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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Plant Sci.</journal-id>
<journal-title>Frontiers in Plant Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Plant Sci.</abbrev-journal-title>
<issn pub-type="epub">1664-462X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fpls.2023.1226041</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Plant Science</subject>
<subj-group>
<subject>Review</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Uncovering the mechanisms of salicylic acid-mediated abiotic stress tolerance in horticultural crops</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Yang</surname>
<given-names>Hua</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2201122"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Fang</surname>
<given-names>Rui</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Luo</surname>
<given-names>Ling</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1819739"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Yang</surname>
<given-names>Wei</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Huang</surname>
<given-names>Qiong</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Yang</surname>
<given-names>Chunlin</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1954664"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Hui</surname>
<given-names>Wenkai</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1880284"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Gong</surname>
<given-names>Wei</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Wang</surname>
<given-names>Jingyan</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
</contrib>
</contrib-group>
<aff id="aff1">
<sup>1</sup>
<institution>Provincial Key Laboratory of Forestry Ecological Engineering of Sichuan Province, College of Forestry, Sichuan Agricultural UR.A.niversity</institution>, <addr-line>Chengdu</addr-line>, <country>China</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>School of Environment, Sichuan Agricultural University</institution>, <addr-line>Chengdu</addr-line>, <country>China</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Mohd Asgher, Baba Ghulam Shah Badshah University, India</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Tasir Sharief Per, Aligarh Muslim University, India; Muhammad Ahsan Altaf, Hainan University, China</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Hua Yang, <email xlink:href="mailto:yanghua151017@163.com">yanghua151017@163.com</email>; <email xlink:href="mailto:yanghua546@gmail.com">yanghua546@gmail.com</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>28</day>
<month>08</month>
<year>2023</year>
</pub-date>
<pub-date pub-type="collection">
<year>2023</year>
</pub-date>
<volume>14</volume>
<elocation-id>1226041</elocation-id>
<history>
<date date-type="received">
<day>20</day>
<month>05</month>
<year>2023</year>
</date>
<date date-type="accepted">
<day>28</day>
<month>07</month>
<year>2023</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2023 Yang, Fang, Luo, Yang, Huang, Yang, Hui, Gong and Wang</copyright-statement>
<copyright-year>2023</copyright-year>
<copyright-holder>Yang, Fang, Luo, Yang, Huang, Yang, Hui, Gong and Wang</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>Salicylic acid (SA) has been recognized as a promising molecule for improving abiotic stress tolerance in plants due to its ability to enhance antioxidant defense system, and promote root architecture system. Recent research has focused on uncovering the mechanisms by which SA confers abiotic stress tolerance in horticultural crops. SA has been shown to act as a signaling molecule that triggers various physiological and morphological responses in plants. SA regulates the production of reactive oxygen species (ROS). Moreover, it can also act as signaling molecule that regulate the expression of stress-responsive genes. SA can directly interact with various hormones, proteins and enzymes involved in abiotic stress tolerance. SA regulates the antioxidant enzymes activities that scavenge toxic ROS, thereby reducing oxidative damage in plants. SA can also activate protein kinases that phosphorylate and activate transcription factors involved in stress responses. Understanding these mechanisms is essential for developing effective strategies to improve crop resilience in the face of changing environmental conditions. Current information provides valuable insights for farmers and plant researchers, offering new strategies to enhance crop resilience and productivity in the face of environmental challenges. By harnessing the power of SA and its signaling pathways, farmers can develop more effective stress management techniques and optimize crop performance. Plant researchers can also explore innovative approaches to breed or engineer crops with enhanced stress tolerance, thereby contributing to sustainable agriculture and food security.</p>
</abstract>
<kwd-group>
<kwd>poor yield</kwd>
<kwd>photosynthetic impairments</kwd>
<kwd>stomatal conductance</kwd>
<kwd>signaling molecules</kwd>
<kwd>stunted growth</kwd>
</kwd-group>
<counts>
<fig-count count="5"/>
<table-count count="4"/>
<equation-count count="0"/>
<ref-count count="186"/>
<page-count count="14"/>
<word-count count="6134"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Plant Abiotic Stress</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<title>Introduction</title>
<p>Abiotic stresses are major threats to the production of horticulture crops globally (<xref ref-type="bibr" rid="B83">Kreps et&#xa0;al., 2002</xref>). Nevertheless, significant abiotic stresses like heavy metals (HMs), drought, ozone, salinity, ultra-violet radiation, nutrients, temperature fluctuations (low and high), and anthropogenic activities in the developmental period have exacerbated the agriculture system (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>) (<xref ref-type="bibr" rid="B77">Khan et&#xa0;al., 2015</xref>). From the earliest stage of seed germination through maturity, the imposed abiotic stresses can affect practically all the molecular, physiological, biochemical, and metabolic functions in plants (<xref ref-type="bibr" rid="B67">Jia et&#xa0;al., 2013</xref>). This could ultimately result in serious losses in the economic output of horticulture crops (<xref ref-type="bibr" rid="B19">Chaudhry and Sidhu, 2022</xref>; <xref ref-type="bibr" rid="B92">Liu et&#xa0;al., 2022</xref>). The approach that plants interact with numerous environmental prompts is a furthermost vital question for plant researchers. Drought, heat, and salt stresses are the most prevalent and vital environmental stresses which have adverse effects on plant growth, yield, and quality (<xref ref-type="bibr" rid="B5">Altaf et&#xa0;al., 2022a</xref>). Phytohormones are crucial in coordinating several signal transduction pathways under stress environments (<xref ref-type="bibr" rid="B76">Khan et&#xa0;al., 2020</xref>). These hormones perform a vigorous function in the plants&#x2019; development and play an important part in the plants&#x2019; external and internal factors (<xref ref-type="bibr" rid="B131">Ryu and Cho, 2015</xref>). Phytohormones are recognized as environmentally acceptable for improving horticulture crops&#x2019; tolerance against abiotic stress (<xref ref-type="bibr" rid="B96">Mantri et al., 2012</xref>). Plants&#x2019; growth is regulated by the phytohormones as they are released in small quantities as a chemical regulator (<xref ref-type="bibr" rid="B6">Altaf et&#xa0;al., 2022b</xref>).</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Physiological and molecular responses of the horticultural crops under abiotic stress.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-14-1226041-g001.tif"/>
</fig>
<p>SA is the hormone that has been shown to activate several stress-reactive genes. Every aspect of the plant&#x2019;s life is controlled by the biological substances known as plant hormones containing seed germination and distribution as well as the development of fruit (<xref ref-type="bibr" rid="B80">Klessig and Malamy, 1994</xref>). Under abiotic stress tolerance and regulation of the hormones involves in the cascade of the composite signaling from the stimulus sensitivity to the expression of the gene (<xref ref-type="bibr" rid="B153">Souza et&#xa0;al., 2017</xref>). According to previous research, the ROS-dependent signaling system and hormone responses interact to allow plants to acclimatize to biotic and abiotic challenges by improving performance (<xref ref-type="bibr" rid="B159">Tognetti et&#xa0;al., 2012</xref>).</p>
