedlings. Sci Rep 2021;11:9913. https://doi.org/10.1038/s41598-021-89322-0

5. Zhang H, Xu Z, Guo K, Huo Y, He G, Sun H, et al. Toxic effects of heavy metal Cd and Zn on chlorophyll, carotenoid metabolism and photosynthetic function in tobacco leaves revealed by physiological and proteomics analysis. Ecotoxicol Environ Saf 2020;202:110856. https://doi.org/10.1016/j.ecoenv.2020.110856

6. Rizvi A, Zaidi A, Ameen F, Ahmed B, AlKahtani MD, Khan MS. Heavy metal induced stress on wheat: Phytotoxicity and microbiological management. RSC Adv 2020;10:38379-403. https://doi.org/10.1039/D0RA05610C

7. Galal TM, Hassan LM, Ahmed DA, Alamri SA, Alrumman SA, Eid EM. Heavy metals uptake by the global economic crop (Pisum sativum L.) grown in contaminated soils and its associated health risks. PLoS One 2021;16:e0252229. https://doi.org/10.1371/journal.pone.0252229

8. El Sabagh A, Mbarki S, Hossain A, Iqbal MA, Islam MS, Raza A, et al. Potential role of plant growth regulators in administering crucial processes against abiotic stresses. Front Agron 2021;3:1-28. https://doi.org/10.3389/fagro.2021.648694

9. Iqbal N, Khan NA, Ferrante A, Trivellini A, Francini A, Khan MI. Ethylene role in plant growth, development and senescence: Interaction with other phytohormones. Front Plant Sci 2017;8:475. https://doi.org/10.3389/fpls.2017.00475

10. Bishop GJ, Koncz C. Brassinosteroids and plant steroid hormone signaling. Plant Cell 2002;14:S97-110. https://doi.org/10.1105/tpc.001461

11. Backer R, Rokem JS, Ilangumaran G, Lamont J, Praslickova D, Ricci E, et al. Plant growth-promoting rhizobacteria: Context, mechanisms of action, and roadmap to commercialization of biostimulants for sustainable agriculture. Front Plant Sci 2018;9:1473. https://doi.org/10.3389/fpls.2018.01473

12. Cornforth JW, Milborrow BV, Ryback G, Wareing PF. Chemistry and physiology of 'dormins' in sycamore: Identity of sycamore 'dormin' with abscisin II. Nature 1965;205:1269-70. https://doi.org/10.1038/2051269b0

13. Guschina IA, Harwood JL, Smith M, Beckett RP. Abscisic acid modifies the changes in lipids brought about by water stress in the moss Atrichum androgynum. New Phytologist 2002;156:255-64. https://doi.org/10.1046/j.1469-8137.2002.00517.x

14. Taylor IB, Burbidge A, Thompson AJ. Control of abscisic acid synthesis. J Exp Bot 2000;51:1563-74. https://doi.org/10.1093/jexbot/51.350.1563

15. Takino J, Kozaki T, Sato Y, Liu C, Ozaki T, Minami A, et al. Unveiling biosynthesis of the phytohormone abscisic acid in fungi: Unprecedented mechanism of core scaffold formation catalyzed by an unusual sesquiterpene synthase. J Am Chem Soc 2018;140:12392-5. https://doi.org/10.1021/jacs.8b08925

16. Lievens L, Pollier J, Goossens A, Beyaert R, Staal J. Abscisic acid as pathogen effector and immune regulator. Front Plant Sci 2017;8:587. https://doi.org/10.3389/fpls.2017.00587

17. Rasool N. Plant hormones: Role in alleviating biotic stress. In: Plant Hormones-Recent Advances, New Perspectives and Applications. London: IntechOpen; 2022. https://doi.org/10.5772/intechopen.102689

18. Verma V, Ravindran P, Kumar PP. Plant hormone-mediated regulation of stress responses. BMC Plant Biol 2016;16:86. https://doi.org/10.1186/s12870-016-0771-y

19. Rock CD, Quatrano RS. The role of hormones during seed development. In: Plant Hormones. Netherlands, Dordrecht: Springer; 1995. p. 671-97. https://doi.org/10.1007/978-94-011-0473-9_31

20. Miransari M, Smith DL. Plant hormones and seed germination. Environ Exp Bot 2014;99:110-21. https://doi.org/10.1016/j.envexpbot.2013.11.005

