What Is the Use of Plant Melatonin?

N ° de catalogue ° de cA A A A A A atalogue ° de catalogue ° de cA A A A A A A A atalogueelatL L L L L L L L L L L L L L lur leduns, P P P P P ProdusL lur le chimiquesly known En tunt que N-acetyl-5-methoxytryptamdunse, wEn tunt que first dI Il estcovered duns Le cL lur leseil des mdansIl esttres pdanseal glEt en plus tIl estsue D D D D D D D D D D D D D D D D De la cows dans 1958. It belL lur legs À propos de Le cL lur leseil des mdansistres dansdole derivatives De la trypÀ propos dephan. Due À propos de sL lur le ability À propos de lighten Le cSur leseil des mdansistres skdans colOu bien De la certadans amphibians Et en plus fish, it is named melaÀ propos dendans [1].Initially, researchers believed that melaÀ propos dendans wEn tant que an active substance exclusive À propos de animals. H H H H H H H H H However, dans 1993, melaÀ propos dendans wEn tant que detected dans J J J J J J J J J J J J J J J J J J J J J J J J J J J J Japanese morndansg glory (P P Pharbitis nil) voir aussi: dans Japan, confirmdansg son presence dans Les planteeeeeeeeeeeeeeees. L lubsequently, melaÀ propos dendans wEn tant que quantified dans various plant species [2-7].

L lith Le conseil des mdansistres discovery De la Le conseil des mdansistres first melaÀ propos dendans récepteur dans Les plantes (C C C C C CA AND2/PN ° de catalogueT T T T T TR R R R R R R R R R R R R R1) voir aussi: Et en plus son physiologique fonctions, such as promotdansg plant La croissance, root Le développement, anti-agdansg, Et en plus sÀ propos dematal closure, melaÀ propos dendans has also been recognized as a Nouveau: type De la plant hormone [8,9]. L ldansce Le conseil des mdansistres discovery De la melaÀ propos dendans dans Les plantes, research on plant melaÀ propos dendans has primarily focused on endogène Contenu du site, biosynLe conseil des mdansistrestic parcourss, Et en plus physiological fonctions. In light De la this, this review summarizes Le conseil des mdansistres current État d’avancement De la research on plant melaÀ propos dendans À partir de Le conseil des mdansistres three aspects mentioned above, aimdansg to provide certadanss reference pour Le conseil des mdansistres application De la melatondans dans plant La La La production practices.

1. N ° de catalogueelatondans content dans Les plantes

Endogenous melatondans niveaux dans Les plantes are gènerally hauteer than those dans animals. On one hEt en plus, Les plantes encounter various adverse environmental conditions pendant La croissance, Et en plus at this time, Le conseil des mdansistres plant body requires numerous bioactive substances, danscluddansg melatondans, to enhance Le conseil des mdansistres plant' L l tolérance to Le Le Le Le Le Le Le Le stress À travers physiological réglementation mécanismes. On Le conseil des mdansistres oLe conseil des mdansistresr hEt en plus, Les plantes can contdansuously produce melatondans precursors, such as tryptophan, À travers Le conseil des mdansistres shikimic acideeeeeee parcours.while animals lack this metabolic parcours Et en plus must obtadans melatondans À partir de Le conseil des mdansistresir diet [10-12].

For Le conseil des mdansistres quantitative detection De la melatondans content dans Les plantes, commonly used methods dansclude radioimmunoassay (RIA), haute-perpourmance liquideee chromatography-electrochemilumdansescence detection (HPLC-ECD) Analyse des données, haute-perpourmance liquid chromatography-fluorescence detection (HPLC-FD) Analyse des données, high-perpourmance liquid chromatography-mass spectrometry (HPLC-N ° de catalogueL l) Analyse des données,gas chromatography-mass spectrometry (G GC-N ° de catalogueL l) Analyse des données, Et en plus enzyme-ldansked immunosorbent assay (ELIL lA), among oLe conseil des mdansistresrs. B B Based on Le conseil des mdansistresse detection methods, melatondans has been found to be widely distributed dans various plant organs such as Les racdanses, stems, flowers, feuilles, seeds, Et en plus fruson; however, son content exhibson significant danster- Et en plus danstra-species variations, Et en plus even différent tissue parts De la Le conseil des mdansistres same plant may show distdansct differences (Table 1).

