P U B L I C A T I O N S CODON Italian Journal of Food Science, 2023; 35 (1): 79–90 ISSN 1120-1770 online, DOI 10.15586/ijfs.v35i1.2332 79 P U B L I C A T I O N S CODON Use of salicylic acid during cultivation of plants as a strategy to improve its metabolite profile and beneficial health effects Humberto Ramos-Sotelo, Marely G. Figueroa-Pérez* Agricultura Sustentable y Protegida. Universidad Tecnológica de Culiacán, Culiacán, México *Corresponding Author: Marely G. Figueroa-Pérez, Culiacán-Imala, Km 2, Los Ángeles, 80014 Culiacán Rosales, Sinaloa. Email: marely_100@hotmail.com Received: 2 February 2023; Accepted: 13 February 2023; Published: 9 March 2023 © 2023 Codon Publications OPEN ACCESS REVIEW Abstract Chemical elicitors in plants during cultivation have been applied in soil, hydroponic solutions, or sprayed on the leaves to induce physiological changes and stimulate the production of bioactive compounds. Salicylic acid (SA) is a phenolic compound present in plants with multiple functions, including stimulus of plant growth and induc- tion of plant defense responses under conditions of stress. Recently, the use of SA as elicitor has generated much interest, due to the growing number of studies demonstrating its positive effects in fruits, vegetables, and herbs on the induction of phytochemicals, mainly phenolic compounds, alkaloids, saponins, carotenoids, among others. The health benefits of plant materials treated with SA are mainly their antioxidant capacities determined by the 2,2'-azinobis-(3-ethylbenzothiazoline-6-sulfonic acid) (ABTS) and 2,2-diphenyl-1-picrilhidrazil (DPPH) assays and anti-inflammatory properties determined in vitro, as well as hypoglycemic and hypolipidemic properties and renal protection evaluated by in vivo studies. Therefore, the exogenous application of SA during cultivation of different plants could be an alternative to increase their economic value and could be the basis for designing stan- dardized procedures in the production of bioactive compounds. Keywords: antioxidant capacity; bioactive compounds; elicitation; health; salicylic acid Introduction Secondary metabolites of plants comprise a large group of compounds that are synthesized during their growth and do not play any role in development, photosynthe- sis, or reproduction. However, these compounds are cru- cial in several important processes, such as plant defense against environmental stresses or attack by pathogens (Cohen and Kennedy, 2010). Plants are an important source of secondary metabolites, which have been used for the production of drugs, due to their beneficial health properties, which are associ- ated with a protective effect against oxidative processes (Baenas et al., 2014). Plants growing under unfavorable environments, such as water deficit, extreme heat or cold, oxygen deficiency, among others, result in an accelerated expression of some genes related to the synthesis of sec- ondary metabolites. This effect has been associated with the generation of reactive oxygen species (ROS) during environmental stress (Jenks, 2007). It has been shown that similar effects can be produced intentionally during the cultivation of plants to improve their content of bio- active metabolites (Baenas et al., 2014). Optimizing the bioactive compounds’ composition of plant food would be a cost-effective strategy for enhanc- ing nutrition and preventing diseases among the pop- ulation. Moreover, this approach could improve the economic value of some medicinal plants (Singh and Dwivedi, 2018). One of the most used strategies for this purpose is the exogenous application of elicitors. mailto:marely_100@hotmail.com 80 Italian Journal of Food Science, 2023; 35 (1) Ramos-Sotelo H et al. and others. For some alkaloids, the nitrogen atom is not in a carbon ring (Bribi, 2018). Most alkaloids are derived from amino acids, including tryptophan, phenylala- nine, lysine, tyrosine, histidine, and ornithine. Some nonamino acid compounds such as terpenoids, purine nucleotide, and polyketide are also precursors of these compounds (Ziegler and Facchini, 2007). Glucosinolates are thioglucosides with a common structure, character- ized by side chain (R) with different aliphatic, aromatic, and heteroaromatic carbon skeletons, all derived from amino acids by a process of long chain elongation, hydroxylation, or oxidation (Vig et al., 2009). Terpenes constitute the largest class of natural prod- ucts, and these are extensively used in the industrial sector. The most important biological terpenes include carotenoids and phytosterols. Carotenoids are a group of plant pigments responsible for bright red, yellow, and orange hues in many fruits and vegetables, includ- ing alpha- carotene and beta-carotene, lutein, lycopene, β-cryptoxanthin, and zeaxanthin, among others (Singh and Sharma, 2015). On the other hand, phenolic compounds include a large group that can be classified according to the number of phenol rings that they contain; phenolic acids, stilbenes, lignans, flavonoids (flavones, flavonols, flavanones, fla- vanols, anthocyanins, chalcones, dihydrochalcones, anthocyanins, and isoflavones), and tannins. Phenolic acids are constituted chemically at least by one aro- matic ring, which has one hydrogen atom substituted by a hydroxyl group. These metabolites are divided in two groups: hydroxybenzoic acids and hydroxycinnamic acids (Vuolo et al., 2019). Stilbenes are a group of phen- ylpropanoid-derived compounds characterized by a 1,2-diphenylethylene backbone (C6-C2-C6). The lignans are a group of polyphenols comprising a large variety of individual structures, mostly consisting of two phenyl- propanoids C6–C3 linked by a bond between the central atoms of the respective side chains (position 8 or β), also called β-β’ bond (Haminiuk et al., 2012). Flavonoids have a C6–C3–C6 general structural back- bone in which two C6 units (Ring A and Ring B) consist- ing of two phenyl rings contain a heterocyclic pyrane ring (C). Due to the hydroxylation pattern and variations in the chromane ring (Ring C), flavonoids are divided into different sub-groups. Chalcones, though lacking the het- erocyclic Ring C, are still categorized as members of the flavonoid family (Tsao, 2010). Flavanols or flavan-3-ols are often commonly called catechins. These compounds and epicatechin can form polymers, which are often referred to as proanthocyanidins. These are tradition- ally considered to be condensed tannins and the hydro- lysable tannins are gallotannin or tannic acid (Haminiuk et al., 2012). Many of these compounds are naturally synthesized by plants in response to attack of pathogens (Baenas et al., 2014). Among the elicitors mostly studied in recent years is sal- icylic acid (SA), an endogenous regulator of growth in plants that controls several physiological processes, such as systemic defense signaling against biotic and abiotic stress. Several studies have shown that exogenous appli- cation of SA also generates different changes in plant physiological processes and reactions, such as prevention of ethylene production (Khan et al., 2013), increase in the growth parameters (Khandaker et al., 2011), and mod- ulation of the bioactive metabolite synthesis (Ananieva et al., 2004). In this review, we will discuss the studies conducted in the last decades regarding the exogenous application of SA during the cultivation of plants and the effect of this treat- ment on bioactive metabolites of the crop. It describes the broad variety of plant secondary metabolites induced by SA, such as phenolic compounds, terpenoids, alka- loids, among others, and their health beneficial proper- ties, which can provide an overview of the possible fields of application of the SA-elicited plant foods. Finally, we conclude our review by providing