21. Ghosh UK, Islam MN, Siddiqui MN, Khan MA. Understanding the roles of osmolytes for acclimatizing plants to changing environment: A review of potential mechanism. Plant Signal Behav 2021;16:1913306. https://doi.org/10.1080/15592324.2021.1913306

22. Rabbani G, Choi I. Roles of osmolytes in protein folding and aggregation in cells and their biotechnological applications. Int J Biol Macromol 2018;109:483-91. https://doi.org/10.1016/j.ijbiomac.2017.12.100

23. Sano N, Marion-Poll A. ABA metabolism and homeostasis in seed dormancy and germination. Int J Mol Sci 2021;22:5069. https://doi.org/10.3390/ijms22105069

24. Daszkowska-Golec A. The role of abscisic acid in drought stress: How ABA helps plants to cope with drought stress. In: Drought Stress Tolerance in Plants. Vol. 2. Cham: Springer International Publishing; 2016. p. 123-51. https://doi.org/10.1007/978-3-319-32423-4_5

25. Kuromori T, Seo M, Shinozaki K. ABA transport and plant water stress responses. Trends Plant Sci 2018;23:513-22. https://doi.org/10.1016/j.tplants.2018.04.001

26. Nakashima K, Yamaguchi-Shinozaki K. ABA signaling in stress-response and seed development. Plant Cell Rep 2013;32:959-70. https://doi.org/10.1007/s00299-013-1418-1

27. Pirasteh-Anosheh H, Saed-Moucheshi A, Pakniyat H, Pessarakli M. Stomatal responses to drought stress. In: Water Stress and Crop Plants. Chichester, UK: John Wiley and Sons Ltd; 2016. p. 24-40. https://doi.org/10.1002/9781119054450.ch3

28. Hsu PK, Dubeaux G, Takahashi Y, Schroeder JI. Signaling mechanisms in abscisic acid-mediated stomatal closure. Plant J 2021;105:307-21. https://doi.org/10.1111/tpj.15067

29. Sehler R, Li J, Reager J, Ye H. Investigating relationship between soil moisture and precipitation globally using remote sensing observations. J Contemp Water Res Educ 2019;168:106-18. https://doi.org/10.1111/j.1936-704X.2019.03324.x

30. Pantin F, Monnet F, Jannaud D, Costa JM, Renaud J, Muller B, et al. The dual effect of abscisic acid on stomata. New Phytol 2013;197:65-72. https://doi.org/10.1111/nph.12013

31. Xu Z, Jiang Y, Zhou G. Response and adaptation of photosynthesis, respiration, and antioxidant systems to elevated CO2 with environmental stress in plants. Front Plant Sci 2015;6:701. https://doi.org/10.3389/fpls.2015.00701

32. Bharath P, Gahir S, Raghavendra AS. Abscisic acid-induced stomatal closure: An important component of plant defense against abiotic and biotic stress. Front Plant Sci 2021;12:615114. https://doi.org/10.3389/fpls.2021.615114

33. El Sabagh A, Islam MS, Hossain A, Iqbal MA, Mubeen M, Waleed M, et al. Phytohormones as growth regulators during abiotic stress tolerance in plants. Front Agron 2022;4:1-16. https://doi.org/10.3389/fagro.2022.765068

34. Hedden P. The current status of research on gibberellin biosynthesis. Plant Cell Physiol 2020;61:1832-49. https://doi.org/10.1093/pcp/pcaa092

35. Wang Y, Zhao J, Lu W, Deng D. Gibberellin in plant height control: Old player, new story. Plant Cell Rep 2017;36:391-8. https://doi.org/10.1007/s00299-017-2104-5

36. Salazar-Cerezo S, Martínez-Montiel N, García-Sánchez J, Pérez-y-Terrón R, Martínez-Contreras R.D. Gibberellin biosynthesis and metabolism: A convergent route for plants, fungi and bacteria. Microbiol Res 2018;208:85-98. https://doi.org/10.1016/j.micres.2018.01.010

37. Yoshida H, Hirano K, Sato T, Mitsuda N, Nomoto M, Maeo K, et al. DELLA protein functions as a transcriptional activator through the DNA binding of the Indeterminate Domain family proteins. Proc Natl Acad Sci 2014;111:7861-6. https://doi.org/10.1073/pnas.1321669111

