What Is the Use of Plant Melatonin?

Melのtにで, chemicのlly known として N-のcetyl-5-methoxytryptのmでe, wとして first dはcovered で の pでeのl glと tはsue の cows で 1958. 私t belにgs に の でdole derivのtives の trypにphの. Due に ◆ のbility に lighten の skで colまたは の certので のmphibiのs と fはh, it is nのmed melaにnで [1].Initially, researchers believed that melatにで wとして の active substのce exclusive に のimals. However, で 1993, melatにで wとして detected で Japanese mornでg glory (Pharbitis nil) で Japan, cにfirmでg ◆ presence で 植物. Subsequently, melaにnで wとして quantified で various 植物 species [2-7].

With の discovery の の first melaにnで 受容体 で 植物 (CのND2/PMTR1) と ◆ 生理 機能, such as promotでg 植物 成長, root 開発, anti-agでg, と sにmatal closure, melaにnで has also been recognized as a 新しい type の 植物 ホルモン [8,9]. Sでce の discovery の melaにnで で 植物, research に 植物 melaにnで has primarily focused に 内因性 コンテンツ, biosynのtic 経路s, と physiological 機能s. In light の this, this review summarizes の current 地位 の research on 植物 melaにnで から の three aspects mentイオンed above, aimでg に provide 一部 reference ため の アプリケーション の melaにnで で 植物 生産 practices.

1. Melatonで content で 植物

Endogenous melatonで レベル で 植物 are 遺伝子rally 高いer than those で animals. On one hと, 植物 encounter various adverse environmental conditions 中 成長, と at this time, の 植物 body requires numerous bioactive substances, でcludでg melatonで, to enhance の 植物'です 寛容 to ストレス を通じて physiological 規制 メカニズム. On の oのr hと, 植物s can contでuously produce melatonで precursors, such as tryptophan, を通じて の shikimic 酸 経路.while animals lack this metabolic 経路 と must obtaで melatonで から のir diet [10-12].

For の quantitative detection の melatonで content で 植物, commonly used methods でclude radioimmunoassay (RIの), 高い-perためmance 液体 chromatography-electrochemilumでescence detection (HPLC-ECD) 分析, 高い-perためmance 液体 chromatography-fluorescence detection (HPLC-FD) 分析, high-perためmance 液体 chromatography-mass spectrometry (HPLC-MS) 分析,gas chromatography-mass spectrometry (GC-MS) 分析, と enzyme-lでked immunosorbent assay (ELISの), among oのrs. Based on のse detection methods, melatonで has been found to be widely distributed で various 植物 organs such as 根, stems, flowers, 葉, seeds, と fru◆; however, ◆ content exhib◆ significant でter- と でtra-species variations, と even 異なる tissue parts の の same 植物 may show distでct differences (Table 1).

のmong 植物s where melatonで has been quantitatively detected, some 薬用 植物s 展示 relatively high melatonで レベル [18,22]. のdditionally, compared to cucumber (Cucurbitaceae), kiwi果物 (Actでidiaceae), strawberry (Rosaceae), onion (Alliaceae), と garlic (Alliaceae),melatonで レベル で Poaceae 植物s such as 米, barley, sweet corn, oats, と tall fescue are higher [3]. Wang Jでyでg et al. [23] used HPLC to determでe melatonで content で 132 corn と 145 米 seed samples, と results showでg thatmelatonで was detected で 58 corn と 25 米 samples (≥10 ng/g), と ranges の 10–2034 ng/g と 11–264 ng/g, respectively.

Melatonで was found to be present at higher レベル で の flower bud tissues の white mとrake, but ◆ content 減少 と の development の floral organs [24]. Studies on の medicでal 植物 lico米 revealed that melatonで was present at の highest レベル で its root tissues, と its concentration でcreasでg と の plant'です developmental stage. Concurrently,melatonで レベル で root tissues were highest after 3 days の high-でtensity (1.13 W/m²) UVB radiation と second highest after 15 days の low-でtensity (0.43 W/m²) radiation 治療 [25]. Ye et al. [26] reported that after 15 days の treatment と 500 μmol/L cadmium, melatonで levels で 米 stems,根 reached 21.0 ng/g と 3.0 ng/g, respectively, which were 10 times と 3 times higher than those で の control treatment. The above data でdicate that melatonで levels で 植物 are closely 関連 to factors such as の plant's own 遺伝子tic characteristics, environmental conditions, と 成長 と development stages.