Among Les plantes where melatondans has been quantitatively detected, some médicdansal Les plantes exposer relatively high melatondans niveaux [18,22]. Additionally, compared to cucumber (Cucurbitaceae), kiwifruitss (Actdansidiaceae), strawberry (Rosaceae), onion (Alliaceae), Et en plus garlic (Alliaceae),melatondans niveaux dans Poaceae plants such as riz, barley, sweet corn, oats, Et en plus tall fescue are higher [3]. L lang Jdansydansg et al. [23] used HPLC to determdanse melatondans content dans 132 corn Et en plus 145 riz seed samples, avec results showdansg thatmelatondans was detected dans 58 corn Et en plus 25 riz samples (≥10 ng/g), avec ranges De la 10–2034 ng/g Et en plus 11–264 ng/g, respectively.

N ° de catalogueelatondans was found to be present at higher niveaux dans Le conseil des mdansistres flower bud tissues De la white mEt en plusrake, but its content dimdansué avec Le conseil des ministres development De la floral organs [24]. L ltudies on Le conseil des ministres medicinal plant licoriz revealed that melatonin was present at Le conseil des ministres highest niveaux in its root tissues, avec its concentration increasing avec Le conseil des ministres plant' L l developmental stage. Concurrently,melatonin niveaux in root tissues were highest after 3 days De la high-intensity (1.13 L l/m²) UV VB radiation Et en plus second highest after 15 days De la low-intensity (0.43 L l/m²) radiation Le traitement [25]. Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Ye et al. [26] reported that after 15 days De la treatment avec 500 μmol/L cadmium, melatonin levels in riz stems,Les racines reached 21.0 ng/g Et en plus 3.0 ng/g, respectively, which were 10 times Et en plus 3 times higher than those in Le conseil des ministres control treatment. The above data indicate that melatonin levels in plants are closely liés to factors such as Le conseil des ministres plant's own gènetic characteristics, environmental conditions, Et en plus La croissance Et en plus development stages.

2. L lynLe conseil des ministressis De la plant melatonin

2.1 N ° de catalogueelatonin L lynthesis Pathway

The elucidation De la the melatonin synthèse parcours in plants is a hot research area in melatonin studies. N ° de catalogueurch et al. [27] were the first to investigate the melatonin synthèse pathway in plants. They detected indoleacetic acid (IAA), tryptophan (TAN ° de catalogue), 5-hydroxytryptophan (5-HTP), Et en plus 5-hydroxytryptamine (5-HT), which are intermediate products required in the melatonin biosynthèse process in animals. Based on these findings, it was speculated that plants Et en plus animals may share similar melatonin synthèse Les voies d’accès.However, subsequent experimental studies En utilisant moléculaire biology Et en plus enzyme-catalyzed reaction kinetics methods revealed that the melatonin synthesis pathway in plants is far more complex than that in animals, Et en plus significant differences exist between the deux (Figures 1 Et en plus 2). This suggests that the mechanisms underlying melatonin synthesis may have evolved differently in plants Et en plus animals.

Based on current research findings, in the melatonin synthesis pathway De la plants,the conversion De la tryptophan to 5-hydroxytryptophan is uncontested, while the final synthesis De la melatonin from 5-hydroxytryptophan remains controversial. Specifically, 5-hydroxytryptophan is first acetylated to pourm N-acetyl-5-hydroxytryptophan,which is then catalyzed Par: a methyltransferase to ultimately synthesize melatonin (referred to as the NN ° de catalogue pathway, similar to the melatonin synthesis pathway in animals) or whether 5-hydroxytryptophan is first methylated to form 5-methoxytryptophan, which is then acetylated to produce melatonin (referred to as the N ° de catalogueN pathway) (Figure 2) [10].

In the melatonin synthesis process De la plants, the NN ° de catalogue Et en plus N ° de catalogueN pathways may coexist in parallel. Recent studies have shown that under normal La croissance conditions, melatonin synthesis in plants is dominated Par: the NN ° de catalogue pathway, while under non-biotic stress conditions, the N ° de catalogueN pathway is predominant [28]. This may be related to the presence De la multiple SNAT Et en plus ASN ° de catalogueT subtypes in plants.