38. Coll-Martínez B, Crosas B. How the 26S proteasome degrades ubiquitinated proteins in the cell. Biomolecules 2019;9:395. https://doi.org/10.3390/biom9090395

39. Silverstone AL, Jung HS, Dill A, Kawaide H, Kamiya Y, Sun TP. Repressing a repressor: Gibberellin-induced rapid reduction of the RGA protein in Arabidopsis. Plant Cell 2001;13:1555-66. https://doi.org/10.1105/TPC.010047

40. Asif R, Yasmin R, Mustafa M, Ambreen A, Mazhar M, Rehman A, et al. Phytohormones as plant growth regulators and safe protectors against biotic and abiotic stress. In: Plant Hormones-Recent Advances, New Perspectives and Applications. London: IntechOpen; 2022. https://doi.org/10.5772/intechopen.102832

41. Devireddy AR, Tschaplinski TJ, Tuskan GA, Muchero W, Chen JG. Role of reactive oxygen species and hormones in plant responses to temperature changes. Int J Mol Sci 2021;22:8843. https://doi.org/10.3390/ijms22168843

42. Liu Y, Huang W, Xian Z, Hu N, Lin D, Ren H, et al. Overexpression of SlGRAS40 in tomato enhances tolerance to abiotic stresses and influences auxin and gibberellin signaling. Front Plant Sci 2017;8:1659. https://doi.org/10.3389/fpls.2017.01659

43. Huang W, Peng S, Xian Z, Lin D, Hu G, Yang L, et al. Overexpression of a tomato MiR171 target gene SlGRAS24 impacts multiple agronomical traits via regulating gibberellin and auxin homeostasis. Plant Biotechnol J 2017;15:472-88. https://doi.org/10.1111/pbi.12646

44. Farhangi-Abriz S, Ghassemi-Golezani K. Jasmonates: Mechanisms and functions in abiotic stress tolerance of plants. Biocatal Agric Biotechnol 2019;20:101210. https://doi.org/10.1016/j.bcab.2019.101210

45. Siddiqi KS, Husen A. Plant response to jasmonates: Current developments and their role in changing environment. Bull Natl Res Cent 2019;43:153. https://doi.org/10.1186/s42269-019-0195-6

46. Goossens J, Fernández-Calvo P, Schweizer F, Goossens A. Jasmonates: Signal transduction components and their roles in environmental stress responses. Plant Mol Biol 2016;91:673-89. 47. Wani AB, Chadar H, Wani AH, Singh S, Upadhyay N. Salicylic acid to decrease plant stress. Environ Chem Lett 2017;15:101-23. https://doi.org/10.1007/s10311-016-0584-0

48. Santino A, Taurino M, De Domenico S, Bonsegna S, Poltronieri P, Pastor V, et al. Jasmonate signaling in plant development and defense response to multiple (a)biotic stresses. Plant Cell Rep 2013;32:1085-98. https://doi.org/10.1007/s00299-013-1441-2

49. Hu Z, Shen Y, Shen F, Luo Y, Su X. Evidence for the signaling role of methyl jasmonate, methyl salicylate and benzothiazole between poplar (Populus simonii×P. pyramidalis 'opera 8277') cuttings. Trees 2009;23:1003-11. https://doi.org/10.1007/s00468-009-0342-z

50. Dathe W, Ronsch H, Preiss A, Schade W, Sembdner G, Schreiber K. Endogenous plant hormones of the broad bean, Vicia faba L. (-)-jasmonic acid, a plant growth inhibitor in pericarp. Planta 1981;153:530-5. https://doi.org/10.1007/BF00385537

51. Ueda J, Kato J. Isolation and identification of a senescence-promoting substance from wormwood (Artemisia absinthium L.). Plant Physiol 1980;66:246-9. https://doi.org/10.1104/pp.66.2.246

52. Cipollini D. Interactive effects of lateral shading and jasmonic acid on morphology, phenology, seed production, and defense traits in Arabidopsis thaliana. Int J Plant Sci 2005;166:955-9. https://doi.org/10.1086/432896

53. Popova LP, Vaklinova SG. Effect of jasmonic acid on the synthesis of ribulose-1,5-bisphosphate carboxylase-oxygenase in barley leaves. J Plant Physiol 1988;133:210-5. https://doi.org/10.1016/S0176-1617(88)80139-1

