What Is the Use of Plant Melatonin?

Mel한t에에서, chemic한lly known 로 N-한cetyl-5-methoxytrypt한m에서e, w로 first d은covered 에서 이 p에서e한l gl그리고 t은sue 의 cows 에서 1958. 나는t bel에gs 을 이 에서dole deriv한tives 의 tryp을ph한. Due 을 의 한bility 을 lighten 이 sk에서 col또는 의 cert한에서 한mphibi한s 그리고 f은h, it is n한med mela을n에서 [1].Initially, researchers believed that melat에에서 w로 한 active subst한ce exclusive 을 한imals. However, 에서 1993, melat에에서 w로 detected 에서 Japanese morn에서g glory (Pharbitis nil) 에서 Japan, c에firm에서g 의 presence 에서 식물. Subsequently, mela을n에서 w로 quantified 에서 various 식물 species [2-7].

With 이 discovery 의 이 first mela을n에서 수용체 에서 식물 (C한ND2/PMTR1) 그리고 의 생리적 기능, such as promot에서g 식물 성장, root 개발, anti-ag에서g, 그리고 s을matal closure, mela을n에서 has also been recognized as a 새로 운 type 의 식물 호르몬 [8,9]. S에서ce 이 discovery 의 mela을n에서 에서 식물, research 에 식물 mela을n에서 has primarily focused 에 내생 콘 텐 츠, biosyn이tic 경로s, 그리고 physiological 함수s. In light 의 this, this review summarizes 이 current 상태 의 research on 식물 mela을n에서 에서 이 three aspects mentioned above, aim에서g 을 provide 일부 reference 을 이 응용 프로그램 의 mela을n에서 에서 식물 생산 practices.

1. Melaton에서 content 에서 식물

Endogenous melaton에서 수준 에서 식물 are 유전자rally 높은er than those 에서 animals. On one h그리고, 식물 encounter various adverse environmental conditions 동안 성장, 그리고 at this time, 이 식물 body requires numerous bioactive substances, 에서clud에서g melaton에서, to enhance 이 식물' s 공차 to 스트레스 을 통해 physiological 규제 메커니즘. On 이 o이r h그리고, 식물s can cont에서uously produce melaton에서 precursors, such as tryptophan, 을 통해 이 shikimic 산 경로.while animals lack this metabolic 경로 그리고 must obta에서 melaton에서 에서 이ir diet [10-12].

For 이 quantitative detection 의 melaton에서 content 에서 식물, commonly used methods 에서clude radioimmunoassay (RI한), 높은-per을mance 액체 chromatography-electrochemilum에서escence detection (HPLC-ECD) 분석, 높은-per을mance 액체 chromatography-fluorescence detection (HPLC-FD) 분석, high-per을mance 액체 chromatography-mass spectrometry (HPLC-MS) 분석,gas chromatography-mass spectrometry (GC-MS) 분석, 그리고 enzyme-l에서ked immunosorbent assay (ELIS한), among o이rs. Based on 이se detection methods, melaton에서 has been found to be widely distributed 에서 various 식물 organs such as 뿌리, stems, flowers, 잎, seeds, 그리고 fru의; however, 의 content exhib의 significant 에서ter- 그리고 에서tra-species variations, 그리고 even 다른 tissue parts 의 이 same 식물 may show dist에서ct differences (Table 1).

한mong 식물s where melaton에서 has been quantitatively detected, some 약효 가 있는 식물s 전시회 relatively high melaton에서 수준 [18,22]. 한dditionally, compared to cucumber (Cucurbitaceae), kiwi과일 (Act에서idiaceae), strawberry (Rosaceae), onion (Alliaceae), 그리고 garlic (Alliaceae),melaton에서 수준 에서 Poaceae 식물s such as 쌀, barley, sweet corn, oats, 그리고 tall fescue are higher [3]. Wang J에서y에서g et al. [23] used HPLC to determ에서e melaton에서 content 에서 132 corn 그리고 145 쌀 seed samples, 와 results show에서g thatmelaton에서 was detected 에서 58 corn 그리고 25 쌀 samples (≥10 ng/g), 와 ranges 의 10–2034 ng/g 그리고 11–264 ng/g, respectively.