2.2 Regulation De la melatonin synthesis

La recherche on the réglementation De la melatonin biosynthèse in plants is still in its infancy, Et en plus the underlying mechanisms remain poorly understood. K K K K Kolár et al. [29] conducted early studies on the temporal patterns De la melatonin levels in the short-day plant red-feuille amaranth, revealing that melatonin levels exposered circadian rhythms similar to those observed in animals, suggesting that light inhibits melatonin biosynthèse in plants.However, the findings De la two research teams led Par: N ° de catalogueurch [27] Et en plus Tan [30] indicate that light does not inhibit melatonin biosynthèse in plants, Et en plus that the rate De la melatonin biosynthèse is positively correlated avec light intensity. In addition to light intensity, non-biotic stress factors such as light wavelength, high temperature, low temperature, drought, high salinity, lead, Et en plus cadmium are also major regulatory factors for melatonin biosynthèse in plants [31-34].

2.3 Sites De la melatonin biosynthèse

N ° de catalogueelatonin, as a hydrophilic Et en plus lipophilic small molecule, can freely transport between tissue celluleuleules; on the other hEt en plus, the biosynthèse De la plant melatonin is significantly influenced Par: external environmental conditions, which Les augmentations the difficulty De la locating the sites De la melatonin biosynthèse. Since chloroplasts Et en plus mitochondria are the primary sites for reactive oxygen species (ROS) production [35,36], it is speculated that chloroplasts Et en plus mitochondria may be the primary sites for melatonin biosynthesis in plants.

The findings De la Byeon et al. [37] provided preliminary evidence supporting this hypothesis. They discovered that SNAT, one De la the Clé: Les enzymes in the melatonin biosynthetic pathway, is localized in chloroplasts, while ASN ° de catalogueT is present in the cytoplasm.Further studies revealed that specific overExpression: De la COMT in chloroplasts significantly augmenté endogenous melatonin levels, whereas overExpression: De la COMT in the cytoplasm did not cause significant Les changements in melatonin levels [38], indicating that the synergistic action De la COMT Et en plus SNAT in chloroplasts plays a crucial Rôle de la commission in plant melatonin biosynthesis.

The most compelling evidence supporting the Rôle de la commission De la chloroplasts as the site De la melatonin biosynthesis in plants comes from the findings De la Zheng et al. [39], who added 5-hydroxytryptophan to isolated Et en plus purified apple chloroplasts, resulting in melatonin production in a dose-dépendante manner.Recent subcellulaire localization experiments have further confirmed that chloroplasts may be the site De la melatonin biosynthesis [40,41].In addition to chloroplasts, mitochondria have also been preliminarily confirmed Par: experimental results as sites De la melatonin synthesis in plants. Wang et al. [42] found that isolated apple mitochondria can produce melatonin, Et en plus the apple SNAT subtype MzSNAT5 was localized in the mitochondria De la Arabidopsis protoplasts Et en plus apple callus cells.

3. Physiological fonctions De la plant melatonin

On the one hEt en plus, melatonin itself possesses strong reducing Capacité d’accueil, capable De la dépouillement free radicals gènerated within plants Et en plus maintaining the metabolic balance De la ROS within cells; on the other hEt en plus, tryptophan serves as a common precursor for the biosynthesis De la melatonin Et en plus IAA, Et en plus both play similar regulatory roles in plant La croissance Et en plus development [43-45]. Previous studies have shown that melatonin, as an important signalisation molecule Et en plus antioxydant, participates in La réglementation plant La croissance Et en plus development as well as responses to various environmental stresses.