54. Weidhase RA, Kramell HM, Lehmann J, Liebisch HW, Lerbs W, Parthier B. Methyljasmonate-induced changes in the polypeptide pattern of senescing barley leaf segments. Plant Sci 1987;51:177-86. https://doi.org/10.1016/0168-9452(87)90191-9

55. Dar TA, Uddin M, Khan MM, Hakeem KR, Jaleel H. Jasmonates counter plant stress: A review. Environ Exp Bot 2015;115:49-57. https://doi.org/10.1016/j.envexpbot.2015.02.010

56. Song SK, Choi Y, Moon YH, Kim SG, Choi YD, Lee JS. Systemic induction of a Phytolacca insularis antiviral protein gene by mechanical wounding, jasmonic acid, and abscisic acid. Plant Mol Biol 2000;43:439-50. https://doi.org/10.1023/A:1006444322626

57. Ahmadi FI, Karimi K, Struik PC. Effect of exogenous application of methyl jasmonate on physiological and biochemical characteristics of Brassica napus L. Cv. talaye under salinity stress. South Afr J Bot 2018;115:5-11. https://doi.org/10.1016/j.sajb.2017.11.018

58. Salimi F, Shekari F, Hamzei J. Methyl jasmonate improves salinity resistance in german chamomile (Matricaria chamomilla L.) by increasing activity of antioxidant enzymes. Acta Physiol Plant 2016;38:1. https://doi.org/10.1007/s11738-015-2023-4

59. Munns R. Genes and salt tolerance: Bringing them together. New Phytologist 2005;167:645-63. https://doi.org/10.1111/j.1469-8137.2005.01487.x

60. Ma C, Wang ZQ, Zhang LT, Sun MM, Lin TB. Photosynthetic responses of wheat (Triticum aestivum L.) to combined effects of drought and exogenous methyl jasmonate. Photosynthetica 2014;52:377-5. https://doi.org/10.1007/s11099-014-0041-x

61. Qiu Z, Guo J, Zhu A, Zhang L, Zhang M. Exogenous jasmonic acid can enhance tolerance of wheat seedlings to salt stress. Ecotoxicol Environ Saf 2014;104:202-8. https://doi.org/10.1016/j.ecoenv.2014.03.014

62. Yau SK, Ryan J. Boron toxicity tolerance in crops: A viable alternative to soil amelioration. Crop Sci 2008;48:854. https://doi.org/10.2135/cropsci2007.10.0539

63. Karabal E, Yücel M, Öktem HA. Antioxidant responses of tolerant and sensitive barley cultivars to boron toxicity. Plant Sci 2003;164:925-33. https://doi.org/10.1016/S0168-9452(03)00067-0

64. Papadakis IE, Dimassi KN, Bosabalidis AM, Therios IN, Patakas A, Giannakoula A. Effects of B excess on some physiological and anatomical parameters of 'navelina' orange plants grafted on two rootstocks. Environ Exp Bot 2004;51:247-57. https://doi.org/10.1016/j.envexpbot.2003.11.004

65. Zhang X, Sheng J, Li F, Meng D, Shen L. Methyl jasmonate alters arginine catabolism and improves postharvest chilling tolerance in cherry tomato fruit. Postharvest Biol Technol 2012;64:160-7. https://doi.org/10.1016/j.postharvbio.2011.07.006

66. Jin P, Duan Y, Wang L, Wang J, Zheng Y. Reducing chilling injury of loquat fruit by combined treatment with hot air and methyl jasmonate. Food Bioproc Tech 2014;7:2259-66. https://doi.org/10.1007/s11947-013-1232-3

67. Ding CK, Wang CY, Gross KC, Smith DL. Reduction of chilling injury and transcript accumulation of heat shock proteins in tomato fruit by methyl jasmonate and methyl salicylate. Plant Sci 2001;161:1153-9. https://doi.org/10.1016/S0168-9452(01)00521-0

68. Dhankhar R, Solanki, R. Effect of copper and zinc toxicity on physiological and biochemical parameters in Vigna mungo (L.) hepper. Int J Pharma Bio Sci 2011;2:553-65.