suggestions that can be applied during plant cultivation to increase the production of specific secondary metabolites. Classification and Synthesis of Bioactive Compounds Plant-based foods have generated great interest in research in recent years, in addition to providing mac- ronutrients and micronutrients. These are a rich source of bioactive compounds, which, although not being clas- sified as nutrients, or considered essential for human health, have an important beneficial impact on some dis- eases (Guerriero et al., 2018). In nature, there are three large groups of bioactive compounds, which include nitrogenous and sulfur (S) substances, terpenoid com- pounds, and the bioactive widely studied, the phenolic compounds (Cohen and Kennedy, 2010). Nitrogen (N) and sulfur are the main plant nutri- ents and serve as constituents of proteins and several other important organic compounds that exert bio- logical activities (such as alkaloids and glycosinolates). Furthermore, these compounds control yield and quality of plants (Ibrahim et al., 2012). Alkaloids are N-containing organic compounds that often contain one or more rings of carbon atoms, where nitrogen atoms are usually located and whose position of those in the carbon ring varies with different alkaloids; pyrroli- dine, pyridine, quinoline, indole, steroidal, diterpenoid, Italian Journal of Food Science, 2023; 35 (1) 81 Use of salicylic acid during cultivation The term elicitor originally included only molecules capable of inducing the synthesis of phytoalexins; how- ever, nowadays, this concept is used for all compounds that induce any type of plant defense. Elicitors can be classified as biotic or abiotic, physical or chemical, and depending on their origin and molecular structure. Biotic elicitors include lipopolysaccharides, oligosac- charides, and polysaccharides, such as pectin and cel- lulose, chitosan, chitin and glucans, galacturonides, some proteins including cellulase, cryptogein, glycopro- teins, oligandrin and pectolyase, as well as other com- pounds of complex composition, such as fungal spores, mycelia cell wall, and microbial cell wall. On the other hand, abiotic elicitors can be chemicals, for example, acetic acid, benzothiadiazole, silicon, ethanol, ethene, hydrogen peroxide, inorganic salts, and metal ions or physical as an altered gas composition, chilling, CO2, drought, extreme temperatures, high pressure, high or low osmolarity, UV irradiation, saline stress, wound- ing and ozone, among others. Furthermore, some plant hormones acts as elicitors, either when they are pro- duced by the plant in response to some pathogen attack or when they are applied exogenously in low concentra- tions to the plants during cultivation. Examples of these compounds are jasmonic acid, methyl jasmonate, eth- ylene, cytokinin, gibberellin GA3 methyl salicylate, and SA (Baenas et al., 2014). Salicylic Acid SA is a phenolic compound, consisting of a ring linked to a hydroxyl and a carboxyl group. This acid regulates physiological functions in plants, when it is exogenously Regarding phenolic compounds, these are basically derived of the malonic and shikimic acid pathways, whose precursors are derived from glycolysis and the pentose phosphate pathway, resulting in the produc- tion of different compounds, such as simple flavonoids, phenolic acids, tannins, coumarins, and anthocya- nins. Nitrogen-containing compounds, such as alkaloids, glucosinolates, and cyanogenic glycosides, are synthe- sized from amino acids like tryptophan, tyrosine, and tryptophan, also derived of the shikimic acid pathway. Terpenoids are units of isoprene, synthesized from acetyl CoA or 3-phosphoglycerate (Cohen and Kennedy, 2010) (Figure 1). Classification of Elicitors in Plants Secondary metabolites are ubiquitous in plants and serve different purposes; for example, they provide the blue color to blueberries and the red color to blood-oranges. They also contribute to the bitterness of grapefruits, to the astringency of unripe persimmon, and the texture of red wines (Vuolo et al., 2019). During plant growth, these phytochemicals are synthesized as a defense mechanism; therefore, when plants are exposed to an adverse situa- tion, such as pathogens attack, ultraviolet (UV) radiation, drought, heavy metals, nutrient deficiency, increased soil salinity, and other types of environmental stresses, the synthesis rate of secondary metabolites increases (Figure  2). These produce changes in the phytochemi- cal profile after the stress situation, and these changes depend on several factors, such as the type and intensity of the stress, the plant species, and the type of metabo- lites involved (Kulbat, 2016). Figure 1. General outline of biosynthetic pathways of secondary metabolites in plants. Primary carbon metabolism Erytrose-4- phosphate Acetyl CoA 3-phosphoglycerate Shikimic acid pathway Aromatic amino acids Mevalonic acid pathway Phenolic compounds (Flavonoids, coumarins, lignin, tannins) Nitrogen containig secondary metabolites (Alkaloids, glucosionates, cyanogenic glycosides) Terpenoid compounds (Isoprene, saponins, mono, di and triterpenes, carotenoids) Methylerythritol 4-phosphate pathway Malonic acid pathway 82 Italian Journal of Food Science, 2023; 35 (1) Ramos-Sotelo H et al. precursors: in the cytoplasm starting from phenylalanine by the phenylpropanoid pathway and in the chloroplast through the isochorismate pathway (Gondor et al., 2016) (Figure 3). Phenylalanine ammonia-lyase (PAL) is the enzyme that initiates the phenylpropanoid pathway by transforming phenylalanine into trans-cinnamic acid and NH3 through a nonoxidative deamination. Trans- cinnamic acid participates as a precursor of diverse phenolic compounds biosynthesis, such as lignin, lig- nans, and flavonoids. Furthermore, some studies indi- cate that SA is also synthesized from phenylalanine, via trans-cinnamic acid, which is then converted into SA via two intermediates: ortho coumaric acid or benzoic acid, depending on the plant species (Dempsey et al., 2011). Plants can use three biosynthetic routes to pro- duce benzoic acid: a β-oxidative and a nonoxidative route from cinnamoyl Co-A and a nonoxidative route from trans-cinnamic acid (Zhang et al., 2014). Regarding the isochorismate pathway, it is known that chorismate is synthesized in the plastid, and is then con- verted into isochorismate by an isochorismate synthase (ICS). After that, an amino acid conjugation of isocho- rismate, followed by a enzymatic conversion or a spon- taneous decomposition results in the synthesis of SA and the gene responsible for this conversion is PBS3 (Lefevere et al., 2020). Due to the importance of both PAL and ICS in SA accu- mulation, it is possible that the PAL and ICS routes are integrated by a metabolic or regulatory grid in the bio- synthesis of this hormone. The recently identified genes PBS3 and EPS1 have demonstrated to be important in applied, plays an important role in the germination of seeds, either by inhibiting germination or increasing seed vigor, depending on the concentration. Recent studies also suggest that SA is a regulator of photosynthesis, it controls chloroplast and leaf structure, stomatal closure, chlorophyll and carotenoid accumulation, and the activ- ity of important enzymes related to photosynthesis, such as RuBisCO (ribulose-1,5-bisphosphate carboxylase/oxy- genase) and carbonic anhydrase. In addition, SA partici- pates in the regulation of the alternative oxidase (AOX) route both in thermogenic and nonthermogenic plants through the induction of gene expression. AOX com- bines ubiquinol oxidation with the reduction of molec- ular oxygen to produce water in a reaction that is not sensible to inhibitors of the cytochrome oxidase path- way, which allows the