3.1 Melatonin réglementation De la plant La croissance Et en plus development

Murch et al. [46,47] found in early studies that changes in endogenous melatonin concentrations in plants affect root development, cell mitosis, Et en plus the formation De la mitotic spindles. Based on this finding, they proposed the hypothesis that melatonin is a potentiel plant La croissance regulator.Subsequent experiments have confirmed this hypothesis, showing that melatonin is widely involved in regulating plant flowering, fruit ripening, photosynthesis, feuille senescence, root morphology, Et en plus other La croissance Et en plus development processes [35], with its Les effets similar to or synergistic with those De la IAA.Melatonin promotes the elongation De la the hypocotyl in lupine, with an activity equivalent to 63% De la IAA [20]. In a similar study, it was found that melatonin promotes the growth De la the coleoptile sheath in monocotyledonous plants such as oats, blé, barley, Et en plus chickweed,with an activity De la 10% (oats) to 55% (barley) De la IAA.

Additionally, similar to IAA, melatonin exhibits concentration-dependent inhibitory effets on root growth in the aforementioned plants, with an inhibitory effet De la 56% (chickweed) to 86% (wheat) De la IAA [16].

Studies have shown that melatonin at concentrations De la 10⁻⁹ to 10⁻⁶ mol·L⁻¹ can act as an IAA analog to promote the growth De la the primary root system De la Arabidopsis.Further Analyse des données revealed that melatonin Et en plus IAA treatment-induit gène expression changes were moderately correlated, Et en plus most gènes regulated Par: IAA were also regulated Par: melatonin. This suggests that melatonin Et en plus IAA co-regulate a similar subset De la gènes, leading to the inference that melatonin promotes Arabidopsis primary root growth in an IAA-dependent manner [48].In riz, overexpression De la the sheep 5-hydroxytryptophan-N-acetyltransferase gene resulted in enhanced root development Et en plus increased adventitious root numbers in transgénique plants. Additionally, exogenous melatonin treatment promoted root growth in sauvage-type riz plants under continuous light conditions [49].

Furthermore, studies in Brassica rapa [50], cherry [51], sunflower [52], La tomate [53], Et en plus some monocotyledonous plants [16,54] have shown that melatonin's regulatory effet on plant growth Et en plus development is concentration-dependent.At a concentration De la 0.1 μmol.L-1, melatonin promotes root growth in rapeseed, while at 100 μmol.L-1, it exhibits an inhibitory effet. Concurrently, low-concentration melatonin treatment leads to an increase in endogenous IAA levels,suggesting that the promoting effect De la low melatonin concentrations is related to the changes in endogenous IAA levels [50]. Hernández Ruiz et al. [16] found that the optimal melatonin concentration for promoting root growth in monocotyledonous plants such as oats, wheat, barley, Et en plus chickweed was 10⁻⁷ mol/L.In riz, melatonin treatment at concentrations De la 10–50 μmol.L-1 inhibited hypocotyl growth but promoted lateral root formation Et en plus development [54].

Coating Le soja seeds with a coating agent containing 50 or 100 μmol.L-1 melatonin significantly promoted plant growth Et en plus development,increased soybean yield Et en plus gras acid content [55]. Zhong et al. [56] found that exogenous melatonin treatment promoted the growth Et en plus development De la grape seedlings Par: enhancing the photosynthetic performance De la feuille blades Et en plus increasing plant biomass. Additionally, exogenous melatonin treatment also increased the yield De la corn, mung beans, Et en plus cucumbers [57–59].Melatonin primarily exerts its fonctions Par: regulating the transcription De la numerous gènes involved in cell division, photosynthesis, carbohydrate métabolisme, fatty acid biosynthesis, Et en plus ascorbic acid métabolisme [60].

Park et al. [61] measured melatonin levels in three different growth stages De la riz—pre-flowering, flowering, Et en plus post-flowering—Et en plus found that melatonin levels in the panicle (flower) were six times higher than those in the flag feuille, suggesting that melatonin may be involved in the development De la floral organs.Studies on the Les effets De la exogenous melatonin on fruit ripening have primarily focused on the interaction between melatonin Et en plus ethylene. Early studies found that after pre-treatment with 50 μmol.L-1 melatonin, parameters related to fruit ripening, such as lycopene levels, fruit sDe latening degree, Et en plus enzymes associés with ethylene signalisation Et en plus biosynthesis, showed significant changes in La tomatees.corresponding proteomics analysis indicated that exogenous melatonin treatment increased the abundance De la protéiness associated with fruit ripening-related pathways Et en plus l’anthocyane l’l’accumulation pathways [62,63].Additionally, exogenous melatonin treatment reduced the weight loss rate Et en plus rot rate De la peach fruits while maintaining fruit firmness, soluble solids content, Et en plus ascorbic acid levels, therePar: effectively delaying the senescence Et en plus decay De la peach fruits from two different genetic backgrounds [64]. Melatonin treatment pendant fruit ripening increased soluble Le sucre content Et en plus single fruit weight in pear fruits [65].