<p>It is challenging to exaggerate the importance of SA spray in preventing abiotic stresses on horticultural crops. SA is essential for increasing crop resistance to numerous abiotic stresses. SA causes a series of physiological processes i.e. better water retention, higher antioxidant activity, and reinforced cell walls. These reactions aid the plants in preventing damage brought on by stress, preserving metabolic balance, and boosting the growth (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>) (<xref ref-type="bibr" rid="B170">Xu et&#xa0;al., 2015a</xref>). SA has a crucial role in protecting horticulture crops, increasing productivity, and maintaining food security in the face of unfavorable environmental conditions by lessening the harmful effects of abiotic stress. The current review aimed to provide a comprehensive understanding of the mechanisms by which SA confers abiotic stress tolerance in horticultural crops and its potential as a tool for improving crop productivity and resilience in the face of changing environmental conditions.</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Antioxidant mechanism under abiotic stress in horticultural crops.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-14-1226041-g002.tif"/>
</fig>
</sec>
<sec id="s2">
<title>Overview of SA and its potential as a tool for improving plant growth and development</title>
<p>SA sometimes referred to technically as 2-hydroxybenzoic acid, is an aromatic phenolic molecule produced by plants that contains a hydroxyl group or its functional derivative. Phenolic compounds, particularly systemic acquired resistance (SAR), have a role in trendy activating plant defenses (<xref ref-type="bibr" rid="B162">Vlot et&#xa0;al., 2009</xref>). SA improves flowering and fruit set even under normal and stress conditions (<xref ref-type="bibr" rid="B119">Pacheco et&#xa0;al., 2013</xref>), promoting root growth, stomatal closure, and reducing transpiration (<xref ref-type="bibr" rid="B44">Gorni and Pacheco, 2016</xref>), reversing ABA effects (<xref ref-type="bibr" rid="B24">Davies, 2004</xref>), regulating gravitropism, and inhibiting fruit ripening (<xref ref-type="bibr" rid="B154">Srivastava and Dwivedi, 2000</xref>). Additionally, it has also been observed that plants of pepper and tomato treated with 05 - 10 &#xb5;M of SA had increased weight of the dry plant shoots along with the area of the leaf (<xref ref-type="bibr" rid="B134">Sardar et&#xa0;al., 2021</xref>).</p>
<p>Nevertheless, on plants growth and gas exchange parameters, the SA stimulatory effects depend on the application technique, exposure time, and ontogenetic plants stages and effective concentrations of the SA vary reliant on species (<xref ref-type="bibr" rid="B60">Horv&#xe1;th et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B134">Sardar et&#xa0;al., 2021</xref>). SA can indirectly affect flowering because it changes production and plant hormone signaling directions such as auxin, ethylene, and jasmonic acid (<xref ref-type="bibr" rid="B162">Vlot et&#xa0;al., 2009</xref>). The exogenous use of SA might cause an uprise in the quantity of endogenous bioactive GA in response, altering the plant&#x2019;s hormonal condition (<xref ref-type="bibr" rid="B106">Mukherjee and Kumar, 2007</xref>). Exogenous application improved the level of phytohormones involved in enhanced plant growth, flowering, fruit set, and ultimately yield (<xref ref-type="bibr" rid="B79">Kim et&#xa0;al., 2009</xref>).</p>
<p>SA is the hormone that controls defense of the plants against pathogens and is connected to phenolic chemicals (<xref ref-type="bibr" rid="B61">Houston et&#xa0;al., 2016</xref>). To modify the tolerance to different abiotic stimuli, SA is also crucial. SA encourages defensive photosynthetic processes, which leads to regulation in stomatal conductance, carbon dioxide (CO<sub>2</sub>) assimilation, ribulose-1,5-bisphosphate carboxylase/oxygenase activity, and chlorophyll and carotenoid content (<xref ref-type="bibr" rid="B37">Feng et&#xa0;al., 2020</xref>). By lowering cellular lipid peroxidation and hydrogen peroxide (H<sub>2</sub>O<sub>2</sub>) buildup, SA application to plants with insufficient water lower cell membrane damage in leaves. Additionally, SA-treated plants have less H<sub>2</sub>O<sub>2</sub> buildup, which boosts the activity of antioxidant enzymes like catalase (CAT), superoxide dismutase (SOD), and guaiacol peroxidase (GPOD) (<xref ref-type="bibr" rid="B3">Ahmad et&#xa0;al., 2023</xref>).</p>
<p>SA performs an important part in the plants&#x2019; developmental processes (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>). Research has shown its functions in stomatal conductance, seeds development, glycolysis, the flowering of the plants, production of the fruit, ion transpiration and accumulation (<xref ref-type="bibr" rid="B50">Harper and Balke, 1981</xref>; <xref ref-type="bibr" rid="B136">Sardar et&#xa0;al., 2022</xref>), and the rate of photosynthesis (<xref ref-type="bibr" rid="B75">Khan, et&#xa0;al., 2003</xref>). SA can regulate the antioxidant defenses and reduce oxidative damage in plants (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>) (<xref ref-type="bibr" rid="B12">Banu et&#xa0;al., 2009</xref>). SA controlled the plant nitrogen, glycine betaine (GB) formation, photosynthesis, proline metabolism, and plant water interactions under abiotic stress (<xref ref-type="bibr" rid="B110">Nazar et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B102">Miura and Tada, 2014</xref>). Additionally, abiotic stressors have been shown to induce defense-related genes and stress tolerance (<xref ref-type="bibr" rid="B127">Rivas-San and Plasencia, 2011</xref>; <xref ref-type="bibr" rid="B84">Kumar, 2014</xref>).</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Role of salicylic acid in abiotic stresses in horticultural crops focusing on sustainable productivity.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="top" align="left">Stress Name</th>
<th valign="top" align="center">Plant Name</th>
<th valign="top" align="center">References</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="left">Heavy Metals Stress</td>
<td valign="middle" align="left">Malting Barley (<italic>Hordeum uhulgare</italic> L.)</td>    <td valign="middle" align="left">
<xref ref-type="bibr" rid="B152">Song et&#xa0;al. (2014)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Cold Stress</td>
<td valign="middle" align="left">Bean (<italic>Phaseolus vulgaris</italic>)</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B116">Orabi et&#xa0;al. (2015)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Drought Stress</td>
<td valign="middle" align="left">
<italic>Portulaca oleracea</italic>
</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B132">Saheri et&#xa0;al. (2020)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Ozone Stress</td>
<td valign="middle" align="left">Strawberry (<italic>Fragaria &#xd7; ananassa</italic>)</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B27">Drogoudi and Ashmore (2000)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">UV-B radiation</td>
<td valign="middle" align="left">Pepper (<italic>Capsicum annuum</italic> L.)</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B93">Mahdavian et at. (2008)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Cold Stress</td>
<td valign="middle" align="left">Cucumber (<italic>Cucumus sativus</italic> L.)</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B54">Hawrylak-Nowak et&#xa0;al. (2010)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Cold stress</td>
<td valign="middle" align="left">Tomato (<italic>Solanum lycopersicum</italic>)</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B116">Orabi et&#xa0;al. (2015)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Salinity stress</td>
<td valign="middle" align="left">Guava (<italic>Psidium guajava</italic> L.)</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B81">Sharma et&#xa0;al. (2020)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Cold stress</td>
<td valign="middle" align="left">Pepper (<italic>Capsicum annuum</italic> L.)</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B81">Korkmaz et&#xa0;al. (2010)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Salinity Stress</td>