69. Maksymiec W, Krupa Z. Effects of methyl jasmonate and excess copper on root and leaf growth. Biol Plant 2007;51:322-6. https://doi.org/10.1007/s10535-007-0062-4

70. Hanaka A, Maksymiec W, Bednarek W. The effect of methyl jasmonate on selected physiological parameters of copper-treated Phaseolus coccineus Plants. Plant Growth Regul 2015;77:167-77. https://doi.org/10.1007/s10725-015-0048-8

71. Turner JG, Ellis C, Devoto A. The jasmonate signal pathway. Plant Cell 2002;14:S153-64. https://doi.org/10.1105/tpc.000679

72. Berni R, Luyckx M, Xu X, Legay S, Sergeant K, Hausman JF, et al. Reactive oxygen species and heavy metal stress in plants: Impact on the cell wall and secondary metabolism. Environ Exp Bot 2019;161:98-106. https://doi.org/10.1016/j.envexpbot.2018.10.017

73. Raja V, Majeed U, Kang H, Andrabi KI, John R. Abiotic stress: Interplay between ROS, hormones and MAPKs. Environ Exp Bot 2017;137:142-57. https://doi.org/10.1016/j.envexpbot.2017.02.010

74. Sachdev S, Ansari SA, Ansari MI, Fujita M, Hasanuzzaman M. Abiotic stress and reactive oxygen species: Generation, signaling, and defense mechanisms. Antioxidants 2021;10:277. https://doi.org/10.3390/antiox10020277

75. Georgiadou EC, Kowalska E, Patla K, Kulbat K, Smoli?ska B, Leszczy?ska J, et al. Influence of heavy metals (Ni, Cu, and Zn) on nitro-oxidative stress responses, proteome regulation and allergen production in basil (Ocimum basilicum L.) plants. Front Plant Sci 2018;9:862. https://doi.org/10.3389/fpls.2018.00862

76. Emamverdian A, Ding Y, Mokhberdoran F, Xie Y. Growth responses and photosynthetic indices of bamboo plant (Indocalamus latifolius) under heavy metal stress. Sci World J 2018;2018:1219364. https://doi.org/10.1155/2018/1219364

77. da Rocha Nina Junior A, Furtunato Maia JM, Vitor Martins SC, Gonçalves JF. Photochemical efficiency and oxidative metabolism of tree species during acclimation to high and low irradiance. Plants (Basel) 2020;9:1047. https://doi.org/10.3390/plants9081047

78. Chen LM, Lin CC, Kao CH. Copper toxicity in rice seedlings: Changes in antioxidative enzyme activities, H2O2 level, and cell wall peroxidase activity in roots. Bot Bull Acad Sin 2000;41:99-103.

79. Ma?ecka A, Konkolewska A, Han? A, Ciszewska L, Staszak AM, Jarmuszkiewicz W, et al. Activation of antioxidative and detoxificative systems in Brassica juncea L. Plants against the toxicity of heavy metals. Sci Rep 2021;11:22345. https://doi.org/10.1038/s41598-021-01827-w

80. Du Y, Zhang D, Zhou D, Liu L, Wu J, Chen H, et al. The growth of plants and indigenous bacterial community were significantly affected by cadmium contamination in soil-plant system. AMB Exp 2021;11:103. https://doi.org/10.1186/s13568-021-01264-y

81. Khan M, Rolly NK, Al Azzawi TN, Imran M, Mun BG, Lee IJ, et al. Lead (Pb)-induced oxidative stress alters the morphological and physio-biochemical properties of rice (Oryza sativa L.). Agronomy 2021;11:409. https://doi.org/10.3390/agronomy11030409

82. Elisa B, Marsano F, Cavaletto M, Berta G. Copper stress in Cannabis sativa roots: Morphological and pro-teomic analysis. Caryologia 2007;60:96-101. https://doi.org/10.1080/00087114.2007.10589553

83. Metwali EM, Gowayed SM, Al-Maghrabi OA, Mosleh YY. Evaluation of toxic effect of copper and cadmium on growth, physiological traits and protein profile of wheat (Triticum aestivium L.), Maize (Zea mays L.) and Sorghum (Sorghum bicolor L.). World Appl Sci J 2013;21:301-14.