plante feuille senescence is a programmed form De la cell death primarily leading to the dégradation De la macromolecules, including chlorophylle [66]. Under drought conditions, exogenous melatonin treatment can inhibit the expression De la apple senescence-related gene 12 (SAG12) Et en plus the polyphenol oxidase gene (PAO) [67].Melatonin pretreatment significantly slows down the aging process De la barley leaves, with the highest chlorophyll content observed in leaves treated with 1 mmol/L melatonin [68]. Liang et al. [69] found in rice that melatonin delays feuille aging Par: inhibiting chlorophyll degradation Et en plus the expression De la aging-related genes.Proteomics analysis revealed that the expression levels De la aging-related protéiness were reduced after melatonin pretreatment [57]. In perennial ryegrass, exogenous melatonin treatment inhibited the transcription De la aging-related genes LpSAG12.1 Et en plus Lph36, therePar: delaying high-temperature stress-induced leaf senescence [70].

In addition to delaying leaf senescence, melatonin appears to enhance plant photosynthetic efficiency through an unconventional biostimulatory pathway. Long-term application De la 100 μmol. L-1 melatonin to growth soil improved the photochemical efficiency De la photosystem II in apples under weak light conditions, allePar l’intermédiaire deted drought stress-induced l’l’inhibition De la photosynthesis,while maintaining higher CO₂ assimilation Capacité d’accueil Et en plus stomatal conductance in plant leaves [71]. Pretreatment with 0.1 mmol/L melatonin increased net photosynthetic rate, transpiration rate, stomatal conductance, photosystem II quantum efficiency, electron transport rate, and maximum photochemical efficiency (Fv/Fm) in La tomate plants [72].

3.2 Role De la plant melatonin in responses to biotic stress

Melatonin significantly enhances plant tolérance to biotic stress. Given its État d’avancement as an environmentally friendly molecule, it is considered the most economical and green alternative for inducing plants to resist biotic stress.Exogenous melatonin exhibits certain effects contre fungal-induced diseases. Treatment with melatonin at concentrations De la 0.05–0.5 mmol/L can enhance resistance to brown spot disease Par: regulating the activity De la antioxydant and défense-related enzymes in apples [73]. Melatonin can also mitigate damage caused by fungal infections to crops such as potatoes, cotton, and white lupins [74-76]. Different concentrations De la exogenous melatonin can inhibit the growth De la fungal pathogens such as Botrytis, Fusarium, and Fusarium [76].

In terms De la its mechanism De la action, melatonin primarily helps plants resist fungal infections, reduce lesions, and inhibit pathogen spread, ultimately mitigating the damage caused by diseases. Arabidopsis thaliana-Pseudomonas syringae La tomate pathogenic strain DC3000 (Pst DC3000) is the most widely used model in studies De la plant-pathogen interactions [77].

Exogenous melatonin pretreatment at certain concentrations enhances the resistance De la Arabidopsis and tobacco to Pst DC3000 [78,79]. Lee et al. [80] found that inactivation De la sérotonine-N-acetyltransferase significantly reduced endogenous melatonin levels in Arabidopsis,resulting in increased susceptibility De la plants to Pst DC3000. Therefore, melatonin can enhance plant tolérance to bactérienne diseases.

Compared to fungal and bacterial diseases, viral infections in plants are more difficult to control once they occur. Zhao et al. [81] first investigated the role De la melatonin in plant-virus interactions, finding that exogenous melatonin application significantly inhibited viral infection De la tobacco seedlings. Additionally, exogenous melatonin pretreatment reduced the incidence De la viral diseases in rice [82].With the deepening De la research on melatonin-médiatisé plant disease resistance mechanisms, melatonin could provide new strategies for the prevention and control De la plant viral diseases.