<td valign="middle" align="left">Mango (<italic>Mangifera indica</italic>)</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B31">Elsheery et&#xa0;al. (2020)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Drought stress</td>
<td valign="middle" align="left">Tomato (<italic>Solanum lycopersicum</italic>)</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B55">Hayat et&#xa0;al. (2008)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">Drought Stress</td>
<td valign="top" align="left">Bean (<italic>Vicia faba</italic> L.)</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B1">Abdelaal, (2015)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">UV-B radiation</td>
<td valign="top" align="left">Mango (<italic>Mangifera indica</italic>)</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B100">Mavri&#x10d; &#x10c;ermelj et&#xa0;al. (2021)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">Heat stress</td>
<td valign="top" align="left">Peas (<italic>Pisum sativum</italic>)</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B120">Pan et&#xa0;al. (2006)</xref>
</td>
</tr>
</tbody>
</table>
</table-wrap>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Exogenous salicylic acid application up-regulated antioxidant defense system and balanced redox homeostasis.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-14-1226041-g003.tif"/>
</fig>
<sec id="s2_1">
<title>SA and drought stress</title>
<p>The most important abiotic stressor limiting crop output is thought to be drought (<xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>). Only 10 - 30% of the available irrigation water is used each year (<xref ref-type="bibr" rid="B156">Szilagyi, 2003</xref>). The rise of the drought was further aided by the amalgamation of high temperatures and low precipitation (<xref ref-type="bibr" rid="B179">Yu et&#xa0;al., 2013</xref>). Global food security is facing a challenge from climate change which pollutes water resources and declining crop production (<xref ref-type="bibr" rid="B57">Hedhly et al., 2009</xref>; <xref ref-type="bibr" rid="B42">Gomes et&#xa0;al., 2019</xref>). Water shortages would impact agricultural land by up to 50% by 2050 (<xref ref-type="bibr" rid="B51">Hasanuzzaman et&#xa0;al., 2019</xref>). Moreover, lack of water is the main environmental barrier to the production of horticulture crops (<xref ref-type="bibr" rid="B160">Ullah et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B148">Sirisuntornlak et&#xa0;al., 2019</xref>). In arid and semiarid environments, the two most significant environmental stresses drought and salinity occur simultaneously (<xref ref-type="bibr" rid="B150">Slama et&#xa0;al., 2008</xref>). Extreme drought and salt stress levels affect a large portion of the earth and harm crop plant output (<xref ref-type="bibr" rid="B36">FAO, 2017</xref>). Drought negatively affects plant development growth, and other metabolic functions, and is a limiting factor in the effective production of crop plants worldwide (<xref ref-type="bibr" rid="B118">Osuagwu et&#xa0;al., 2010</xref>). The weight of the dry roots can be increased due to the drought stress and also had an impact on nutrient uptake, resulting in a decline in the potassium content of the plant&#x2019;s roots and shoots.</p>
<table-wrap id="T2" position="float">
<label>Table&#xa0;2</label>
<caption>
<p>Function of salicylic acid application in horticultural plants.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="top" align="left">Functions</th>
<th valign="top" align="left">References</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="left">Improved CO<sub>2</sub> absorption</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B132">Saheri et&#xa0;al. (2020)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Increased the plant&#x2019;s PAL (phenylalanine lyase)</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B20">Chavoushi et&#xa0;al. (2020)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Mitigates water deficits</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B18">Chakma et&#xa0;al. (2021)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Combat drought problem</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B9">Ashraf et&#xa0;al. (2013)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Opening of the stomata</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B125">Rai, 2002</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Resistance to cold stress</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B137">Sayyari et&#xa0;al. (2013)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Osmolytes Contents and Anti oxidative defense system modification</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B78">Kohli et&#xa0;al. (2018)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Lipid peroxidation and ROS level decline</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B147">Shi and Zhu (2008)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Transpiration and the rate of photosynthesis increases</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B75">Khan, et&#xa0;al., (2003)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Controls the plant nitrogen</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B110">Nazar et&#xa0;al. (2011)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">metabolic and defense pathways</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B16">B&#xfc;cker-Neto et&#xa0;al. (2017)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Antioxidant enzyme activity increase</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B71">Kazemi et&#xa0;al. (2010)</xref>
</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>Phytohormone supplementation is regarded as an efficient adaptation technique (<xref ref-type="bibr" rid="B30">El-Mageed et&#xa0;al., 2017</xref>). SA affects the absorption of ions, the development of the plant, and the transport of substances. It is the crucial plant signaling molecule regarding the plants&#x2019; biotic and abiotic stress tolerance (<xref ref-type="bibr" rid="B72">Kazemi, 2014</xref>; <xref ref-type="bibr" rid="B43">Gorni et&#xa0;al., 2017</xref>). SA treatment had a progressive effect on <italic>Portulaca oleracea</italic> growth and biomass accumulation during water deficit by improving photosynthetic pigments and CO<sub>2</sub> absorption (<xref ref-type="bibr" rid="B132">Saheri et&#xa0;al., 2020</xref>). According to <xref ref-type="bibr" rid="B3">Ahmad et&#xa0;al. (2023)</xref>, the application of SA on <italic>Carthamus tinctorius</italic> L. during a water shortage increased the plants&#x2019; phenylalanine lyase (PAL) enzyme activity and anthocyanin contents but had no impact on the dry matter (<xref ref-type="bibr" rid="B3">Ahmad et&#xa0;al., 2023</xref>). Recent findings indicate that foliar SA spray on tomato mitigates water deficits (<xref ref-type="bibr" rid="B18">Chakma et&#xa0;al., 2021</xref>). The necessity of verifying SA dosages in scenarios that farmers can use is raised by the fact that grape tomatoes are not the most widely farmed tomato in the world and that the investigational circumstances of this research cannot be reconstructed in the field.</p>
<p>Previous research suggested using SA, which is intricate in the stress signaling compared to abiotic and biotic stresses to combat drought problems (<xref ref-type="bibr" rid="B124">Rabnawaz et&#xa0;al., 2020</xref>). SA applied topically to plants stated to cut their susceptibility to stress such as drought and the maximum temperature (<xref ref-type="bibr" rid="B165">Wang and Li, 2006</xref>). Opening of the stomata, ion transport regulation, and cleaning of the HMs adverse effects are all functions of SA (<xref ref-type="bibr" rid="B125">Rai, 2002</xref>).</p>
</sec>
<sec id="s2_2">
<title>SA and cold stress</title>
<p>Environmental stresses including heat, salinity, cold, and drought harm plant growth and production and cause several physiological, molecular, morphological, and biochemical changes in plants that ultimately cause the plants to become unsustainable (<xref ref-type="bibr" rid="B4">Ahmad and Prasad, 2012</xref>). Cold stress inhibits the growth of a plant its development and plant metabolism (<xref ref-type="bibr" rid="B90">Lichtenthaler, 1998</xref>). The sensitivity and intolerance of many crops to low temperatures (cold stress) have an impact on crop yield. A sudden temperature change harms the plants (<xref ref-type="bibr" rid="B139">Schwartz et&#xa0;al., 2006</xref>). According to <xref ref-type="bibr" rid="B129">Ruelland and Zachowski, (2010)</xref>, plants typically succumb to chilling and freezing stress as a result of membrane damage, change in enzyme activity, necrosis, changes in cytoplasm viscosity, and chlorosis.</p>