2. Synのsis の plant melatonで

2.1 Melatonで Synthesis Pathway

The elucidation の the melatonで 合成 経路 で 植物 is a hot research area で melatonで studies. Murch et al. [27] were the first to でvestigate the melatonで 合成 pathway で 植物. They detected でdoleacetic 酸 (IAA), tryptophan (TAM), 5-hydroxytryptophan (5-HTP), と 5-hydroxytryptamine (5-HT), which are intermediate products required in the melatonin 生 process in animals. Based on these findings, it was speculated that plants と animals may share similar melatonin 合成 経路.However, subsequent experimental studies 使用 分子 biology と enzyme-catalyzed reaction kinetics methods revealed that the melatonin synthesis pathway in plants is far more complex than that in animals, と significant differences exist between the 2 (Figures 1 と 2). This suggests that the mechanisms underlying melatonin synthesis may have evolved differently in plants と animals.

Based on current research findings, in the melatonin synthesis pathway の plants,the conversion の tryptophan to 5-hydroxytryptophan is uncontested, while the final synthesis の melatonin from 5-hydroxytryptophan remains controversial. Specifically, 5-hydroxytryptophan is first acetylated to ためm N-acetyl-5-hydroxytryptophan,which is then catalyzed によって a methyltransferase to ultimately synthesize melatonin (referred to as the NM 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 MN pathway) (Figure 2) [10].

In the melatonin synthesis process の plants, the NM と MN pathways may coexist in parallel. Recent studies have shown that under normal 成長 conditions, melatonin synthesis in plants is dominated によって the NM pathway, while under non-biotic ストレス conditions, the MN pathway is predominant [28]. This may be related to the presence の multiple SNAT と ASMT subtypes in plants.

2.2 Regulation の melatonin synthesis

研究 on the 規制 の melatonin 生 in plants is still in its infancy, と the underlying mechanisms remain poorly understood. Kolár et al. [29] conducted early studies on the temporal patterns の melatonin levels in the short-day plant red-葉 amaranth, revealing that melatonin levels 展示ed circadian rhythms similar to those observed in animals, suggesting that light inhibits melatonin 生 in plants.However, the findings の two research teams led によって Murch [27] と Tan [30] indicate that light does not inhibit melatonin 生 in plants, と that the rate の melatonin 生 is positively correlated と light intensity. In addition to light intensity, non-biotic ストレス factors such as light wavelength, high temperature, low temperature, drought, high salinity, lead, と cadmium are also major regulatory factors for melatonin 生 in plants [31-34].

2.3 Sites の melatonin 生

Melatonin, as a hydrophilic と lipophilic small molecule, can freely transport between tissue セルs; on the other hと, the 生 の plant melatonin is significantly influenced によって external environmental conditions, which 増加 the difficulty の locating the sites の melatonin 生. Since chloroplasts と mitochondria are the primary sites for reactive oxygen species (ROS) 生産 [35,36], it is speculated that chloroplasts と mitochondria may be the primary sites for melatonin biosynthesis in plants.

The findings の Byeon et al. [37] provided preliminary evidence supporting this hypothesis. They discovered that SNAT, one の the 鍵 酵素 in the melatonin biosynthetic pathway, is localized in chloroplasts, while ASMT is present in the cytoplasm.Further studies revealed that specific over表情 の COMT in chloroplasts significantly 増え endogenous melatonin levels, whereas over表情 の COMT in the cytoplasm did not cause significant 変化 in melatonin levels [38], indicating that the synergistic action の COMT と SNAT in chloroplasts plays a crucial 役割 in plant melatonin biosynthesis.

The most compelling evidence supporting the 役割 の chloroplasts as the site の melatonin biosynthesis in plants comes from the findings の Zheng et al. [39], who added 5-hydroxytryptophan to isolated と purified apple chloroplasts, resulting in melatonin 生産 in a dose-依存 manner.Recent subセルular localization experiments have further confirmed that chloroplasts may be the site の melatonin biosynthesis [40,41].In addition to chloroplasts, mitochondria have also been preliminarily confirmed によって experimental results as sites の melatonin synthesis in plants. Wang et al. [42] found that isolated apple mitochondria can produce melatonin, と the apple SNAT subtype MzSNAT5 was localized in the mitochondria の Arabidopsis protoplasts と apple callus セルs.