In addition to phenotypic identification, recent analyses De la gene expression have provided strong evidence for melatonin's implication in regulating plant responses to abiotic stress.Exogenous melatonin treatment or overexpression De la melatonin synthesis-related genes can induce the expression De la disease-related (PR) genes such as PR1, PR5, NPR1, and PDF1.2, as well as activate mitogen-activated protein kinases (MAPKs) and other disease-resistant proteins,thereby enhancing plant disease resistance, indicating that melatonin is an efficient défense agent contre pathogens in plants [78,83-85].

In addition to pathogens, insect pests are another major biotic stress that plants face pendant growth. plante-derived secondary metabolites act as antagonists De la insect juvenile hormone to defend contre insect predation [86].It has been reported that dopamine, which has a similar structure to melatonin, plays an important role in plant défense contre herbivores [87], suggesting that melatonin may also exert similar defensive effects [88].

3.3 Role De la plant melatonin in responses to abiotic stress

Under normal growth conditions, the production and removal De la ROS in plant cells are in dynamic equilibrium. When plants are exposed to adverse environmental factors such as drought, high salinity, extreme temperatures, heavy metals, and ultraviolet radiation, this equilibrium is disrupted, leading to oxydatif stress damage in plant cells [89-91].To eliminate ROS within the body, plants have evolved an efficient enzymatic and non-enzymatic antioxidant defense system to protect cells from or mitigate damage caused by oxydatif stress.

Melatonin is currently the strongest endogenous free radical scavenger with antioxidant activity. It is estimated that a single melatonin molecule can eliminate 10 free radicals through a cascade reaction,while classic antioxidants typically only eliminate one free radical per molecule [92]. Therefore, it is inferred that melatonin's primary function in organisms is as an antioxidant to eliminate various ROS and reactive nitrogen species (RNS), thereby protecting plants from oxydatif stress [92-95].

Melatonin may control the burst De la hydrogen Le peroxyde pendant non-biotic stress responses in plants by directly dépouillement excess ROS, enhancing antioxidant enzyme activity, and improving the ascorbic acid-glutathione (AsA-GSH) cycle capacity [67].Exogenous melatonin treatment enhances antioxidant enzyme activity in apples, grapes, corn, sunflowers, La tomatees, and wheat, while reducing the concentrations De la superoxide, hydrogen peroxide, and malondialdehyde [59,67,72,96-99].Additionally, exogenous melatonin treatment reduces the accumulation De la oxidized proteins in plants, accelerates the occurrence De la autophagy induced by oxidative stress, and allePar l’intermédiaire detes photooxidative damage [100].

4. Conclusion and Outlook

Since the discovery De la melatonin in plants, its diverse physiological fonctions and significant potentiel applications have attracted increasing attention in plant melatonin research. However, compared with other plant hormones, there are still many unresolved issues in plant melatonin research.For example, while the interactions between plant melatonin and other hormones have been studied directly or indirectly, the signal transduction models underlying these interactions remain unclear. Therefore, further investigation into the mechanisms by which melatonin coordinates with other plant hormones to regulate plant growth and development and responses to abiotic stress is a key direction for future research on plant melatonin.

By elucidating the interactions between melatonin and other hormones, we can also gain insights into the moléculaire mechanisms De la melatonin signaling and its role as a plant growth regulator. Additionally, how plants perceive melatonin signals and how downstream signal transduction regulates plant responses to stress at physiological and metabolic levels remain unresolved questions.However, the discovery De la Arabidopsis CAND2/PMTR1 as melatonin signal transduction récepteurs has laid a foundation for further research into melatonin signal transduction in plants [8].

Melatonin is widely distributed in various plant tissues; however, it remains unclear whether all plant organs possess melatonin synthesis capabilities. The mechanisms and pathways De la melatonin transport within plants require further investigation.The heterologous overexpression De la melatonin synthesis-related genes can significantly increase the endogenous melatonin levels in plants, while transgénique plants exhibit enhanced tolérance to various abiotic stresses [40,49,101-110].In the future, genetic engineering could serve as an important means to increase the endogenous melatonin content in crops, thereby promoting plant growth and development, enhancing tolérance to abiotic stress, and ultimately improving crop yields.

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