84. Hakmaoui A, Ater M, Bóka K, Barón M. Copper and cadmium tolerance, uptake and effect on chloroplast ultrastructure. Studies on Salix purpurea and Phragmites australis. Zeitschr Naturforsch C 2007;62:417-26. https://doi.org/10.1515/znc-2007-5-616

85. Martins JP, Conde LT, Falqueto AR, Gontijo AB. Selenium biofortified Aechmea blanchetiana (Bromeliaceae) can resist lead-induced toxicity during in vitro culture. Acta Physiol Plant 2021;43:149. https://doi.org/10.1007/s11738-021-03323-0

86. Aftab T, Masroor M, Khan A, Naeem M. Effects of aluminium exposures on growth, photosynthetic efficiency, lipid peroxidation, antioxidant enzymes and artemisinin content of Artemisia annua L. J Phytol 2010;28:23-37.

87. Sen A, Shukla KK, Singh S, Tejovathi G. Impact of heavy metals on root and shoot length of Indian mustard: An initial approach for phytoremediation. Sci Secure J Biotechnol 2014;2:48-55.

88. Jung HI, Lee TG, Lee J, Chae MJ, Lee EJ, Kim MS, et al. Foliar-applied glutathione mitigates cadmium-induced oxidative stress by modulating antioxidant-scavenging, redox-regulating, and hormone-balancing systems in Brassica napus. Front Plant Sci 2021;12:700413. https://doi.org/10.3389/fpls.2021.700413

89. Mukhtar N, Hameed M, Ashraf M, Ahmed R. Modifications in stomatal structure and function in Cenchrus ciliaris L. and Cynodon dactylon (L.) Pers. in response to cadmium stre. Pak J Bot 2013;45:351-7.

90. Ahmed SM, Alsamman AM, Jighly A, Mubarak MH, Al-Shamaa K, Istanbuli T, et al. Genome-wide association analysis of chickpea germplasms differing for salinity tolerance based on DArTseq markers. PLoS One 2021;16:e0260709. https://doi.org/10.1371/journal.pone.0260709

91. Baweja P, Kumar D, Maheshwari V, Singla M, Ritu, Parul, et al. Effect of cadmium stress on growth and development of Cicer arietinum (Fabaceae). Environ We Int J Sci Tech 2020;15:49-59.

92. Malik DK, Sindhu SS. Production of indole acetic acid by Pseudomonas Sp.: Effect of coinoculation with Mesorhizobium Sp. cicer on nodulation and plant growth of chickpea (Cicer arietinum). Physiol Mol Biol Plants 2011;17:25-32. https://doi.org/10.1007/s12298-010-0041-7

93. Shiyab S. Morphophysiological effects of chromium in sour orange (Citrus aurantium L.). Hort Sci 2019;54:829-34. https://doi.org/10.21273/HORTSCI13809-18

94. Malar S, Vikram SS, Favas PJ, Perumal V. Lead heavy metal toxicity induced changes on growth and antioxidative enzymes level in water hyacinths [Eichhornia crassipes (Mart.)]. Bot Stud 2016;55:54. https://doi.org/10.1186/s40529-014-0054-6

95. Joardar JC, Afrin N, Halder M. Arsenic stress on photosynthesis and growth in ipomoea aquatica. Plant Sci Today 2019;6:420. https://doi.org/10.14719/pst.2019.6.4.589

96. Dias MC, Monteiro C, Moutinho-Pereira J, Correia C, Gonçalves B, Santos C. Cadmium toxicity affects photosynthesis and plant growth at different levels. Acta Physiol Plant 2013;35:1281-9. https://doi.org/10.1007/s11738-012-1167-8

97. Ma T, Zhou W, Chen L, Wu L, Christie P, Liu W. Toxicity of phthalate esters to lettuce (Lactuca sativa) and the soil microbial community under different soil conditions. PLoS One 2018;13:e0208111. https://doi.org/10.1371/journal.pone.0208111

98. Rocha A, Canal E, Campostrini E, Reis F, Cuzzuol G. Influence of chromium in Laguncularia racemosa (L). gaertn f. physiology. Braz J Plant Physiol 2009;21:87-94. https://doi.org/10.1590/S1677-04202009000200001

99. Li J, Zhao Q, Xue B, Wu H, Song G, Zhang X. Arsenic and nutrient absorption characteristics and antioxidant response in different leaves of two ryegrass (Lolium perenne) species under arsenic stress. PLoS One 2019;14:e0225373. https://doi.org/10.1371/journal.pone.0225373

100. Januskaitiene I. The effect of cadmium on several photosynthetic parameters of pea (Pisum sativum L.) at two growth stages. Zemdirbyste Agric 2012;99:71-6.