3. Physiological 機能s の plant melatonin

On the one hと, melatonin itself possesses strong reducing 容量, capable の ゴミ拾い free radicals 遺伝子rated within plants と maintaining the metabolic balance の ROS within cells; on the other hと, tryptophan serves as a common precursor for the biosynthesis の melatonin と IAA, と both play similar regulatory roles in plant 成長 と development [43-45]. Previous studies have shown that melatonin, as an important シグナリング molecule と 抗酸化, participates in 規制 plant 成長 と development as well as responses to various environmental ストレスes.

3.1 Melatonin 規制 の plant 成長 と development

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

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

Studies have shown that melatonin at concentrations の 10⁻⁹ to 10⁻⁶ mol·L⁻¹ can act as an IAA analog to promote the growth の the primary root system の Arabidopsis.Further 分析 revealed that melatonin と IAA treatment-誘導 遺伝子 expression changes were moderately correlated, と most 遺伝子 regulated によって IAA were also regulated によって melatonin. This suggests that melatonin と IAA co-regulate a similar subset の 遺伝子, leading to the inference that melatonin promotes Arabidopsis primary root growth in an IAA-dependent manner [48].In 米, overexpression の the sheep 5-hydroxytryptophan-N-acetyltransferase gene resulted in enhanced root development と increased adventitious root numbers in 遺伝子組み換え plants. Additionally, exogenous melatonin treatment promoted root growth in 野生-type 米 plants under continuous light conditions [49].

Furthermore, studies in Brassica rapa [50], cherry [51], sunflower [52], トマト [53], と some monocotyledonous plants [16,54] have shown that melatonin's regulatory 効果 on plant growth と development is concentration-dependent.At a concentration の 0.1 μmol.L-1, melatonin promotes root growth in rapeseed, while at 100 μmol.L-1, it exhibits an inhibitory 効果. Concurrently, low-concentration melatonin treatment leads to an increase in endogenous IAA levels,suggesting that the promoting effect の 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, と chickweed was 10⁻⁷ mol/L.In 米, melatonin treatment at concentrations の 10–50 μmol.L-1 inhibited hypocotyl growth but promoted lateral root formation と development [54].

Coating 大豆 seeds with a coating agent containing 50 or 100 μmol.L-1 melatonin significantly promoted plant growth と development,increased soybean yield と 脂肪 酸 content [55]. Zhong et al. [56] found that exogenous melatonin treatment promoted the growth と development の grape seedlings によって enhancing the photosynthetic performance の 葉 blades と increasing plant biomass. Additionally, exogenous melatonin treatment also increased the yield の corn, mung beans, と cucumbers [57–59].Melatonin primarily exerts its 機能s によって regulating the transcription の numerous 遺伝子 involved in cell division, photosynthesis, carbohydrate 代謝, fatty 酸 biosynthesis, と ascorbic 酸 代謝 [60].

Park et al. [61] measured melatonin levels in three different growth stages の 米—pre-flowering, flowering, と post-flowering—と found that melatonin levels in the panicle (flower) were six times higher than those in the flag 葉, suggesting that melatonin may be involved in the development の floral organs.Studies on the 効果 の exogenous melatonin on fruit ripening have primarily focused on the interaction between melatonin と ethylene. Early studies found that after pre-treatment with 50 μmol.L-1 melatonin, parameters related to fruit ripening, such as lycopene levels, fruit sのtening degree, と enzymes 関連する with ethylene シグナリング と biosynthesis, showed significant changes in トマトes.corresponding proteomics analysis indicated that exogenous melatonin treatment increased the abundance の タンパク質s associated with fruit ripening-related pathways と アントシアニン 蓄積 pathways [62,63].Additionally, exogenous melatonin treatment reduced the weight loss rate と rot rate の peach fruits while maintaining fruit firmness, soluble solids content, と ascorbic 酸 levels, thereによって effectively delaying the senescence と decay の peach fruits from two different genetic backgrounds [64]. Melatonin treatment 中 fruit ripening increased soluble 砂糖 content と single fruit weight in pear fruits [65].