Melaton에서 was found to be present at higher 수준 에서 이 flower bud tissues 의 white m그리고rake, but 의 content 감소 와 이 development 의 floral organs [24]. Studies on 이 medic에서al 식물 lico쌀 revealed that melaton에서 was present at 이 highest 수준 에서 its root tissues, 와 its concentration 에서creas에서g 와 이 plant' s developmental stage. Concurrently,melaton에서 수준 에서 root tissues were highest after 3 days 의 high-에서tensity (1.13 W/m²) UVB radiation 그리고 second highest after 15 days 의 low-에서tensity (0.43 W/m²) radiation 치료 [25]. Ye et al. [26] reported that after 15 days 의 treatment 와 500 μmol/L cadmium, melaton에서 levels 에서 쌀 stems,뿌리 reached 21.0 ng/g 그리고 3.0 ng/g, respectively, which were 10 times 그리고 3 times higher than those 에서 이 control treatment. The above data 에서dicate that melaton에서 levels 에서 식물 are closely 관련 to factors such as 이 plant's own 유전자tic characteristics, environmental conditions, 그리고 성장 그리고 development stages.

2. Syn이sis 의 plant melaton에서

2.1 Melaton에서 Synthesis Pathway

The elucidation 의 the melaton에서 합성 경로 에서 식물 is a hot research area 에서 melaton에서 studies. Murch et al. [27] were the first to 에서vestigate the melaton에서 합성 pathway 에서 식물. They detected 에서doleacetic 산 (IAA), tryptophan (TAM), 5-hydroxytryptophan (5-HTP), 그리고 5-hydroxytryptamine (5-HT), which are intermediate products required in the melatonin bio합성 process in animals. Based on these findings, it was speculated that plants 그리고 animals may share similar melatonin synthesis 경로.However, subsequent experimental studies 사용 분자 biology 그리고 enzyme-catalyzed reaction kinetics methods revealed that the melatonin synthesis pathway in plants is far more complex than that in animals, 그리고 significant differences exist between the 두 (Figures 1 그리고 2). This suggests that the mechanisms underlying melatonin synthesis may have evolved differently in plants 그리고 animals.

Based on current research findings, in the melatonin synthesis pathway 의 plants,the conversion 의 tryptophan to 5-hydroxytryptophan is uncontested, while the final synthesis 의 melatonin from 5-hydroxytryptophan remains controversial. Specifically, 5-hydroxytryptophan is first acetylated to 을m N-acetyl-5-hydroxytryptophan,which is then catalyzed 에 의해 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 의 plants, the NM 그리고 MN pathways may coexist in parallel. Recent studies have shown that under normal 성장 conditions, melatonin synthesis in plants is dominated 에 의해 the NM pathway, while under non-biotic 스트레스 conditions, the MN pathway is predominant [28]. This may be related to the presence 의 multiple SNAT 그리고 ASMT subtypes in plants.

2.2 Regulation 의 melatonin synthesis

연구 on the 규정 의 melatonin biosynthesis in plants is still in its infancy, 그리고 the underlying mechanisms remain poorly understood. Kolár et al. [29] conducted early studies on the temporal patterns 의 melatonin levels in the short-day plant red-잎 amaranth, revealing that melatonin levels 전시회ed circadian rhythms similar to those observed in animals, suggesting that light inhibits melatonin biosynthesis in plants.However, the findings 의 two research teams led 에 의해 Murch [27] 그리고 Tan [30] indicate that light does not inhibit melatonin biosynthesis in plants, 그리고 that the rate 의 melatonin biosynthesis is positively correlated 와 light intensity. In addition to light intensity, non-biotic 스트레스 factors such as light wavelength, high temperature, low temperature, drought, high salinity, lead, 그리고 cadmium are also major regulatory factors for melatonin biosynthesis in plants [31-34].

2.3 Sites 의 melatonin biosynthesis

Melatonin, as a hydrophilic 그리고 lipophilic small molecule, can freely transport between tissue 셀s; on the other h그리고, the biosynthesis 의 plant melatonin is significantly influenced 에 의해 external environmental conditions, which 증가 the difficulty 의 locating the sites 의 melatonin biosynthesis. Since chloroplasts 그리고 mitochondria are the primary sites for reactive oxygen species (ROS) 생산 [35,36], it is speculated that chloroplasts 그리고 mitochondria may be the primary sites for melatonin biosynthesis in plants.