<p>Low-temperature stress is most dangerous for vegetable crops regarding their productivity and quality (<xref ref-type="bibr" rid="B173">Yadav, 2010</xref>). The low temperature of about 0 - 15 &#xb0;C is vulnerable for summer vegetables, for the whole plant life cycle including germination of seeds, vegetative growth, and reproduction. Cold stress plants reproductive stages, cause them to blossom later and produce sterile pollen, which has a significant negative impact on agricultural productivity (<xref ref-type="bibr" rid="B155">Suzuki et&#xa0;al., 2008</xref>; <xref ref-type="bibr" rid="B173">Yadav, 2010</xref>). Among abiotic stresses, the low temperature has a significant impact on the existence and geographic spread of the plants, as well as on the agricultural production of plants in mountainous terrain. Disruptions to plant growth and productivity lead to significant crop failure (<xref ref-type="bibr" rid="B53">Hatfield and Prueger, 2015</xref>). Low-temperature initial effect on the plant is the membrane fluidity decrease may be the harm perception of the potential site (<xref ref-type="bibr" rid="B117">Orvar et&#xa0;al., 2000</xref>). ROS damaged the membrane due to freezing temperature with cell wall and membrane adhesions and causes the rupture of the cell (<xref ref-type="bibr" rid="B114">Olien and Smith, 1997</xref>).</p>
<p>SA application enhanced the number of fruits in cucumber and pepper (<xref ref-type="bibr" rid="B32">Elvwan and Hamahyomy, 2009</xref>; <xref ref-type="bibr" rid="B56">Hayat et&#xa0;al., 2010</xref>). SA applied on the cucumber leaves enhanced their resistance to cold (<xref ref-type="bibr" rid="B137">Sayyari et&#xa0;al., 2013</xref>). Lower acetyl SA and SA concentrations (0.1 mM and 0.5 mM) have been revealed to be highly effective against low-temperature stress in bean and tomato (<xref ref-type="bibr" rid="B116">Orabi et&#xa0;al., 2015</xref>).</p>
</sec>
<sec id="s2_3">
<title>SA and heat stress</title>
<p>When plants are subjected to abiotic stress, they cannot relocate to more hospitable conditions, which has a substantial adverse effect on the growth of the plant, development, and yield (<xref ref-type="bibr" rid="B91">Lippmann et&#xa0;al., 2019</xref>). High temperatures (HT) are a significant stress, and in recent decades, global warming has increased the rise in air temperatures (<xref ref-type="bibr" rid="B91">Lippmann et&#xa0;al., 2019</xref>). The processes by which plants adapt to extreme temperatures are therefore quite interesting. One of the most harmful pressures among the continually shifting environmental factors is the steadily rising ambient temperature. By 2100, temperatures are expected to be 1.8 &#x2013; 4.0&#xb0;C greater than they are now due to the global air temperature increase of 0.2&#xb0;C every decade (<xref ref-type="bibr" rid="B64">IPCC, 2007</xref>). Plants that are subjected to heat stress (HS) experience significant, and occasionally fatal, negative impacts. Plants have developed sophisticated HS response mechanisms to deal with such circumstances. HS influences many essential physiological functions of plants, including photosynthesis, water metabolism, and respiration. Characterizing HS reactive genes, non-coding RNAs, DNA methylation, and histone amendments have made significant strides (<xref ref-type="bibr" rid="B129">Ruelland and Zachowski, 2010</xref>).</p>
<p>For plant scientists, developing new crop cultivars resistant to HT is a big task (<xref ref-type="bibr" rid="B182">Zhang et&#xa0;al., 2006</xref>; <xref ref-type="bibr" rid="B104">Moreno and Orellana, 2011</xref>). HT affects plants in numerous means, depending on their intensity, duration, the kinds of plants present, and other environmental conditions (<xref ref-type="bibr" rid="B128">Rodr&#xed;guez et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B163">Wahid et&#xa0;al., 2007</xref>). However, the features that confer resistance to HT have yet to be identified and confirmed. Researchers studying plants under HT stress are attempting to understand the plant retorts that result in heat tolerance as well as how plants might be controlled in HT situations. Omics techniques and the development of transgenic plants by modifying target genes are two recently highly explored molecular approaches (<xref ref-type="bibr" rid="B138">Schoffl et&#xa0;al., 1999</xref>; <xref ref-type="bibr" rid="B82">Kosov&#xe1; et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B28">Duque et&#xa0;al., 2013</xref>). Investigating these underlying molecular mechanisms may help to develop stress-resistant crop variants and enable the HT cultivation of vital crop plants.</p>
<p>SA has a role in altering resistance in plants against biotic stress and abiotic stress due to the part that plays in immune system activation (<xref ref-type="bibr" rid="B69">Kawano and Bouteau, 2013</xref>). Researchers are very interested in the use of SA to enhance plant thermo tolerance as well as the physiological and biochemical mechanisms involved. SA foliar spraying promoted heat tolerance in essential horticultural plants like cucumbers and grapes (<xref ref-type="bibr" rid="B164">Wang et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B177">Yang et&#xa0;al., 2022</xref>). By controlling enzymatic activity, SA application boosts the rate of photosynthesis and preserves cell membrane stability (<xref ref-type="bibr" rid="B166">Waqas et&#xa0;al., 2021</xref>). The formation of cardenolides and antioxidants is also stimulated by SA treatments, which may be crucial in boosting resistance to HT stress (<xref ref-type="bibr" rid="B181">Zhang et&#xa0;al., 2020</xref>).</p>
</sec>
<sec id="s2_4">
<title>SA and salinity stress</title>
<p>Salinity had numerous detrimental effects on the germination of seeds, plant development, and crop yield (<xref ref-type="bibr" rid="B108">Munns and Tester, 2008</xref>). Various irrigated regions get salinized, primarily as a result of the usage of saline water. Approximately, 45 million hectares of land are damaged due to salinity (<xref ref-type="bibr" rid="B108">Munns and Tester, 2008</xref>). Plants are impacted by high salinity in a variety of aspects, including oxidative stress, water stress, nutritional issues, metabolic process changes, ion toxicity, decreased expansion and cell division, membrane disruption, and genotoxicity (<xref ref-type="bibr" rid="B52">Hasegawa et&#xa0;al., 2000</xref>; <xref ref-type="bibr" rid="B107">Munns, 2002</xref>; <xref ref-type="bibr" rid="B184">Zhu, 2007</xref>). These effects collectively by inhibiting the survival of the plant, its growth, and its development (<xref ref-type="bibr" rid="B135">Sardar et&#xa0;al., 2023</xref>).</p>
<p>A major factor restricting agricultural production is salinity (<xref ref-type="bibr" rid="B150">Slama et al., 2008</xref>). It will be difficult to feed the 9.3 billion people on the planet in 2050. According to estimates, agricultural production must increase by more than 60% from 2005 to 2007 levels by 2050. (<xref ref-type="bibr" rid="B38">Fita et&#xa0;al., 2015</xref>). However, stress factors like salinity are a constant danger to effective agricultural productivity. Climate change, groundwater penetration, seawater backflow, and human activities like fertilizer application and irrigation have all contributed to an increase in soil salinity in recent years (<xref ref-type="bibr" rid="B8">Arani et&#xa0;al., 2023</xref>). The salinity of the soil reduces plant product quality, productivity, and yield (<xref ref-type="bibr" rid="B175">Yang and Guo, 2018a</xref>). Salinity stress affects the land by about 950 million hectares, although the percentage of soil salinity is rising in the future (<xref ref-type="bibr" rid="B176">Yang and Guo, 2018b</xref>).</p>