101. Zemanová V, Pavlíková D, Hnili?ka F, Pavlík M. Arsenic toxicity-induced physiological and metabolic changes in the shoots of Pteris cretica and Spinacia oleracea. Plants (Basel) 2021;10:2009. https://doi.org/10.3390/plants10102009

102. Anuradha S, Rao S. Amelioration of lead toxicity in radish (Raphanus sativus L) plants by brassinolide. J Appl Biol Sci 2011;5:43-8.

103. Kumar MV, Ramya V, Govindaraj M, Kumar CV, Maheshwaramma S, Gokenpally S, et al. Harnessing sorghum landraces to breed high-yielding, grain mold-tolerant cultivars with high protein for drought-prone environments. Front Plant Sci 2021;12:659874. https://doi.org/10.3389/fpls.2021.659874

104. Lamhamdi M, El Galiou O, Bakrim A, Nóvoa-Muñoz JC, Arias-Estévez M, Aarab A, et al. Effect of lead stress on mineral content and growth of wheat (Triticum aestivum) and spinach (Spinacia oleracea) seedlings. Saudi J Biol Sci 2013;20:29-36. https://doi.org/10.1016/j.sjbs.2012.09.001

105. Singh V, Bell M. Genotypic variability in architectural development of mungbean (Vigna radiata L.) root systems and physiological relationships with shoot growth dynamics. Front Plant Sci 2021;12:725915. https://doi.org/10.3389/fpls.2021.725915

106. Li C, Xiong Y, Huang Q, Xu X, Huang G. Impact of irrigation and fertilization regimes on greenhouse gas emissions from soil of mulching cultivated maize (Zea Mays L.) field in the upper reaches of yellow river, China. J Clean Prod 2020;259:120873. https://doi.org/10.1016/j.jclepro.2020.120873

107. Ling T, Gao Q, Du H, Zhao Q, Ren J. Growing, physiological responses and Cd uptake of corn (Zea Mays L.) under different Cd supply. Chem Speciat Bioavail 2017;29:216-21. https://doi.org/10.1080/09542299.2017.1400924

108. Várallyay S, Bódi É, Garousi F, Veres S, Kovács B. Effect of arsenic on dry weight and relative chlorophyll content in greening maize and sunflower tissues. J Microbiol Biotechnol Food Sci 2015;4:167-9. https://doi.org/10.15414/jmbfs.2015.4.special3.167-169

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Characterization of indole-3-acetic acid in cambial zone tissue by ultra-performance liquid chromatography-tandem mass spectrometry

Renan Marcel Bonilha Dezena, Renata Mauri, Fabio Alessandro Proença Barros, Paulo César Pires Rosa

The effects of diet and temperature on enzymes, hormones, and fecundity of the African Catfish Clarias gariepinus (Burchell 1822)

Waleed Abdul-Aziz A. Al-Deghayem, El Amin Mohamed Suliman

Screening and evaluation of PGPR strains having multiple PGP traits from hilly terrain

Teg Bahadur Singh, Vikram Sahai, Akbar Ali, Mrinalini Prasad, Arti Yadav, Preksha Shrivastav, Deepika Goyal, Prem Kumar Dantu

Applications of bacterial endophytes and their advanced identification methodologies

R. Renugadevi, M. P. Ayyappadas, V. Subha Priya, M. Flory Shobana, K. Vivekanandhan

Vitamin K2: An emerging essential nutraceutical and its market potential

Sunita Aggarwal, Siddharth Gupta, Shalini Sehgal, Praneeta Srivastava, Aparajita Sen, Garishma Gulyani, Anmol Jain

In silico identification of target fetal protein(s) in the development of polycystic ovarian syndrome

Pawan Kumar, Deepak, Vandana Saini, Annu, Neelesh Yadav, Ajit Kumar

Assessing the role of temperature as an elicitor for indole-3-acetic acid production in cyanobacterial species

Priyanka Behera, Dhanesh Kumar,, Shubhashree Mahalik

Effect of salicylic acid, jasmonic acid, and a combination of both on andrographolide production in cell suspension cultures of Andrographis paniculata (Burm.f.) Nees

Nihal Ahmed, N. Praveen