control of ATP synthesis to keep the homeostasis and regulate the growth of plants. These growth-promoting effects of SA in plants are also related to the increase in photosynthesis, to changes in the hor- monal status, transpiration, and stomatal conductance (Lefevere et al., 2020). Other studies have demonstrated that SA is involved in the regulation of flowering by interacting with components of the photoperiod pathway through a CO-independent branch. SA is also involved in the regulation of senescence, which is characterized by a decrease in the photosynthetic activity, a loss of antioxidant capacity, and therefore higher levels of ROS (Dempsey et al., 2011). SA is widely distributed in plants at different basal lev- els depending on plant species. SA is synthesized by two different compartmentalized routes with different Figure 2. Production of bioactive metabolites in plants under environmental stress factors. UV radiation Chemical priming agents pathogen and herbivore attack Cold heat Drought Heavy metals Salinity Nutrient deficiency Italian Journal of Food Science, 2023; 35 (1) 83 Use of salicylic acid during cultivation the pathogen-induced SA production and to encode enzymes that catalyze reactions involved in the synthesis of SA (Dempsey et al., 2011). Pathtways Induced by SA as Elicitor SA has been intensively studied in recent years due to its function as an endogenous signal mediating local and systemic plant defense responses against pathogens (Aftab et al., 2010). In addition, it has also been demon- strated that SA is involved in the plant response to dif- ferent types of abiotic stresses such as drought, chilling, heavy metal toxicity, heat, and osmotic stress, playing a crucial role in the regulation of physiological and bio- chemical processes in plants (Vicente and Plasencia, 2011). It has been demonstrated in different plant species that exogenous application of SA induces synthesis and accumulation of antioxidant compounds. The molecular mechanism of elicitation by SA is complex and depends on the concentration applied, the growth stage and nutri- tional uptake by plants, environmental conditions, etc. SA regulates the PAL enzyme activity, which catalyzes key biosynthetic reactions to produce secondary metab- olites that act as a defense mechanism against environ- mental stresses (Puthusseri et al., 2012). Signal recognition of SA is mediated by receptors and binding sites located on the plasma membrane, such as NPR1, NPR3, and NPR4, which activate a complex cas- cade of events that is initiated by inhibiting the activity of catalase, the enzyme responsible for breaking down hydrogen peroxide in water and oxygen. This produces an increase in the H2O2 levels, which produces an elevation in OH radicals that generate an oxidative stress status in cells, which in turn initiates a series of signaling cas- cades that involve the activation of Mitogen-Activated Protein Kinases (MAPKs) and G proteins, which leads to an increase in the production of secondary metabo- lites (Khalil et al., 2018). Calcium flux also participates as signaling in plant cells after SA exogen application (Rodas-Junco et al., 2013). In unstimulated cells, Ca2+ concentrations in cytosol are maintained at lower levels; when plants are attacked by a pathogen, their abscisic acid levels increase in response to the attack, which activates calcium channels, causing a Ca2+ influx into cytosol. It is known that exogenous SA also produces an increase in abscisic acid levels in the plant, which causes the same effect as a pathogen attack, increasing the cal- cium concentrations in the cytosol. This Ca2+ influx is involved in diverse physiological and cellular processes, associated with Ca2+-binding proteins, calcium-depen- dent kinases (CDPKs), phospholipases, and through sec- ondary messengers such as inositol 1,4,5- triphosphate (IP3) and diacylglycerol (DAG). CDPKs trigger diverse signaling cascades to coordinate cellular processes such as regulation of the oxidative stress, hormonal signal- ing, and gene expression (Herrera-Vásquez et al., 2015). Furthermore, studies have shown that G-proteins play an important role in stimulating ion channels, phos- pholipases A, C, and D, ROS generation, and apopto- sis. G-protein activation stimulates the accumulation of cAMP, IP3 and DAG, which triggers the activation of PKA and PKC. This causes the phosphorylation of MAPKs, resulting in gene expression that leads to sev- eral enzymatic reactions involved in secondary metabo- lite production (Rodas-Junco et al., 2015; Ruelland et al., 2014) (Figure 4). Figure 3. Biosynthesis of SA in plants. PAL: Phenylalanine ammonia-lyase; PBS3: Proteasome subunit beta type-3; EPS1: ER-retained Pma1 Suppressing; ICS: Isochorismate synthase. Salicylic acid Chorismate Cinnamate Isochorismate Benzoate Phenylalanine Salicylic acid Salicylic acid Phenylpropanoid pathway Isochorismate pathway PAL ICS PBS3 EPS1 PBS3/EPS1 84 Italian Journal of Food Science, 2023; 35 (1) Ramos-Sotelo H et al. SA and Bioactive Compounds There are a large number of studies that demonstrate the beneficial health properties of secondary metabolites in plants, which vary depending on the compound’s struc- ture and concentration. It has been reported that some nitrogen compounds, such as alkaloids, among which are choline, trigonelline, sitsirikine, and others, exert several pharmacological effects, including anti-inflammatory, anti-diarrheal, anti-cancer activities, and therapeutic potential for hypertension, hyperlipemia, diabetes, car- diovascular system, and central nervous system diseases (Qian et al., 2017). Sulfur substances predominate in some vegetables of the cabbage family, onions, garlic, etc. Garlic and onion contain sulfur compounds, such as allicin (2-propene1- thiolsulfinate), thiosulfinates, diallyl disulfide, and S-alk(en)yl-L-cysteine sulfoxides, which have been associated with health properties such as antioxidants, antibacterial, anti-inflammatories, and inhibitors of the proliferation of human tumor cells (Higuchi et al., 2003). Many terpenoids have biological activities, particularly against certain cancers and eye disease (Johnson, 2002). On the other hand, phytosterols, mainly sitosterol, stigmasterol, and campesterol, are associated with the reduction of risk of coronary heart disease by improving LDL cholesterol concentrations. They also have anti-can- cerous properties and are immune system modulators (Chawla and Goel, 2014). Phenolic compounds have shown various beneficial effects on human health, such as anti-aging, anti-in- flammatory, antioxidant, and anti-proliferative activi- ties. So they have been pointed out as an alternative to improve the incidence of certain chronic diseases, such as diabetes, cancer, and cardiovascular diseases, through the control of oxidative stress (Lin et al., 2016). There are an increasing number of studies that evaluate the exogenous application of SA to different plant species and the effects on its content of bioactive compounds (Table 1). The foliar application of SA has increased the total phenolic compounds and total flavonoids on Solanum lycopersicum (Javanmardi and Akbari, 2016), Amaranthus tricolor L (Khandaker et al., 2011), Ocimum basilicum (Gharib, 2007), Achillea millefolium (Gorni and Pacheco, 2016), and Mentha pipperita (Figueroa- Perez et al., 2014, 2015, 2018). Figure 4. General mechanism induced by elicitation with SA. MAPKs: Mitogen-Activated Protein Kinases, cAMP: Cyclic adenosine monophosphate, IP 3 : Inositol 1,4,5-triphosphate, DAG: Diacylglycerol, and PKA and PKC: Protein kinase A and C. Abscisic acid G-protien activation MAPKs activation MAPKs activation Oxidative stress Secondary metabolism Gene expression transcriptional reprogramming Citoplasmatic