<p>When salt stress initially appears, excessive salt and chloride ions restrict root water because it decreases the water potential surrounding the roots of the plants (<xref ref-type="bibr" rid="B112">Negr&#xe3;o et&#xa0;al., 2016</xref>). Second, too much salt and chloride in plants cause toxicity of the ion. It not only interferes with the homeostasis of ions like Na<sup>+</sup> and K<sup>+</sup> but also averts the efficient uptake of nutrients like Ca<sup>2+</sup>, leaving plants deficient in vital nutrients (<xref ref-type="bibr" rid="B183">Zhu, 2002</xref>; <xref ref-type="bibr" rid="B169">Wu, 2018</xref>). Plants experience oxidative stress due to the overabundance of ROS brought on by osmotic and ionic stressors (<xref ref-type="bibr" rid="B185">Zhu, 2016</xref>). In chloroplasts and mitochondria, excessive H<sub>2</sub>O<sub>2</sub> accumulation impairs respiration and photosynthesis under salt stress (<xref ref-type="bibr" rid="B10">Balal et&#xa0;al., 2011</xref>).</p>
<p>To address the salt issue, a variety of phenolic compounds and plant growth regulators (PGRs) are used to decrease their effects. SA is an endogenous signal molecule that functions in plants&#x2019; ability to withstand environmental challenges. Plants must be able to respond to abiotic and biotic stresses (<xref ref-type="bibr" rid="B105">Moustafa-Farag et&#xa0;al., 2020</xref>). SA also promotes several physiological processes important for plant growth (<xref ref-type="bibr" rid="B14">Basit et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B49">Guo et&#xa0;al., 2022</xref>). Antioxidant SA is regarded as having hormone-like properties. Abiotic stressors can raise the SA concentration in the plant, strengthening its ability to withstand these challenging circumstances (<xref ref-type="bibr" rid="B180">Yuan and Lin, 2008</xref>). SA is essential for stomatal conductance, photosynthesis, and transpiration. Additionally, it decreases Na<sup>+</sup> and Cl<sup>-</sup> buildup in plant tissues, enhancing antioxidant defense (<xref ref-type="bibr" rid="B136">Sardar et&#xa0;al., 2022</xref>). SA can thereby increase salt tolerance, reducing the adverse effects that occur on plants (<xref ref-type="bibr" rid="B95">Manaa et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B141">Semida et&#xa0;al., 2014</xref>). The upregulation of anti-stress mechanisms and the restoration of the development process are two examples of SA&#x2019;s beneficial effects  (<xref ref-type="bibr" rid="B94">Mahmood, 2006</xref>).</p>
</sec>
<sec id="s2_5">
<title>SA and heavy metals</title>
<p>HMs have surpassed pesticides and eminent pollutants like sulfur dioxide (SO<sub>2</sub>) and CO<sub>2</sub> (<xref ref-type="bibr" rid="B21">Chen et&#xa0;al., 2007</xref>). It is expected that HMs could overtake solid waste and nuclear waste as the most harmful contaminant (<xref ref-type="bibr" rid="B86">Lajayer et&#xa0;al., 2017</xref>). Eating is the primary way that 70% of all HMs and their derivatives enter human bodies (<xref ref-type="bibr" rid="B65">Jaishankar et&#xa0;al., 2014</xref>). HMs are emitted into the air, land, and water from numerous sources and as a result of human activity (<xref ref-type="bibr" rid="B158">Tchounwou et&#xa0;al., 2012</xref>). HMs toxicity has grown to be a significant issue in many terrestrial ecosystems all over the globe. Recently, excessive industrialization damages soil quality and agricultural yield by collecting HMs (<xref ref-type="bibr" rid="B143">Shahid et&#xa0;al., 2015</xref>).</p>
<p>Changes in the soil pH which damage the texture of the soil, the presence of various elements, and the HMs accumulation, may delay a variety of molecular as well as plant physiological processes, which may either directly or indirectly limit plant development (<xref ref-type="bibr" rid="B121">Panuccio et&#xa0;al., 2009</xref>). Several developmental and biological processes need HMs such as Cu, Mo, Ni, Mn, Co, and Zn at very low amounts (<xref ref-type="bibr" rid="B133">Salla et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B143">Shahid et&#xa0;al., 2015</xref>). However, when their concentration exceeds suboptimal levels, these metals along with four other extremely poisonous heavy metals, Be (beryllium), As (arsenic), Al (aluminum), Cd (cadmium), Pb (lead), and Hg (mercury) can significantly decrease the crop yield (<xref ref-type="bibr" rid="B122">Pierart et&#xa0;al., 2015</xref>). These harmful substances result in aberrant plant morphology and metabolic issues that lower plant production (<xref ref-type="bibr" rid="B7">Amari et&#xa0;al., 2017</xref>). ROS such as OH<sup>-</sup>, O<sub>2</sub>
<sup>-</sup>, and H<sub>2</sub>O<sub>2</sub> are produced as a result of these irregularities, which disturb the redox balance of the cells (<xref ref-type="bibr" rid="B62">Ibrahim et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B143">Shahid et&#xa0;al., 2015</xref>). This redox state imbalance is known to have a significant role in the HMs toxicity in plants. Earlier studies showed that HMs accumulation in horticulture products harms human health (<xref ref-type="bibr" rid="B161">Uzu et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B143">Shahid et&#xa0;al., 2015</xref>).</p>
<p>Metal absorption and sequestration capabilities vary across plant types and plant regions. There are considerable changes in the absorption and transport of metals across species of plants and numerous varieties of the same species plant species (<xref ref-type="bibr" rid="B142">Shahid et&#xa0;al., 2017</xref>). The majority of cereal crop plants&#x2019; roots are located 25&#xa0;cm below the surface of the soil, where the plant absorbs HMs. Metals like Cd and Pb are primarily collected in roots, which are the major target for metal accumulation (<xref ref-type="bibr" rid="B85">Kumar et&#xa0;al., 2012</xref>). The cellular transport system is one method that plants have progressed to lessen the toxicity of the metal ion.</p>
<p>Phytohormones are small signaling molecules that have a significant impact on virtually every aspect of plant development and growth. Distinct hormones can have quite different mechanisms of action for various functions. So, although several hormones may be involved in the regulation of a single function, this is shown that a single hormone may govern the range of developmental and cellular processes (<xref ref-type="bibr" rid="B45">Gray, 2004</xref>). Phytohormones such as IAA (indole-3-acetic acid), also known as auxin, abscisic acid (ABA), cytokinin (CK), gibberellin (GA), ethylene (ET), jasmonic acid (JA), brassinosteroids (BRs), cytokinin (CK), salicylic acid (SA) and the most recent discovered strigolactones (SLs) are essential for plant growth and development, signaling, and crosstalk. Phytohormones are much important for plants to protect them from biotic stress as well as abiotic stress (<xref ref-type="bibr" rid="B113">Nishiyama et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B22">Colebrook et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B172">Xu et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B16">B&#xfc;cker-Neto et&#xa0;al., 2017</xref>). In agronomical crop management strategies, phytohormones have been used as regulators of HM absorption to reduce metal toxicity (<xref ref-type="bibr" rid="B123">Piotrowska-Niczyporuk et&#xa0;al., 2012</xref>). Exogenous phytohormone application demonstrated to be safe to employ and to produce favorable outcomes for plants exposed to HM toxicity (<xref ref-type="bibr" rid="B186">Zhu et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B185">Zhu et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B98">Masood et&#xa0;al., 2016</xref>). Phytohormones like IAAs, BRs, SAs, CKs, GAs, and much more play important signaling roles, metabolic and defense pathways, but the way they alleviate HM stress is at this time a subject of great attention worldwide (<xref ref-type="bibr" rid="B16">B&#xfc;cker-Neto et&#xa0;al., 2017</xref>). Phytohormone priming is carried out to promote imminent research for crop stress treatment.</p>
<p>Under HM stress, naturally accruing SA phenolic molecule is associated with the defense response in plants (<xref ref-type="table" rid="T3">
<bold>Table&#xa0;3</bold>