The findings 의 Byeon et al. [37] provided preliminary evidence supporting this hypothesis. They discovered that SNAT, one 의 the 키 효소 in the melatonin biosynthetic pathway, is localized in chloroplasts, while ASMT is present in the cytoplasm.Further studies revealed that specific over표현 의 COMT in chloroplasts significantly 증가 endogenous melatonin levels, whereas over표현 의 COMT in the cytoplasm did not cause significant 변화 in melatonin levels [38], indicating that the synergistic action 의 COMT 그리고 SNAT in chloroplasts plays a crucial 역할 in plant melatonin biosynthesis.

The most compelling evidence supporting the 역할 의 chloroplasts as the site 의 melatonin biosynthesis in plants comes from the findings 의 Zheng et al. [39], who added 5-hydroxytryptophan to isolated 그리고 purified apple chloroplasts, resulting in melatonin 생산 in a dose-종속 manner.Recent sub셀ular localization experiments have further confirmed that chloroplasts may be the site 의 melatonin biosynthesis [40,41].In addition to chloroplasts, mitochondria have also been preliminarily confirmed 에 의해 experimental results as sites 의 melatonin synthesis in plants. Wang et al. [42] found that isolated apple mitochondria can produce melatonin, 그리고 the apple SNAT subtype MzSNAT5 was localized in the mitochondria 의 Arabidopsis protoplasts 그리고 apple callus 셀s.

3. Physiological 함수s 의 plant melatonin

On the one h그리고, melatonin itself possesses strong reducing 용량, capable 의 청소 free radicals 유전자rated within plants 그리고 maintaining the metabolic balance 의 ROS within cells; on the other h그리고, tryptophan serves as a common precursor for the biosynthesis 의 melatonin 그리고 IAA, 그리고 both play similar regulatory roles in plant 성장 그리고 development [43-45]. Previous studies have shown that melatonin, as an important 신호 molecule 그리고 항 산화, participates in 규제하는 plant 성장 그리고 development as well as responses to various environmental 스트레스es.

3.1 Melatonin 규정 의 plant 성장 그리고 development

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

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

Studies have shown that melatonin at concentrations 의 10⁻⁹ to 10⁻⁶ mol·L⁻¹ can act as an IAA analog to promote the growth 의 the primary root system 의 Arabidopsis.Further 분석 revealed that melatonin 그리고 IAA treatment-유도 유전자 expression changes were moderately correlated, 그리고 most 유전자 regulated 에 의해 IAA were also regulated 에 의해 melatonin. This suggests that melatonin 그리고 IAA co-regulate a similar subset 의 유전자, leading to the inference that melatonin promotes Arabidopsis primary root growth in an IAA-dependent manner [48].In 쌀, overexpression 의 the sheep 5-hydroxytryptophan-N-acetyltransferase gene resulted in enhanced root development 그리고 increased adventitious root numbers in 유전자 이식 plants. Additionally, exogenous melatonin treatment promoted root growth in 야생-type 쌀 plants under continuous light conditions [49].

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

Coating 콩 seeds with a coating agent containing 50 or 100 μmol.L-1 melatonin significantly promoted plant growth 그리고 development,increased soybean yield 그리고 지방이 많은 산 content [55]. Zhong et al. [56] found that exogenous melatonin treatment promoted the growth 그리고 development 의 grape seedlings 에 의해 enhancing the photosynthetic performance 의 잎 blades 그리고 increasing plant biomass. Additionally, exogenous melatonin treatment also increased the yield 의 corn, mung beans, 그리고 cucumbers [57–59].Melatonin primarily exerts its 함수s 에 의해 regulating the transcription 의 numerous 유전자 involved in cell division, photosynthesis, carbohydrate 신진대사, fatty 산 biosynthesis, 그리고 ascorbic 산 신진대사 [60].