acidification Phospholipases A,C and D cAMP, IP3 and DAG accumulation PKA and PKC activation Enzymetic reactions Catalase H2O2 OH O2 P H2O Receptors NPR1, NPR3/NPR4 H+ Ca2+ Cu2+ Fe2+ Ca2+ Ca2+ Ca2+ K+/Cl– Salicyclic acid Italian Journal of Food Science, 2023; 35 (1) 85 Use of salicylic acid during cultivation On the other hand, it has been shown that SA elicita- tion could improve other bioactive compounds. For example, saponins and alkaloids have been detected in Mentha pipperita (Figueroa-Pérez et al., 2018). Isatis tinctoria L. hairy root cultures treated with SA increased the alkaloid content (Qing-Yan et al., 2019). SA applied to root cultures of Stemona curtisii significantly improved the production of the alkaloids, namely, oxy- protostemonine, stemocurtisine, and stemocurtisinol (Chotikadachanarong et al., 2011). It has been found that foliar application of 2 mM SA in peppermint plants increased their phytosterols contents, such scholine, trigonelline and vinblastine (Figueroa- Perez et al., 2015). Carotenoid content, mainly crocin, was highly improved in Crocus sativus plants after SA application (Tajik et al., 2015). Likewise, foliar appli- cation of SA in yarrow (Achillea millefolium L.) plants resulted in linear increases in chlorophyll content, as well as a high production of essential oils. Furthermore, addi- tion of SA to the medium of Alternanthera tenella leaves cultured in vitro induced an increase in betacyanins of the leaves (Gorni and Pacheco, 2016). The molecular response mechanism underlying SA elic- itation involves the upregulation of some genes related to the alkaloids and flavonoids synthesis, such as aro- matic amino acid decarboxylase (AADC), YUCCA monooxygenase (YUCCA), 4-coumarate coenzyme A ligase (4CL), chalcone synthase (CHS), chalcone isomerase (CHI), and flavonoid 3′-hydroxylase (F3′H). Specifically, YUCCA gene exhibits the greatest tran- scriptional abundance for the maximal alkaloid pro- duction in Isatis tinctoria L. hairy root cultures after SA elicitation, which suggested that this gene might be more sensitive and key for inducing alkaloid biosynthe- sis (Qing-Yan et al., 2019). Health Benefits of licited Plants with SA There are several studies on the effect of elicitation with SA on the metabolite content of plants. The main effects with health benefits determined in these plants have been related to their antioxidant capacities. It has been shown that the application of salicylic acid 1 mM during cultivation of Saffron (Crocus sativus L.) increased the crocin content (a carotenoid responsible for the color of flowers) and improves the antioxidant activity of stigmas.(Tajik et al., 2015). Furthermore, elic- itation with 300 μM SA in plant cell suspension cultures of Thevetia peruviana increased the antioxidant capacity by 1.66-fold determined by the ABTS assay, compared to the control culture (Mendoza et al., 2018). Blanch et al. (2020) found that foliar application of SA (100 mg/L) during cultivation of Vitis vinifera cv Syrah plants, significantly increased (3-fold) the anti- oxidant capacity of the fruits, this effect was associ- ated to increases of some phenolic compounds, such as myricetin, trans-resveratrol and phenolic acids, mainlygallic, chlorogenic, caffeic and trans-ferulic acids. Furthermore, it has been shown that 12 mM SA applied postharvest to Kinnow mandarin under cold stor- age increased twofold the total antioxidant capacity of the fruits determined by DPPH test. SA treatment also improved the activity of the antioxidant enzymes cat- alase, peroxidase, and superoxide dismutase. These enzymes help to scavenge free radicals in the fruit, which can damage the cells under stress. These effects were related to phenolic compounds and ascorbic acid content (Haider et al., 2020). Lee et al. (2013) showed that SA-treated Aloe vera adven- titious roots cultured on MS liquid media decreased the anti-inflammatory activity in UVB-treated mouse skin cells, suppressing the activity of COX-2, NF-kB, and AP-1. Foliar spraying of 2 mM SA in Ammi visnaga potentiated the radical scavenging activity of plant extracts using DPPH assay and these effects were higher for drought stressed aerial parts sprayed with 2 mM SA. The cyto- toxic activity of extracts of Ammi visnaga were evalu- ated against different cell lines as liver cancer (HepG2), breast cancer (MCF-7), lung cancer (A549), and colon cancer (Caco2), and the major effect was observed for the methanolic extracts of the fruits, roots, aerial parts, and umbels against MCF7 cell line and fruits for HepG2 cell line (Osama, 2019). It also has been reported that appli- cation of 2 mM SA during 60 min to Centella asiatica (L.) leaves effectively inhibited nitric oxide (NO) production in LPS-stimulated RAW 264.7 macrophage cells, related to the reduction in the transcription level expression of iNOS in a dose-dependent manner, which suggests that elicitation with SA increased anti-inflammatory activ- ity in Centella asiatica leaves (Buraphaka and Putalun, 2020). On the other hand, it has been demonstrated that the administration of infusion prepared with 2 mM SA-treated peppermint to diabetic rats for 4 weeks, decreased serum glucose (up to 25%) and increased serum insulin levels (up to 75%) as compared to diabetic controls. Furthermore, these infusions prevented oxida- tive damage on pancreas β-cells, improved serum lipid profile, and decreased hepatic damage in diabetic rats (Figueroa-Perez et al., 2015). Also, they decreased the renal accumulation of 14 inflammation-related proteins, associated with glomerular hypertrophy, tubular damage, expansion of mesangial matrix, and cell death in diabetic rats (Figueroa-Pérez et al., 2018). In addition, it has been 86 Italian Journal of Food Science, 2023; 35 (1) Ramos-Sotelo H et al. Ta bl e 1. B io ac tiv e m et ab ol ite s’ c on te nt o f fo od p la nt s in r es po ns e to th e ap pl ic at io n of s al ic yl ic a ci d du ri ng it s cu lti va tio n. N o. P la nt s pe ci es D os e an d m od e of a pp lic at io n of S A Ta rg et c om po un ds a nd in cr ea se R ef er en ce 1 To m at o (S ol an um ly co pe rs ic um ) Fo lia r a pp lic at io n at 4 50 m g/ L, 3 w ee ks a fte r f ru iti ng u nd er gr ee nh ou se c on di tio ns To ta l p he no lic c om po un ds (2 .1 -fo ld ); to ta l fl av on oi ds (1 .2 -fo ld ); vi ta m in C (2 .8 -fo ld ) (J av an m ar di a nd A kb ar i, 20 16 ) 2 C hi ne se c hi ve (A lli um tu be ro su m ) Fo lia r a pp lic at io n of 5 00 a nd 1 50 μ M S A C hl or op hy ll, p he no ls a nd fl av on oi ds , v ita m in C , a nd v ol at ile co m po ne nt s (W an g et a l., 2 02 2) 3 (C or ia nd ru m S at iv um ) S up pl em en ta tio n w ith 2 25 m g/ L S A o n th e gr ow th m ed iu m d ur in g 30 d ay s G al lic , b en zo ic , f er ul ic a nd 3 -O - C af fe oy lq ui ni c ac id s, Q ue re ce tin - 3- O -r ut in oo si de , a nd g lu cr on id e an d ka em pf er ol -3 -O -r ut in os id e (K dh im e t a l., 2 02 0) 4 R ed a m ar an th (A m ar an th us tri co lo r L .) Fo lia r a pp lic at io n at 1 0− 5 M u nd er g re en ho us e co nd iti on s 1 w ee k af te r s ow in g To ta l p he no lic c om po un ds (1 .2 -fo ld ); be ta cy an in s (1 .3 -fo ld ); ch lo ro ph yl l ( 1. 