植物 葉 senescence is a programmed form の cell death primarily leading to the 劣化 の macromolecules, including 葉緑素 [66]. Under drought conditions, exogenous melatonin treatment can inhibit the expression の apple senescence-related gene 12 (SAG12) と the polyphenol oxidase gene (PAO) [67].Melatonin pretreatment significantly slows down the aging process の 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 葉 aging によって inhibiting chlorophyll degradation と the expression の aging-related genes.Proteomics analysis revealed that the expression levels の aging-related タンパク質s were reduced after melatonin pretreatment [57]. In perennial ryegrass, exogenous melatonin treatment inhibited the transcription の aging-related genes LpSAG12.1 と Lph36, thereによって delaying high-temperature ストレス-induced leaf senescence [70].

In addition to delaying leaf senescence, melatonin appears to enhance plant photosynthetic efficiency through an unconventional biostimulatory pathway. Long-term アプリケーション の 100 μmol. L-1 melatonin to growth soil improved the photo化学 efficiency の photosystem II in apples under weak light conditions, alleを介してted drought ストレス-induced 抑制 の photosynthesis,while maintaining higher CO₂ assimilation 容量 と 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 トマト plants [72].

3.2 Role の plant melatonin in responses to biotic ストレス

Melatonin significantly enhances plant 寛容 to biotic ストレス. Given its 地位 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 反対 fungal-induced diseases. Treatment with melatonin at concentrations の 0.05–0.5 mmol/L can enhance resistance to brown spot disease によって regulating the activity の 抗酸化 and 国防-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 の exogenous melatonin can inhibit the growth の fungal pathogens such as Botrytis, Fusarium, and Fusarium [76].

In terms の its mechanism の 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 トマト pathogenic strain DC3000 (Pst DC3000) is the most widely used model in studies の plant-pathogen interactions [77].

Exogenous melatonin pretreatment at certain concentrations enhances the resistance の Arabidopsis and tobacco to Pst DC3000 [78,79]. Lee et al. [80] found that inactivation の セロトニン-N-acetyltransferase significantly reduced endogenous melatonin levels in Arabidopsis,resulting in increased susceptibility の plants to Pst DC3000. Therefore, melatonin can enhance plant 寛容 to 細菌 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 の melatonin in plant-virus interactions, finding that exogenous melatonin アプリケーション significantly inhibited viral infection の tobacco seedlings. Additionally, exogenous melatonin pretreatment reduced the incidence の viral diseases in rice [82].With the deepening の research on melatonin-仲介 plant disease resistance mechanisms, melatonin could provide new strategies for the prevention and control の plant viral diseases.

In addition to phenotypic identification, recent analyses の gene expression have provided strong evidence for melatonin's 介入 in regulating plant responses to abiotic stress.Exogenous melatonin treatment or overexpression の melatonin synthesis-related genes can induce the expression の 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 国防 agent 反対 pathogens in plants [78,83-85].

In addition to pathogens, insect pests are another major biotic stress that plants face 中 growth. 植物-derived secondary metabolites act as antagonists の insect juvenile ホルモン to defend 反対 insect predation [86].It has been reported that dopamine, which has a similar structure to melatonin, plays an important role in plant 国防 反対 herbivores [87], suggesting that melatonin may also exert similar defensive effects [88].

3.3 Role の plant melatonin in responses to abiotic stress

Under normal growth conditions, the production and removal の 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 酸化 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 酸化 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 酸化 stress [92-95].

Melatonin may control the burst の hydrogen 過酸化 中 non-biotic stress responses in plants by directly ゴミ拾い excess ROS, enhancing antioxidant enzyme activity, and improving the ascorbic 酸-glutathione (AsA-GSH) cycle capacity [67].Exogenous melatonin treatment enhances antioxidant enzyme activity in apples, grapes, corn, sunflowers, トマトes, and wheat, while reducing the concentrations の superoxide, hydrogen peroxide, and malondialdehyde [59,67,72,96-99].Additionally, exogenous melatonin treatment reduces the 蓄積 の oxidized proteins in plants, accelerates the occurrence の autophagy induced by oxidative stress, and alleを介してtes photooxidative damage [100].

4. Conclusion and Outlook

Since the discovery の melatonin in plants, its diverse physiological 機能 and significant 潜在 アプリケーションs 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 分子 mechanisms の 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 の Arabidopsis CAND2/PMTR1 as melatonin signal transduction 受容体 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 の melatonin transport within plants require further investigation.The heterologous overexpression の melatonin synthesis-related genes can significantly increase the endogenous melatonin levels in plants, while 遺伝子組み換え plants exhibit enhanced 寛容 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 寛容 to abiotic stress, and ultimately improving crop yields.

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