</xref>). SA contributes to coordinating plant development, ripening, and abiotic stress responses (<xref ref-type="bibr" rid="B127">Rivas-San and Plasencia, 2011</xref>). Both the ABA and SA are engaged in managing the HMs response (<xref ref-type="bibr" rid="B102">Miura and Tada, 2014</xref>). The presence of SA can also control a plant&#x2019;s tolerance to salt, heat, and cell death under heavy metals toxicity (<xref ref-type="bibr" rid="B109">Naz et&#xa0;al., 2022</xref>). SA shown to activate the antioxidant defense system and regulate Cd detoxification pathways, hence reducing Cd toxicity. Cd exposure increases barley SA content (Hordeum vulgare) roots. SA and other phytohormones, such as JA and ET, commonly interact with one another (<xref ref-type="bibr" rid="B67">Jia et&#xa0;al., 2013</xref>). Plant stress response as a part of hormone production, transport, and storage result in a cascade of signaling pathways (<xref ref-type="bibr" rid="B99">Matilla-Vazquez and Matilla, 2014</xref>). ET biosynthesis enzyme activity increases in the presence of HM, and MAPKs phosphorylate ACS2 and ACS6 (<xref ref-type="bibr" rid="B149">Skottke et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B16">B&#xfc;cker-Neto et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B2">Ahmad and Anjum, 2023</xref>).</p>
<table-wrap id="T3" position="float">
<label>Table&#xa0;3</label>
<caption>
<p>SA enhanced HMs stress in different Horticultural plants.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left">Plant Name</th>
<th valign="middle" align="center">Heavy Metal</th>
<th valign="middle" align="center">Plants Adaptive Reaction</th>
<th valign="middle" align="center">Reference</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="left">Cucumber</td>
<td valign="middle" align="center">Mn</td>
<td valign="middle" align="center">Lipid peroxidation and ROS level decline.<break/>Antioxidants (Glutathione and Ascorbate) increase</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B147">Shi and Zhu (2008)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Peanut</td>
<td valign="middle" align="center">Cd</td>
<td valign="middle" align="center">Metal Toxicity alleviation.<break/>Photosynthesis, Mineral nutrients, Growth, and Chlorophyll enhancement</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B170">Xu et&#xa0;al (2015a</xref>; <xref ref-type="bibr" rid="B171">2015b</xref>)</td>
</tr>
<tr>
<td valign="middle" align="left">Onion</td>
<td valign="middle" align="center">Pb</td>
<td valign="middle" align="center">Minimize the DNA damage, Increased the CAT (Catalases), APX (Ascorbate), SOD (Guaiacol Peroxidases), and GPX (Superoxide Dismutases)</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B68">Kaur et&#xa0;al. (2021)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Spinach</td>
<td valign="middle" align="center">B</td>
<td valign="middle" align="center">Decreased the Lipid peroxidation, H<sub>2</sub>O<sub>2</sub> activity, and MDA contents, While increasing the Chlorophyll contents, CAT, and SOD activity.</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B34">Eraslan et&#xa0;al. (2008)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Tomato</td>
<td valign="middle" align="center">Cd</td>
<td valign="middle" align="center">Enhanced the plant leaves, stem and root biomass, and Photosynthetic rate</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B167">Wei et&#xa0;al. (2018)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Cucumber</td>
<td valign="middle" align="center">Mn</td>
<td valign="middle" align="center">Reduced the Mn transport from roots to the plant above parts, Lipid peroxidation, ROS contents, APX, and CAT activities. Increased the Ascorbate, SOD, GR (Glutathione reductase), POD, and DHAR (Dehydroascorbate reductase).</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B147">Shi and Zhu (2008)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Pea</td>
<td valign="middle" align="center">B</td>
<td valign="middle" align="center">Enhanced ROS production and degradation, Plant fresh and dry biomass, Carotenoids, and Chlorophyll contents. Reduction of the H<sub>2</sub>O<sub>2</sub> and MDA contents.</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B115">Oliveira et&#xa0;al. (2020)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Potato</td>
<td valign="middle" align="center">Cd</td>
<td valign="middle" align="center">Increased the antioxidant defense system, RWC (relative water contents), and chlorophyll and proline.<break/>While decreasing the O<sub>2</sub>
<sup>-</sup> (Superoxide Anion redicle), MDA (Malondialdehyde), and H<sub>2</sub>O<sub>2</sub> (Hydrogen Peroxide).</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B89">Li et&#xa0;al. (2019)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Lemon balm</td>
<td valign="middle" align="center">Ni</td>
<td valign="middle" align="center">Improved the plant fresh and dry shoot and root biomass, Carotenoid contents, and decline in the electrolyte leakage.</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B151">Soltani Maivan et&#xa0;al. (2017)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Safflower</td>
<td valign="middle" align="center">Zn</td>
<td valign="middle" align="center">Reduced oxidative stress activity. Increased in the ASC (Ascorbate) level, PC (Phytochelatin), GSH (Glutathione)</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B29">Eisalou et&#xa0;al. (2021)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Cucumber</td>
<td valign="middle" align="center">Cu</td>
<td valign="middle" align="center">Minimized the GSH, Proline, and MDA contents. Increased plant growth.</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B17">&#xc7;anakci-G&#xfc;leng&#xfc;l et&#xa0;al. (2017)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Lettuce</td>
<td valign="middle" align="center">Cd</td>
<td valign="middle" align="center">Enhancement of proline contents, Peroxidase enzyme activity, and ascorbic acid,</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B157">Talabany and Albarzinji (2023)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Pepper</td>
<td valign="middle" align="center">As</td>
<td valign="middle" align="center">Increased glutamine synthetase, Nitrogen metabolism, Nitrate reductases, and Glutamate synthase activity.</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B70">Kaya et&#xa0;al. (2022)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Periwinkle</td>
<td valign="middle" align="center">Ni</td>
<td valign="middle" align="center">Improvement of Alkaloid contents, Vincristine, and anticancer alkaloids vincristine</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B63">Idrees et&#xa0;al. (2013)</xref>
</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="s2_6">
<title>SA and ozone stress</title>
<p>Tropospheric ozone (O<sub>3</sub>) is a naturally occurring element of the atmosphere; however, since industrialization, anthropogenic emissions of NO (nitrogen oxides), NMVOCs (Non-methane volatile organic compounds), CO (Carbon monoxide), and CH<sub>4</sub> have caused the O<sub>3</sub> to be produced as a result of photochemical reactions. O<sub>3</sub> levels have increased as a result, rising from pre-industrial levels of less than 20 ppb to contemporary levels, which have boosted the average of Northern Mid-Latitude O<sub>3</sub> concentrations (30-50 ppb) (<xref ref-type="bibr" rid="B23">Cooper et&#xa0;al., 2014</xref>). Since the 1950s, mean O<sub>3</sub> levels have been increasing, with the Northern Hemisphere (NH) seeing an increase of 1 to 5 ppb/decade and the Southern Hemisphere (SH) seeing an increase of 2 ppb/decade (<xref ref-type="bibr" rid="B23">Cooper et&#xa0;al., 2014</xref>). Over the past 50 years, evidence of O<sub>3</sub> harm and damage to vegetation as a result of O<sub>3</sub> phytotoxicity has increased significantly along with these concentrations (<xref ref-type="bibr" rid="B47">Grulke and Heath, 2020</xref>). O<sub>3</sub> is a secondary pollutant, so its concentrations vary a lot in space and time. This is because of things like the season, interactions between land and coast, monsoon systems, and the exchange of air between the stratosphere and troposphere. Accordingly, in semi-rural and rural areas O<sub>3</sub> concentrations are typically maximum that are downwind of the industrial and the urbanization sources of precursor emissions, i.e., in areas vital to agriculture, forestry, and grasslands and the ecosystem services they provide (<xref ref-type="bibr" rid="B101">Mills et&#xa0;al., 2013</xref>). When O<sub>3</sub> degradation (via chemical transformation and deposition) is slower than O<sub>3</sub> creation, background O<sub>3</sub> concentrations will be increased. O<sub>3</sub> and its precursors are also carried around the planet in air masses. The duration, frequency, and magnitude of O<sub>3</sub> levels are all crucial factors in determining how plants will react, and these global processes have an impact on local to regional O<sub>3</sub> concentrations.</p>