3- fo ld ) (K ha nd ak er e t a l., 2 01 1) 5 S w ee t b as il (O ci m um ba si lic um L .) Fo lia r a pp lic at io n at 1 m M to 1 -m on th o ld p la nt s un de r c on tro lle d en vi ro nm en ta l c on di tio ns To ta l fl av on oi ds (1 .9 -fo ld ); to ta l p he no lic c om po un ds (1 .3 -fo ld ); to ta l fl av an ol s (1 .7 -fo ld ) (K ar al ija a nd P ar ić , 2 01 7) 6 Pe pp er m in t ( M en th a pi pe rit a) Fo lia r a pp lic at io n at 0 .5 a nd 2 m M , t w o do se s 45 a nd 6 0 da ys af te r p la nt in g R os m ar in ic a ci d (1 .7 -fo ld ); he sp er id in (1 .5 -fo ld ); ga lla to ca te ch in - ga lla te (2 .8 -fo ld ); qu er ce tin (1 .6 -fo ld ); se rja ni c ac id 3β -a ra bi no py ra no si de (9 -fo ld ); st ig m as te ry l 3 β- D -g lu co py ra no si de (2 -fo ld ); tri go ne lli ne (1 .8 -fo ld ) (F ig ue ro a Pe re z et a l., 2 01 4, 20 15 ) 7 B ro cc ol i ( B ra ss ic a ol er ac ea e) D ai ly e xo ge no us s pr ay in g at 1 00 µ M to 7 -d ay -o ld s pr ou ts o n da ys 3, 5 a nd 7 In do le g lu co si no la te (1 .3 -fo ld ) (P ér ez -B al ib re a et a l., 2 01 1) 8 Ya rr ow (A ch ill ea m ill ef ol iu m ) Fo lia r a pp lic at io n at 0 .5 m M 2 0 da ys a fte r t ra ns pl an tin g th e se ed lin gs C hl or op hy ll (1 .6 -fo ld ); es se nt ia l o ils (2 -fo ld ); to ta l p he no lic co m po un ds (1 .5 -fo ld ) (G or ni a nd P ac he co , 2 01 6) 9 C ha m om ile (M at ric ar ia ch am om ill a) Fo lia r a pp lic at io n at 7 m M o n 6- w ee k- ol d pl an ts in s ta ge o f le af ro se tte H er ni ar in (2 .4 -fo ld ); Z) - a nd (E )- 2- β- d- gl uc op yr an os yl ox y- 4- m et ho xy ci nn am ic (1 .8 -fo ld ) (D uč ai ov á et a l., 2 01 3) 10 S t J oh n’ s- w or t ( H yp er ic um pe rfo ra tu m ) Fo lia r a pp lic at io n at 2 m M in a s in gl e do se a nd h ar ve st ed 7 d ay s la te r To ta l p he no lic c om po un ds (1 .9 -fo ld ); ul ig in os in B (1 .6 -fo ld ) (d e M at os N un es e t a l., 20 14 ) 11 M ar jo ra m (O rig an um m aj or an a) Fo lia r a pp lic at io n at 1 m M o n 2- m on th - o ld p la nt s, a t t w o do se s: af te r 7 5 da ys a fte r s ow in g an d 1 w ee k la te r C hl or op hy ll (1 .4 -fo ld ); pr ol in e (1 .6 -fo ld ); m ic ro el em en ts c on te nt (1 .2 -2 -fo ld ) (G ha rib e t a l., 2 00 7) 12 G in ge r ( Zi ng ib er o ffi ci na le R os co e) Fo lia r a pp lic at io n at 1 m M a t t he s ec on d le af s ta ge o nc e a w ee k fo r 4 w ee ks M yr ic et in (2 -fo ld ); fis et in (2 -fo ld ); m or ie n (1 .6 -fo ld ); an th oc ya ni n (2 -fo ld ) (G ha se m za de h et a l., 2 01 2) 13 S ca rle t s ag e (S al vi a co cc in ea ) Fo lia r a pp lic at io n at 1 m M S A th re e tim es , e ve ry 7 d ay s to p la nt s gr ow in g un de r s al t s tre ss c on di tio ns To ta l p he no lic c om po un ds (1 .2 -fo ld ); to ta l c ar ot en oi ds (1 .4 -fo ld ) (G rz es zc zu k et a l., 2 01 8) 14 Th ym e (T hy m us v ul ga ris ) Fo lia r a pp lic at io n at 3 m M e ve ry 2 1 da ys fo r 2 m on th s to 1 -m on th - ol d pl an ts To ta l fl av on oi ds (2 -fo ld ); to ta l p he no lic c om po un ds (2 -fo ld ); ka em pf er ol -3 -g lu co si de (2 -fo ld ); ap ig en in 7 -O -g lu cu ro ni de (2 -fo ld ); ch lo ro ge ni c ac id (1 .6 -fo ld ); ro sm ar in ic a ci d (1 .3 -fo ld ); er io ci tri n (2 -fo ld ); di hy dr oq ue rc et in (2 .3 -fo ld ) (K ha lil e t a l., 2 01 8) 15 S w ee t w or m w oo d (A rt em is ia an nu a L. ) Tw en ty -o ne -d ay -o ld p la nt s gr ow in g in a h yd ro po ni c so lu tio n su pp le m en te d w ith 1 00 µ M S A fo r 5 d ay s C ar ot en oi ds (1 .2 -fo ld ); ar te m is in in (1 4- fo ld ); di hy dr o ar te m is in ic ac id (5 -fo ld ) (K um ar i e t a l., 2 01 8) 16 A lo e ve ra (A sp ho de lo id ea e) Th irt y- fiv e- da y- ol d ad ve nt iti ou s ro ot s gr ow in g in a h yd ro po ni c so lu tio n su pp le m en te d w ith 2 m M S A fo r 7 d ay s A lo e em od in (1 1- fo ld ); ch ry so ph an ol (1 3- fo ld ) (L ee e t a l., 2 01 3) (c on tin ue s) Italian Journal of Food Science, 2023; 35 (1) 87 Use of salicylic acid during cultivation shown that hypolipidemic properties of common beans sprouts can be significantly improved by elicitation with 1 and 2 mM SA, which decreased TAG intestinal absorp- tion in rats fed with a high fat and fructose (HFF) diet and supplemented with bean sprouts (10%), this bene- ficial effect was associated to an increase in hesperidin and soysaponin-I contents of elicited sprouts (Mendoza- Sanchez et al., 2019). Future Trends Research oriented to the production of functional foods has been growing over the last years due to the increas- ing interest of people to consume natural products. Bioactive metabolites extracted from medicinal plants have a great therapeutic value for which they are used all over the world. The food industry continues to look for ingredients with nutritional and nutraceutical prop- erties, to develop functional products with elevated health beneficial properties (Singh and Dwivedi, 2018). However, in many cases, the cultivation conditions of the raw material used to produce nutraceutical foods are not controlled, which generate variations in its bioactive metabolite content, and therefore, its health beneficial properties. Furthermore, in some cases, the plant has a low potential for chemical synthesis of these compounds; thus, it is important to establish a regulation of natural products used in the food industry and generate strate- gies to produce quality nutraceutical foods (Baenas et al., 2014). SA as an elicitor may be a complementary tool to breed- ing programs, production management, or genetic engineering applications. The controlled short-time elicitation with SA at low doses, during the cultivation of some medicinal plants, can be used by the producers to obtain healthier products with enhanced bioactive metabolites content. Also, these preharvest treatments with SA can be of great interest for the pharmaceuti- cal industry as tools to enhance the extractable yields of specific active compounds in plants with medicinal properties. Understanding how a plant changes its con- tent of bioactive metabolites in response to a specific SA treatment would increase the economic value of medici- nal plants and could be the basis for designing standard- ized procedures that generate high-quality nutraceutical foods. On the other hand, it would be of great interest for the evaluation of nutraceutical properties of SA-elicited plants, including biological studies, to demonstrate the potential to produce safe and valuable nonpharmacolog- ical alternatives for human health through this strategy, which may provide a new approach for disease preven- tion and treatment.Ta bl e 1. C on tin ue d. N o. P la nt s pe ci es D os e an d m od e of a pp lic at io n of S A Ta rg et c om po un ds a nd in cr ea se R ef er en ce 17 C an ol a (B ra ss ic a na pu s L. ) O ne -w ee k- ol d se ed lin gs g ro w in g in n ut rie nt s ol ut io n su pp le m en te d w ith 5 µ M S A C hl or op hy ll a (2 -fo ld ) (M on ire h et a l., 2 01 1) 18 Q ui no a (C he no po di um qu in oa ) Fo lia r a pp lic at io n at 4 00 m g/ L du rin g ve ge ta tiv e gr ow th a t 4 5 an d 60 d ay s af te r s ow in g C hl or op hy ll a (1 .5 -fo ld ) a nd b (2 .5 -fo ld ); to ta l c ar ot en oi ds (2 .4 - fo ld ); to ta l p he no lic c om po un ds (1 .5 -fo ld ) (A bd A lla h et a l., 2 01 5) 19 A rt em is in in (A rt em is ia a nn ua L) Fo lia r a pp lic at io n at 1 m M a t 1 0- da y in te rv al s st ar tin g fro m 3 0 da ys a fte r p la nt in g an d en di ng a t d ay 9 0 A rt em is in in (1 .5 -fo ld ); to ta l c hl or op hy lls (1 .3 -fo ld ); to ta l ca ro te no id s (1 .2 -fo ld ) (A fta b et a l., 2 01 0) 20 W he at (T rit ic um a es tiv um ) 0. 5 m M S A w as a dd ed to th e hy dr op on ic s ol ut io n of 2 -w ee k- ol d pl an ts a nd c ol le ct ed 7 d ay s la te r Q ue rc et in (9 -fo ld ); m yr ic et in (4 .8 -fo ld ); ru tin (1 .9 -fo ld ) (G on do r e t a l., 2 01 6) 21 B er ga m ot (M on ar da d id ym a) Fo lia r a pp lic at io n of s al ic yl ic a ci d at a c on ce nt ra tio n of 1 m M H yd ro xy ci nn am ic a ci ds , fl av on oi ds , a nd p he no lic c om po un ds (S kr yp ni k et a l., 2 02 2) 22 C en te lla a si at ic a (L .) 2 m M S A w as a dd ed to le av es fo r 4 0 m in Tr ite rp en oi ds a ro un d 1- 3- fo ld (A si at ic a ci d, m ad ec as si c ac id , as ia tic os id e, a nd m ad ec as so si de ) (B ur ap ha ka a nd P ut al un , 20 20 ) 23 A lo e ve ra (A sp ho de lo id ea e) 4 m M S A w as a dd ed to th e nu tri tiv e so lu tio n of 3 -w ee k- ol d pl an ts an d co lle ct ed 1 4 da ys la te r A lo e em od in (5 .6 -fo ld ); ch ry so ph an ol (1 2- fo ld ) (L ee e t a l., 2 01 3) 24 N ia ga ra R os ad a (V iti s la br us ca ) g ra pe Fo lia r a pp lic at io n of 1 a nd 2 m m ol L −1 S A in th e pr eh ar ve st pe rio d R ut in , c ya ni di n- 3, 5- di gl uc os id e an d 3- O -g ly co si di c de lp hi ni di n (G om es e t a l., 2 02 1) 25 Fl am e gr ap es (V iti s vi ni fe ra ) S ix a pp lic at io ns o f 0. 25 , 1 , a nd 2 m M o f S A in th e ve ra is on s ta ge . P he no lic c om po un ds , a nt ho cy an in s, a nd fl av on oi ds (V az qu ez e t a l., 2 02 2) 88 Italian Journal of Food Science, 2023; 35 (1) Ramos-Sotelo H et al. Conclusion The controlled use of elicitors of SA as preharvest treatment of some medicinal plants could be an effec- tive strategy to obtain tailored foods with enhanced health-promoting compounds. Exogenous application of SA to fruits, vegetables, and herbs during cultivation produces significant increases in the content of bioac- tive metabolites, such as phenolic compounds, alkaloids, saponins, vitamins, carotenoids, among others. These changes in the plant could improve its health beneficial properties, such as antioxidant, antidiabetic, anticancer, and antiobesogenic, among others. However, since the effects of these elicitors depend on many factors, includ- ing the plant species, it is important to conduct studies oriented to elucidate the specific effect of SA on the plant of interest, to establish protocols that result in the con- trolled production of bioactive compounds. References Abd Allah, M.M.S., El-Bassiouny, H.M.S., Elewa, T.A.E. and El-Sebai, T.N., 2015. Effect of salicylic acid and benzoic acid on growth, yield and some biochemical aspects of quinoa plant grown in sandy soil. International Journal of ChemTech Research 8(12): 216–225. https://sphinxsai.com/2015/ch_vol8_ no12/1/(216-225)V8N12CT.pdf Aftab, T., Khan, M.M.A., Idrees, M., Naeem, M. and Moinuddin. (2010). Salicylic acid acts as potent enhancer of growth, pho- tosynthesis and artemisinin production in Artemisia annua L. Journal of Crop Science and Biotechnology 13(3): 183–188. https://doi.org/10.1007/s12892-010-0040-3 Ananieva, E.A., Christov, K.N. and Popova, L.P., 2004. Exogenous treatment with salicylic acid leads to increased antioxi- dant capacity in leaves of barley plants exposed to paraquat. Journal of Plant Physiology 161(3): 319–328. https://doi. org/10.1078/0176-1617-01022 Baenas, N., García-Viguera, C. and Moreno, D.A., 2014. Elicitation: a tool for enriching the bioactive composition of foods. Molecules 19(9): 13541–13563. https://doi.org/10.3390/molecules190913541 Blanch, G.P., Gómez-Jiménez, M.C. and Del Castillo, M.L.R., 2020. Exogenous salicylic acid improves phenolic content and antioxidant activity in table grapes. Plant Foods for Human Nutrition (Dordrecht, Netherlands) 75(2): 177–183. https://doi. org/10.1007/s11130-019-00793-z Bribi, N., 2018. Pharmacological activity of alkaloids: a review. Asian Journal of Botany 1. Buraphaka, H. and Putalun, W., 2020. Stimulation of health-promot- ing triterpenoids accumulation in Centella asiatica (L.) urban leaves triggered by postharvest application of methyl jasmonate and salicylic acid elicitors. Industrial Crops and Products 146: 112171. https://doi.org/10.1016/j.indcrop.2020.112171 Chawla, R. and Goel, N., 2014. Phytosterols and its effect on human health. pp. 139–145. Chotikadachanarong, K., Dheeranupattana, S., Jatisatienr, A., Wangkarn, S., Mungkornasawakul, P., Pyne, S.G., et al. 2011. Influence of salicylic acid on alkaloid production by root cul- tures of Stemona curtisii Hook. F. Current Research Journal of Biological Science 3(4): 322–325. https://ro.uow.edu.au/cgi/view- content.cgi?article=2209&context=scipapers Cohen, S.D. and Kennedy, J.A., 2010. Plant metabolism and the environment: implications for managing phenolics. Critical Reviews in Food Science and Nutrition 50(7): 620–643. https:// doi.org/10.1080/10408390802603441 de Matos Nunes, J., Bertodo, L.O.O., da Rosa, L.M.G., Von Poser,  G.L. and Rech, S.B., 2014. Stress induction of valuable secondary metabolites in Hypericum polyanthemum accli- matized plants. South African Journal of Botany 94: 182–189. https://doi.org/10.1016/j.sajb.2014.06.014 Dempsey, D.A., Vlot, A.C., Wildermuth, M.C., Klessig, D.F. (2011). Salicylic Acid biosynthesis and metabolism. Arabidopsis Book, 9, e0156. https://doi.org/10.1199/tab.0156 Dučaiová, Z., Petruľová, V. and Repčák, M., 2013. Salicylic acid regulates secondary metabolites content in leaves of Matricaria chamomilla. Biologia (Poland) 68(5): 904–909. https://doi. org/10.2478/s11756-013-0217-z Figueroa-Perez, M.G., Gallegos-Corona, M.A., Ramos-Gomez,  M. and Reynoso-Camacho, R., 2015. Salicylic acid elicitation during cultivation of the peppermint plant improves anti-diabetic effects of its infusions. Food and Function 6(6): 1865–1874. https://doi.org/10.1039/C5FO00160A Figueroa-Pérez, M.G., Pérez-Ramírez, I.F., Enciso-Moreno,  J.A., Gallegos-Corona, M.A., Salgado, L.M. and Reynoso- Camacho,  R., 2018. Diabetic nephropathy is ameliorated with peppermint (Mentha piperita) infusions prepared from sali- cylic acid-elicited plants. Journal of Functional Foods 43: 55–61. https://doi.org/10.1016/j.jff.2018.01.029 Figueroa Perez, M.G., Rocha-Guzman, N.E., Mercado-Silva, E., Loarca-Pina, G. and Reynoso-Camacho, R., 2014. Effect of chemical elicitors on peppermint (Mentha piperita) plants and their impact on the metabolite profile and antioxidant capacity of resulting infusions. Food Chemistry 156: 273–278. https:// doi.org/10.1016/j.foodchem.2014.01.101 Gharib, F.A.E., 2007. Effect of salicylic acid on the growth, metabolic activities and oil content of basil and marjoram. International Journal of Agriculture and Biology (Pakistan) 9: 294–301. https:// agris.fao.org/agris-search/search.do?recordID=PK2007000749 Ghasemzadeh, A., Jaafar, H.Z.E., Karimi, E. and Ibrahim, M.H., 2012. Combined effect of CO2 enrichment and foliar application of salicylic acid on the production and antioxidant activities of anthocyanin, flavonoids and isoflavonoids from ginger. BMC Complementary and Alternative Medicine 12(1): 229. https:// doi.org/10.1186/1472-6882-12-229 Gomes, E., P., Borges, C., V., Monteiro, G., C., Belin, M., A., F., Minatel, I., O., Junior, A., P., et al. 2021. Preharvest salicylic acid treatments improve phenolic compounds and biogenic amines in ‘Niagara Rosada’ table grape. Postharvest Biology and Technology 176: 111505. https://doi.org/10.1016/j. postharvbio.2021.111505. https://sphinxsai.com/2015/ch_vol8_no12/1/(216-225)V8N12CT.pdf� https://sphinxsai.com/2015/ch_vol8_no12/1/(216-225)V8N12CT.pdf� https://doi.org/10.1007/s12892-010-0040-3� https://doi.org/10.1078/0176-1617-01022� https://doi.org/10.1078/0176-1617-01022� https://doi.org/10.3390/molecules190913541� https://doi.org/10.1007/s11130-019-00793-z� https://doi.org/10.1007/s11130-019-00793-z� https://doi.org/10.1016/j.indcrop.2020.112171� https://ro.uow.edu.au/cgi/viewcontent.cgi?article=2209&context=scipapers� https://ro.uow.edu.au/cgi/viewcontent.cgi?article=2209&context=scipapers� https://doi.org/10.1080/10408390802603441� https://doi.org/10.1080/10408390802603441� https://doi.org/10.1016/j.sajb.2014.06.014� https://doi.org/10.1199/tab.0156� https://doi.org/10.2478/s11756-013-0217-z� https://doi.org/10.2478/s11756-013-0217-z� https://doi.org/10.1039/C5FO00160A� https://doi.org/10.1016/j.jff.2018.01.029� https://doi.org/10.1016/j.foodchem.2014.01.101� https://doi.org/10.1016/j.foodchem.2014.01.101� https://agris.fao.org/agris-search/search.do?recordID=PK2007000749� https://agris.fao.org/agris-search/search.do?recordID=PK2007000749� https://doi.org/10.1186/1472-6882-12-229� https://doi.org/10.1186/1472-6882-12-229� https://doi.org/10.1016/j.postharvbio.2021.111505� https://doi.org/10.1016/j.postharvbio.2021.111505� Italian