<p>Limited research has been done on the ozone effects on plant reproductive characteristics. <xref ref-type="bibr" rid="B88">Leisner and Ainsworth (2012)</xref> examined 128 peer-reviewed studies to assess the detrimental effects of present ozone levels on the reproductive development and production of several crops. However, current levels of ambient ozone considerably reduced the number of seeds (16%), fruits (9%), and fruit weight (22%). The development and makeup of fruits vary with the growing environment and may be impacted by other environmental factors. Ozone can affect reproductive development at many distinct stages (<xref ref-type="bibr" rid="B15">Black et&#xa0;al., 2000</xref>). It was demonstrated that strawberry fruiting plants bare to ozone (184 g/m<sup>3</sup> O<sub>3</sub> for 69 days) experienced an initial acceleration of inflorescence formation, monitored by a decrease in inflorescence production rate and fruit setting rate (Drogoudi and Ashmore, 2000). Moreover, ozone treatment (156&#xa0;g m<sup>3</sup> ozone for 2 months of exposure) reduced ascorbic acid content, increased peroxidation of lipids, and decreased strawberry fruit taste because it altered the fruit&#x2019;s carbohydrate pool (<xref ref-type="bibr" rid="B73">Keutgen and Pawelzik, 2008</xref>; <xref ref-type="bibr" rid="B178">Yi et&#xa0;al., 2021</xref>). This research showed that ozone modifies the development of reproductive organs to influence fruit yield and quality. Consumers&#x2019; demands for higher-quality fruit have grown.</p>
<p>In response to relating the ozone effects to concentrations averaged at one-hour intervals, which might serve as a substitute for dosage, it is crucial to summarize concentrations in a physiologically significant way (<xref ref-type="bibr" rid="B87">Lefohn, 1992</xref>). The exposure index should, in theory, be based on the idea of effective dose (<xref ref-type="bibr" rid="B130">Runeckles, 1974</xref>), that is, it should capture the exposure features that are most closely related to the quantity of ozone that is absorbed by the plants. In this case, the absorbed dosage would be the integral of the rate of absorption (flux) over time, which could be determined for ozone by dividing the concentration at the leaf surface by the leaf conductance (<xref ref-type="bibr" rid="B39">Fowler and Cape, 1982</xref>).</p>
<p>The conduction of the atmosphere could be incorporated into this conception (<xref ref-type="bibr" rid="B48">Grunhage and Jager, 1994</xref>), when there is adequate air mixing (high air conductivity), ozone concentration, and conductance of the leaf control the diurnal pattern of ozone flux. In open-top contact chambers, this is the case. In crops, the use of radiation as a substitute for leaf conductance has been proposed due to the lack of data on leaf conductance (<xref ref-type="bibr" rid="B126">Rawlings et&#xa0;al., 1988</xref>). It is necessary to characterize exposure by using ozone concentrations during the daylight hours measured (for example, &gt;50 W/m<sup>2</sup> global radiation). However, no such distinction should be established for species that exhibit significant leaf conductivity at night. Although it is also known that other parameters, such as air humidity, soil water availability, and temperature affect leaf conductance, these factors have not yet been employed in long-term research to quantify ozone uptake or dose.</p>
</sec>
<sec id="s2_7">
<title>SA and UV-B radiation</title>
<p>Sunlight, the supreme energy source on Earth, is required for all vital biological activities. On the earth&#x2019;s surface, UV light is one of the principal solar spectrum components (<xref ref-type="bibr" rid="B26">Dotto et&#xa0;al., 2018</xref>). The geomagnetic field and stratospheric ozone layer are only two examples of the many natural barriers that exist to lessen the effects of hazardous natural radiations on biotic components (<xref ref-type="bibr" rid="B58">Herndon et&#xa0;al., 2018</xref>). Approximately, 90% of (290-320 nm) of UV-B and 100% of (100-290 nm) light cannot reach the Earth&#x2019;s surface due to the stratospheric ozone layer. Although over the past two decades, the thinning of the stratospheric ozone layer has gained attention due to the massive buildup of environmental pollutants that eventually cause UV-B rays to reach the earth&#x2019;s surface (<xref ref-type="bibr" rid="B13">Barnes et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B111">Neale et&#xa0;al., 2021</xref>). As a result, concern over UV-B radiation&#x2019;s effects on plants has increased due to its increased penetration.</p>
<p>UV-B-induced morphological alterations in plants may have an impact on how well they compete for light. Deformed morphological characteristics are the result of UV-B radiation&#x2019;s harmful impacts. UV-B exposure decreased plant leaf area, height, and dry weight while increasing auxiliary branching and leaf curling (<xref ref-type="bibr" rid="B46">Greenberg et&#xa0;al., 1997</xref>; <xref ref-type="bibr" rid="B40">Furness et&#xa0;al., 1999</xref>). After a few weeks of UV-B exposure, the leaf area, and plant dry weight of rice plants significantly reduced. <xref ref-type="bibr" rid="B168">Weih et&#xa0;al. (1998)</xref> found that improved UV-A decreased the amount of leaf area per unit plant biomass (leaf area ratio), but increased the amount of biomass produced per unit leaf area (leaf area productivity) and per unit leaf nitrogen (leaf nitrogen productivity). It was clear that these variables affected the relative growth rate and nitrogen productivity since high UV-B levels decreased the leaf area ratio, leaf area productivity, and leaf nitrogen productivity (<xref ref-type="bibr" rid="B97">Manzoor et&#xa0;al., 2023</xref>). The principal causes of UV-B-induced the loss of photosynthetic pigments, thylakoid membrane integrity, reduced rubisco enzymatic activity, and the down-regulation of photosynthesis-related gene transcript levels (<xref ref-type="bibr" rid="B66">Jansen et&#xa0;al., 1996</xref>; <xref ref-type="bibr" rid="B59">Holl&#xf3;sy, 2002</xref>). Adverisites occur from abiotic stress tolerance can be mitigated by SA application (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4</bold>
</xref>; <xref ref-type="table" rid="T4">
<bold>Table&#xa0;4</bold>
</xref>).</p>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Salicylic acid supplementation enhanced abiotic stress tolerance in horticultural crops.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-14-1226041-g004.tif"/>
</fig>
<table-wrap id="T4" position="float">
<label>Table&#xa0;4</label>
<caption>
<p>Effects of Salicylic acid use in different abiotic stresses in the plants.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left">Plant name</th>
<th valign="middle" align="center">Stress Type</th>
<th valign="middle" align="center">Antioxidant Enzyme</th>
<th valign="middle" align="center">Non-enzymatic antioxidant</th>
<th valign="middle" align="center">Stress biomarkers</th>
<th valign="middle" align="center">References</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="left">Tomato</td>
<td valign="middle" align="left">Salinity</td>
<td valign="middle" align="left">APX decreased and GPX increased</td>
<td valign="middle" align="left">Damage DNA, inactivate enzymes, and cause lipid peroxidation</td>
<td valign="middle" align="left">H<sub>2</sub>O<sub>2</sub> increased</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B103">Monteiro et&#xa0;al. (2011)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Tobacoo</td>
<td valign="middle" align="left">Ozone</td>
<td valign="middle" align="left">antioxidant enzyme decreased; enzyme BAZ-hydroxylase increased</td>
<td valign="middle" align="left">Photosynthesis decreased</td>
<td valign="middle" align="left">Antioxidant enzymes (CAT and SOD) decline</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B174">Yalpani et&#xa0;al. (1994)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Bluegrass</td>
<td valign="middle" align="left">UV-B Radiation</td>
<td valign="middle" align="left">Antioxidant enzymes (CAT and SOD) decreased</td>
<td valign="middle" align="left">Inhibition of<break/>photosynthesis, premature senescence, and altered biomass partitioning</td>