Journal of Food Science, 2023; 35 (1) 89 Use of salicylic acid during cultivation Gondor, O.K., Janda, T., Soós, V., Pál, M., Majláth, I., Adak, M.K., et al. 2016. Salicylic acid induction of flavonoid biosynthesis pathways in wheat varies by treatment. Frontiers in Plant Science 7: 1–12. https://doi.org/10.3389/fpls.2016.01447 Gorni, P.H. and Pacheco, C., 2016. Growth promotion and elicitor activity of salicylic acid in Achillea millefolium L. African Journal of Biotechnology 15(16): 657–665. https://doi.org/10.5897/ AJB2016.15320 Grzeszczuk, M., Salachna, P. and Meller, E., 2018. Changes in pho- tosynthetic pigments, total phenolic content, and antioxidant activity of Salvia coccinea Buc’hoz Ex Etl. Induced by exogenous salicylic acid and soil salinity. Molecules (Basel, Switzerland) 23(6), 1296. https://doi.org/10.3390/molecules23061296 Guerriero, G., Berni, R., Munoz-Sanchez, J.A., Apone, F., Abdel- Salam, E.M., Qahtan, A.A., et al. 2018. Production of plant secondary metabolites: examples, tips and suggestions for biotechnologists. Genes 9(6): 309. https://doi.org/10.3390/ genes9060309 Haider, S.-A., Ahmad, S., Sattar Khan, A., Anjum, M.A., Nasir, M. and Naz, S., 2020. Effects of salicylic acid on postharvest fruit quality of “Kinnow” mandarin under cold storage. Scientia Horticulturae 259: 108843. https://doi.org/10.1016/j.scienta.2019.108843 Haminiuk, C.W.I., Maciel, G.M., Plata-Oviedo, M.S.V. and Peralta,  R.M., 2012. Phenolic compounds in fruits – an over- view. International Journal of Food Science and Technology 47: 2023–2044. https://doi.org/10.1111/j.1365-2621.2012.03067.x Herrera-Vásquez, A., Salinas, P. and Holuigue, L., 2015. Salicylic acid and reactive oxygen species interplay in the transcriptional control of defense genes expression. Frontiers in Plant Science 6: 171. https://doi.org/10.3389/fpls.2015.00171 Higuchi, O., Tateshita, K. and Nishimura, H., 2003. Antioxidative activity of sulfur-containing compounds in allium species for human low-density lipoprotein (LDL) oxidation in vitro. Journal of Agricultural and Food Chemistry 51(24): 7208–7214. https:// doi.org/10.1021/jf034294u Ibrahim, M.H., Jaafar, H.Z.E., Rahmat, A. and Rahman, Z.A., 2012. Involvement of nitrogen on flavonoids, glutathione, antho- cyanin, ascorbic acid and antioxidant activities of Malaysian medicinal plant Labisia pumila Blume (Kacip Fatimah). International Journal of Molecular Sciences 13(1): 393–408. https://doi.org/10.3390/ijms13010393 Javanmardi, J. and Akbari, N., 2016. Salicylic acid at different plant growth stages affects secondary metabolites and phisico-chemi- cal parameters of greenhouse tomato. Advances in Horticultural Science 30(3): 151–157. https://oaj.fupress.net/index.php/ahs/ article/view/3063 Jenks, M.A., 2007. Plant abiotic stress edited by. In Stress The International Journal on the Biology of Stress 43(2). Johnson, E.J., 2002. The role of carotenoids in human health. Nutrition in Clinical Care: An Official Publication of Tufts University 5(2): 56–65. https://doi.org/10.1046/j.1523-5408.2002.00004.x Karalija, E. and Parić, A., 2017. Effects of salicylic acid foliar applica- tion on growth and antioxidant potential of basil (Ocimum basi- licum L.). Biologica Nyssana 8: 145–150. https://www.cabdirect. org/cabdirect/abstract/20183136195 Kdhim, S., J., Jassim, E., H., Salih, H., H., 2020. Effect of salicylic acid on increasing of some phenolic acid and flavonoids in corian- drum sativum callus. Plant Archives 20(1): 3007–3014. Khalil, N., Fekry, M., Bishr, M., El-Zalabani, S. and Salama, O., 2018. Foliar spraying of salicylic acid induced accumulation of pheno- lics, increased radical scavenging activity and modified the com- position of the essential oil of water stressed Thymus vulgaris L. Plant Physiology and Biochemistry: PPB 123: 65–74. https://doi. org/10.1016/j.plaphy.2017.12.007 Khan, M.I.R., Iqbal, N., Masood, A., Per, T.S. and Khan, N.A., 2013. Salicylic acid alleviates adverse effects of heat stress on photo- synthesis through changes in proline production and ethylene formation. Plant Signaling and Behavior 8(11): e26374. https:// doi.org/10.4161/psb.26374 Khandaker, L., Akond, A.M. and Oba, S., 2011. Foliar application of salicylic acid improved the growth, yield and leaf ’s bioac- tive compounds in Red Amaranth (Amaranthus tricolor L.). Vegetable Crops Research Bulletin 74(1): 77–86. https://doi. org/10.2478/v10032-011-0006-6 Kulbat, K., 2016. The role of phenolic compounds in plant resis- tance. Biotechnology and Food Science 80(2): 97–108. https:// repozytorium.p.lodz.pl/handle/11652/1613 Kumari, A., Pandey, N. and Pandey-Rai, S., 2018. Exogenous sali- cylic acid-mediated modulation of arsenic stress tolerance with enhanced accumulation of secondary metabolites and improved size of glandular trichomes in Artemisia annua L. Protoplasma 255(1): 139–152. https://doi.org/10.1007/s00709-017-1136-6 Lee, Y.S., Ju, H.K., Kim, Y.J., Lim, T.-G., Uddin, M.R., Kim, Y.B., et al. 2013. Enhancement of anti-inflammatory activity of aloe vera adventitious root extracts through the alteration of primary and secondary metabolites via salicylic acid elicitation. PLoS One 8(12): e82479. https://doi.org/10.1371/journal.pone.0082479 Lefevere, H., Bauters, L., Gheysen, G., 2020. Salicylic Acid Biosynthesis in Plants. Frontiers in Plant Science 11. https://doi. org/10.3389/fpls.2020.00338. Lin, D., Xiao, M., Zhao, J., Li, Z., Xing, B., Li, X., et al. 2016. An overview of plant phenolic compounds and their importance in human nutrition and management of Type 2 diabetes. Molecules (Basel, Switzerland) 21(10): 1374. https://doi.org/10.3390/ molecules21101374 Mendoza, D., Cuaspud, O., Arias, J.P., Ruiz, O. and Arias, M., 2018. Effect of salicylic acid and methyl jasmonate in the production of phenolic compounds in plant cell suspension cultures of Thevetia peruviana. Biotechnology Reports 19: e00273. https:// doi.org/10.1016/j.btre.2018.e00273 Mendoza-Sánchez, M., Pérez-Ramírez, I.F., Wall-Medrano, A., Martinez-Gonzalez, A.I., Gallegos-Corona, M.A. and Rosalía Reynoso-Camacho, R., 2019. Chemically induced common bean (Phaseolus vulgaris L.) sprouts ameliorate dyslipidemia by lipid intestinal absorption inhibition. Journal of Functional Foods 52: 54–62. https://www.sciencedirect.com/science/article/abs/pii/ S1756464618305504 Monireh, R., Hossein, L.Y. and Sheidaboroumand, J., 2011. The effect of salicylic acid on photosynthetic pigments, contents of sugar and antioxidant enzymes under lead stress in Brassica https://doi.org/10.3389/fpls.2016.01447� https://doi.org/10.5897/AJB2016.15320� https://doi.org/10.5897/AJB2016.15320� https://doi.org/10.3390/molecules23061296� https://doi.org/10.3390/genes9060309� https://doi.org/10.3390/genes9060309� https://doi.org/10.1016/j.scienta.2019.108843� https://doi.org/10.1111/j.1365-2621.2012.03067.x� https://doi.org/10.3389/fpls.2015.00171� https://doi.org/10.1021/jf034294u� https://doi.org/10.1021/jf034294u� https://doi.org/10.3390/ijms13010393� https://oaj.fupress.net/index.php/ahs/article/view/3063� https://oaj.fupress.net/index.php/ahs/article/view/3063� https://doi.org/10.1046/j.1523-5408.2002.00004.x� https://www.cabdirect.org/cabdirect/abstract/20183136195� https://www.cabdirect.org/cabdirect/abstract/20183136195� https://doi.org/10.1016/j.plaphy.2017.12.007� https://doi.org/10.1016/j.plaphy.2017.12.007� https://doi.org/10.4161/psb.26374� https://doi.org/10.4161/psb.26374� https://doi.org/10.2478/v10032-011-0006-6� https://doi.org/10.2478/v10032-011-0006-6� https://repozytorium.p.lodz.pl/handle/11652/1613� https://repozytorium.p.lodz.pl/handle/11652/1613� https://doi.org/10.1007/s00709-017-1136-6� https://doi.org/10.1371/journal.pone.0082479� https://doi.org/10.3389/fpls.2020.00338� https://doi.org/10.3389/fpls.2020.00338� https://doi.org/10.3390/molecules21101374� https://doi.org/10.3390/molecules21101374� https://doi.org/10.1016/j.btre.2018.e00273� https://doi.org/10.1016/j.btre.2018.e00273� https://www.sciencedirect.com/science/article/abs/pii/S1756464618305504� https://www.sciencedirect.com/science/article/abs/pii/S1756464618305504� 90 Italian Journal of Food Science, 2023; 35 (1) Ramos-Sotelo H et al. Napus L. Iranian Journal of Plant Biolology 3(9): 39–52. https:// ijpb.ui.ac.ir/article_18820.html?lang=en Osama, S., 2019. Effect of drought stress and salicylic acid on active constituents of Ammi visnaga L. Pharmacognosy Department. Faculty of Pharmacy, Cairo University. Pérez-Balibrea, S., Moreno, D.A. and García-Viguera, C., 2011. Improving the phytochemical composition of broccoli sprouts by elicitation. Food Chemistry 129(1): 35–44. https://doi. org/10.1016/j.foodchem.2011.03.049 Puthusseri, B., Divya, P., Lokesh, V. and Neelwarne, B., 2012. Enhancement of folate content and its stability using food grade elicitors in coriander (Coriandrum sativum L.). Plant Foods for Human Nutrition (Dordrecht, Netherlands) 67(2): 162–170. https://doi.org/10.1007/s11130-012-0285-1 Qian, P., Zhang, Y.B., Yang, Y.F., Xu, W. and Yang, X.W., 2017. Pharmacokinetics studies of 12 alkaloids in rat plasma after oral administration of Zuojin and Fan – Zuojin formulas. Molecules 22(2): 214. https://doi.org/10.3390/molecules22020214 Qing-Yan, G., Jiao, J., Wang, X., Yu-Ping, Z., Li-Li, N. and Yu-Jie, F., 2019. Elicitation of Isatis tinctoria L. hairy root cultures by sal- icylic acid and methyl jasmonate for the enhanced production of pharmacologically active alkaloids and flavonoids. Plant Cell, Tissue and Organ Culture 137: 417. https://doi.org/10.1007/ s11240-019-01610-w Rodas-Junco, B.A., Cab-Guillén, Y., Muñoz-Sánchez, J.A., Vázquez-Flota, F., Monforte-González, M. and Hernández- Sotomayor,  S.M., 2013. Salicylic acid induces vanillin synthesis through the phospholipid signaling pathway in Capsicum chin- ense cell cultures. Plant Signaling and Behavior 8(10): e26752. https://doi.org/10.4161/psb.26752 Rodas-Junco, B.A., Muñoz-Sánchez, J.A., Vázquez-Flota, F. and Hernández-Sotomayor, S.M.T., 2015. Salicylic-acid elicited phospholipase D responses in Capsicum chinense cell cul- tures. Plant Physiology and Biochemistry 90: 32–37. https://doi. org/10.1016/j.plaphy.2015.02.022 Ruelland, E., Pokotylo, I., Djafi, N., Cantrel, C., Repellin, A. and Zachowski, A., 2014. Salicylic acid modulates levels of phos- phoinositide dependent-phospholipase C substrates and products to remodel the Arabidopsis suspension cell transcrip- tome. Frontiers in Plant Science 5. https://doi.org/10.3389/ fpls.2014.00608 Singh, A. and Dwivedi, P., 2018. Methyl-jasmonate and salicylic acid as potent elicitors for secondary metabolite production in medicinal plants: a review. Journal of Pharmacognosy and Phytochemistry 7(1): 750–757. https://www.phytojournal. com/archives/2018.v7.i1.2574/methyl-jasmonate-and-salicylic- acid-as-potent-elicitors-for-secondary-metabolite-production- in-medicinal-plants-a-review Singh, B. and Sharma, R.A., 2015. Plant terpenes: defense responses, phylogenetic analysis, regulation and clinical appli- cations. 3 Biotech 5(2): 129–151. https://doi.org/10.1007/ s13205-014-0220-2 Skrypnik, L., Golovin, A., Savina, T., 2022. Effect of salicylic acid on phenolic compounds, antioxidant and antihyperglycemic activity of Lamiaceae plants grown in a temperate climate. Frontiers in Bioscience-Elite 14(1): 3. https://doi.org/10.31083/j. fbe1401003 Tajik, S., Zarinkamar, F. and Niknam, V., 2015. Effects of salicylic acid on carotenoids and antioxidant activity of saffron (Crocus sativus L.). Applied Food Biotechnology 2(4): 33–37. https:// journals.sbmu.ac.ir/afb/article/view/9739 Tsao, R., 2010. Chemistry and Biochemistry of Dietary Polyphenols. Nutrients 2(12): 1231–1246. https://doi.org/10.3390/ nu2121231 Vazquez, K. E., Flores-Cordova, M. A., Soto-Parra, J. M., Sánchez,  E., Soto-Caballero, M.C., Salas-Salazar, N.A., et al. 2022. Antioxidant activity and bio compounds induced by sali- cylic acid and potassium from ‘Flame’ grapes. Notulae Botanicae Horti Agrobotanici Cluj-Napoca 50(2): 12756. https://doi. org/10.15835/nbha50212756. Vicente, M.R. and Plasencia, J., 2011. Salicylic acid beyond defence: its role in plant growth and development. Journal of Experimental Botany 62(10): 3321–3338. https://doi. org/10.1093/jxb/err031 Vig, A.P., Rampal, G., Thind, T.S. and Arora, S., 2009. Bio-protective effects of glucosinolates – a review. LWT – Food Science and Technology 42(10): 1561–1572. https://doi.org/10.1016/j. lwt.2009.05.023 Vuolo, M.M., Lima, V.S. and Maróstica Junior, M.R., 2019. Chapter 2 – phenolic compounds: structure, classification, and antioxi- dant power. In: Campos (ed.) Bioactive compounds. Woodhead Publishing, pp. 33–50. Available at: https://app.dimensions.ai/ details/publication/pub.1111365242 Zhang, L., Du, L. and Poovaiah, B.W., 2014. Calcium signaling and biotic defense responses in plants. Plant Signaling and Behavior 9(11): e973818. https://doi.org/10.4161/15592324.2014.973818 Ziegler, J. and Facchini, P.J., 2007. Alkaloid biosynthesis: metabolism and trafficking. Annual Review of Plant Biology 59: 735–769. https://doi.org/10.1146/annurev.arplant.59.032607.092730 https://ijpb.ui.ac.ir/article_18820.html?lang=en� https://ijpb.ui.ac.ir/article_18820.html?lang=en� https://doi.org/10.1016/j.foodchem.2011.03.049� https://doi.org/10.1016/j.foodchem.2011.03.049� https://doi.org/10.1007/s11130-012-0285-1� https://doi.org/10.3390/molecules22020214� https://doi.org/10.1007/s11240-019-01610-w� https://doi.org/10.1007/s11240-019-01610-w� https://doi.org/10.4161/psb.26752� https://doi.org/10.1016/j.plaphy.2015.02.022� https://doi.org/10.1016/j.plaphy.2015.02.022� https://doi.org/10.3389/fpls.2014.00608� https://doi.org/10.3389/fpls.2014.00608� https://www.phytojournal.com/archives/2018.v7.i1.2574/methyl-jasmonate-and-salicylic-acid-as-potent-elicitors-for-secondary-metabolite-production-in-medicinal-plants-a-review� https://www.phytojournal.com/archives/2018.v7.i1.2574/methyl-jasmonate-and-salicylic-acid-as-potent-elicitors-for-secondary-metabolite-production-in-medicinal-plants-a-review� https://www.phytojournal.com/archives/2018.v7.i1.2574/methyl-jasmonate-and-salicylic-acid-as-potent-elicitors-for-secondary-metabolite-production-in-medicinal-plants-a-review� https://www.phytojournal.com/archives/2018.v7.i1.2574/methyl-jasmonate-and-salicylic-acid-as-potent-elicitors-for-secondary-metabolite-production-in-medicinal-plants-a-review� https://doi.org/10.1007/s13205-014-0220-2� https://doi.org/10.1007/s13205-014-0220-2� https://doi.org/10.31083/j.fbe1401003 https://doi.org/10.31083/j.fbe1401003 https://journals.sbmu.ac.ir/afb/article/view/9739� https://journals.sbmu.ac.ir/afb/article/view/9739� https://doi.org/10.3390/nu2121231� https://doi.org/10.3390/nu2121231� https://doi.org/10.15835/nbha50212756� https://doi.org/10.15835/nbha50212756� https://doi.org/10.1093/jxb/err031� https://doi.org/10.1093/jxb/err031� https://doi.org/10.1016/j.lwt.2009.05.023� https://doi.org/10.1016/j.lwt.2009.05.023� https://app.dimensions.ai/details/publication/pub.1111365242� https://app.dimensions.ai/details/publication/pub.1111365242� https://doi.org/10.4161/15592324.2014.973818� https://doi.org/10.1146/annurev.arplant.59.032607.092730� _gjdgxs _Hlk44419703 _Hlk44420993 _30j0zll