<td valign="middle" align="left">Anthocyanin and<break/>a-tocopherol contents decreased</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B35">Ervin et&#xa0;al. (2004)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Pea</td>
<td valign="middle" align="left">Heat stress</td>
<td valign="middle" align="left">MDA contents decreased</td>
<td valign="middle" align="left">Promote plant thermotolerance, PAL, and BA2H.</td>
<td valign="middle" align="left">Induce the synthesis of Hsp (Heat shock protein)</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B120">Pan et&#xa0;al. (2006)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Cucumber</td>
<td valign="middle" align="left">Cold Stress</td>
<td valign="middle" align="left">Benzoic acid 2-hydroxylase activity increased</td>
<td valign="middle" align="left">Significantly decreased the shoot biomass</td>
<td valign="middle" align="left">Reduced the H<sub>2</sub>O<sub>2</sub> accumulation</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B25">Dong et&#xa0;al. (2014)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Spinach</td>
<td valign="middle" align="left">Salinity</td>
<td valign="middle" align="left">Reduction of electrolyte linkage</td>
<td valign="middle" align="left">Enhanced root and shoot length and dry weight of the plant</td>
<td valign="middle" align="left">Increased Chlorophyll a and b and total chlorophyll contents</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B41">Gilani et&#xa0;al. (2020)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Potato</td>
<td valign="middle" align="left">HMs</td>
<td valign="middle" align="left">Declined the MDA contents</td>
<td valign="middle" align="left">Increased the RWC (relative water contents), Chlorophyll, and Proline</td>
<td valign="middle" align="left">Enhanced the ROS metabolism, SOD, CAT, EC, GR, and APX contents</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B89">Li et&#xa0;al. (2019)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Carrot</td>
<td valign="middle" align="left">Salinity</td>
<td valign="middle" align="left">Enhanced the total antioxidant activity</td>
<td valign="middle" align="left">Increased the root&#x2019;s fresh and dry weight</td>
<td valign="middle" align="left">Regulation of B and Cl ion and Proline</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B33">Eraslan et&#xa0;al. (2007)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Cucumber</td>
<td valign="middle" align="left">Heat Stress</td>
<td valign="middle" align="left">Increased the CAT and SOD activity</td>
<td valign="middle" align="left">Maximize the Photosystem II photochemical reactions</td>
<td valign="middle" align="left">Decreased the H<sub>2</sub>O<sub>2</sub> and Thiobarbituric acid reactive substances.</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B146">Shi et&#xa0;al. (2006)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Pepper</td>
<td valign="middle" align="left">Heat Stress</td>
<td valign="middle" align="left">Decreased the ROS and Increased the protective enzymes activity</td>
<td valign="middle" align="left">Maintain root vigor and cell structure integrity. Also, inhibit the water boss</td>
<td valign="middle" align="left">Reduced the oxidative damage</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B181">Zhang et&#xa0;al. (2020)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Common Bean</td>
<td valign="middle" align="left">HMs</td>
<td valign="middle" align="left">Increased the Catalase, Peroxidase, Superoxide dismutase and antioxidant enzymes, and proline</td>
<td valign="middle" align="left">Improvements in photosynthetic pigments, Nitrogen contents, and growth traits</td>
<td valign="middle" align="left">Decreased the electrolyte leakage and mineral ions uptake</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B74">Khalil et&#xa0;al. (2021)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<bold>Mung Bean</bold>
</td>
<td valign="middle" align="left">UV B Radiation</td>
<td valign="middle" align="left">Reduce the Superoxide dismutase, peroxidase, and Catalase</td>
<td valign="middle" align="left">Increased the chlorophyll contents an</td>
<td valign="middle" align="left">Enhanced the Protein contents and carotenoids</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B11">Bano and Singh (2017)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<bold>Onion</bold>
</td>
<td valign="middle" align="left">Drought stress</td>
<td valign="middle" align="left">Reduce the osmoprotectants</td>
<td valign="middle" align="left">Increased the RWC, Photosynthetic activity, and membrane stability index</td>
<td valign="middle" align="left">Increased Chlorophyll a contents</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B140">Semida et&#xa0;al. (2017)</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<bold>Garlic</bold>
</td>
<td valign="middle" align="left">Salinity</td>
<td valign="middle" align="left">Increased the bulb diameter and nick bulb diameter</td>
<td valign="middle" align="left">Increase the average clove weight, bulb diameter, number of cloves per bulb, and bulb fresh weight</td>
<td valign="middle" align="left">Enhanced the potassium concentration</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B144">Shama et&#xa0;al. (2016)</xref>
</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
</sec>
<sec id="s3" sec-type="conclusion">
<title>Conclusion and future horizon</title>
<p>SA works as a signaling molecule and regulates physiological activities in plants such as photosynthesis, metabolic, and growth processes (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5</bold>
</xref>). Numerous studies indicate that SA plays a significant part in regulating how plants react to different abiotic stressors. Depending on the plant type, the SA was proven to be efficacious in various foliar sprays/incorporation with growing media treatment forms. SA helps crops withstand the effects of salinity, cold, heat, heavy metals, and drought stressors. Foliar SA treatment proved effective in reducing the negative effects of water deficiency on plants. By boosting the activities of antioxidant enzymes, SA administered to drought-stressed seedlings improved physiological traits and decreased ROS generation, which lessened drought stress and increased plant growth. One method to reduce the low-temperature stress on crops is the application of PGRs and the unitization of genetic tools and plant breeding. PGRs are more important in increasing resistance to cold stress. There has been a lot of research on the use of exogenous SA to reduce heat exhaustion.</p>
<fig id="f5" position="float">
<label>Figure&#xa0;5</label>
<caption>
<p>Salicylic acid application protected photosynthetic system and promoted plant growth.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-14-1226041-g005.tif"/>
</fig>
<p>Future research should focus on how SA controls cross-talk with PGR operating at high, medium, and short ranges (auxins, gibberellins, cytokinins, ethylene, etc.). It will be possible to identify plant systems devoted to maintaining an optimum balance between growth and defense with further study into the double roles in the development and stress response of salicylic acid. Moreover, this innovative growth regulator has prodigious potential as a running tool for allowing crops to attain greater yields.</p>
</sec>
<sec id="s4" sec-type="author-contributions">
<title>Author contributions</title>
<p>HY and RF: Conceptualization, Literature survey, Writing major original draft, Review structure. LL and WY: Literature survey, Writing- review and editing, Figure designing. QH: Literature survey, Writing- review and editing. CY: Reviewing and editing, References collection. HY: Supervision, WH and WG, JW: Preparing figures and tables: All authors contributed to the article and approved the submitted version.</p>
</sec>
</body>
<back>
<sec id="s5" sec-type="funding-information">
<title>Funding</title>
<p>This work was supported financially by the National Key Research and Development Program of China (Program No. 2020YFD1000700) and the Sichuan Characteristic Economic Crops Innovation Team Project of National Modern Agricultural Technology Systems (sccxtd-2020-12).</p>
</sec>
<sec id="s6" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s7" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
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