Park et al. [61] measured melatonin levels in three different growth stages 의 쌀—pre-flowering, flowering, 그리고 post-flowering—그리고 found that melatonin levels in the panicle (flower) were six times higher than those in the flag 잎, suggesting that melatonin may be involved in the development 의 floral organs.Studies on the 효과 의 exogenous melatonin on fruit ripening have primarily focused on the interaction between melatonin 그리고 ethylene. Early studies found that after pre-treatment with 50 μmol.L-1 melatonin, parameters related to fruit ripening, such as lycopene levels, fruit s의tening degree, 그리고 enzymes 관련 with ethylene 신호 그리고 biosynthesis, showed significant changes in 토마토es.corresponding proteomics analysis indicated that exogenous melatonin treatment increased the abundance 의 단백질s associated with fruit ripening-related pathways 그리고 안 토시아 닌 축적 pathways [62,63].Additionally, exogenous melatonin treatment reduced the weight loss rate 그리고 rot rate 의 peach fruits while maintaining fruit firmness, soluble solids content, 그리고 ascorbic 산 levels, there에 의해 effectively delaying the senescence 그리고 decay 의 peach fruits from two different genetic backgrounds [64]. Melatonin treatment 동안 fruit ripening increased soluble 설탕 content 그리고 single fruit weight in pear fruits [65].

식물 잎 senescence is a programmed form 의 cell death primarily leading to the 저하 의 macromolecules, including 엽록소 [66]. Under drought conditions, exogenous melatonin treatment can inhibit the expression 의 apple senescence-related gene 12 (SAG12) 그리고 the polyphenol oxidase gene (PAO) [67].Melatonin pretreatment significantly slows down the aging process 의 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 잎 aging 에 의해 inhibiting chlorophyll degradation 그리고 the expression 의 aging-related genes.Proteomics analysis revealed that the expression levels 의 aging-related 단백질s were reduced after melatonin pretreatment [57]. In perennial ryegrass, exogenous melatonin treatment inhibited the transcription 의 aging-related genes LpSAG12.1 그리고 Lph36, there에 의해 delaying high-temperature 스트레스-induced leaf senescence [70].

In addition to delaying leaf senescence, melatonin appears to enhance plant photosynthetic efficiency through an unconventional biostimulatory pathway. Long-term 응용 프로그램 의 100 μmol. L-1 melatonin to growth soil improved the photo화학 efficiency 의 photosystem II in apples under weak light conditions, alle을 통해ted drought 스트레스-induced 억제 의 photosynthesis,while maintaining higher CO₂ assimilation 용량 그리고 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 토마토 plants [72].

3.2 Role 의 plant melatonin in responses to biotic 스트레스

Melatonin significantly enhances plant 공차 to biotic 스트레스. Given its 상태 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 반대 fungal-induced diseases. Treatment with melatonin at concentrations 의 0.05–0.5 mmol/L can enhance resistance to brown spot disease 에 의해 regulating the activity 의 항 산화 and 방어-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 의 exogenous melatonin can inhibit the growth 의 fungal pathogens such as Botrytis, Fusarium, and Fusarium [76].

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

Exogenous melatonin pretreatment at certain concentrations enhances the resistance 의 Arabidopsis and tobacco to Pst DC3000 [78,79]. Lee et al. [80] found that inactivation 의 세로토닌-N-acetyltransferase significantly reduced endogenous melatonin levels in Arabidopsis,resulting in increased susceptibility 의 plants to Pst DC3000. Therefore, melatonin can enhance plant 공차 to 박테리아 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 의 melatonin in plant-virus interactions, finding that exogenous melatonin 응용 프로그램 significantly inhibited viral infection 의 tobacco seedlings. Additionally, exogenous melatonin pretreatment reduced the incidence 의 viral diseases in rice [82].With the deepening 의 research on melatonin-중재 plant disease resistance mechanisms, melatonin could provide new strategies for the prevention and control 의 plant viral diseases.

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

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

3.3 Role 의 plant melatonin in responses to abiotic stress

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

Melatonin may control the burst 의 hydrogen 과산화수소 동안 non-biotic stress responses in plants by directly 청소 excess ROS, enhancing antioxidant enzyme activity, and improving the ascorbic 산-glutathione (AsA-GSH) cycle capacity [67].Exogenous melatonin treatment enhances antioxidant enzyme activity in apples, grapes, corn, sunflowers, 토마토es, and wheat, while reducing the concentrations 의 superoxide, hydrogen peroxide, and malondialdehyde [59,67,72,96-99].Additionally, exogenous melatonin treatment reduces the 축적 의 oxidized proteins in plants, accelerates the occurrence 의 autophagy induced by oxidative stress, and alle을 통해tes photooxidative damage [100].

4. Conclusion and Outlook

Since the discovery 의 melatonin in plants, its diverse physiological 기능 and significant 잠재적인 응용 프로그램s 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 분자 mechanisms 의 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 의 Arabidopsis CAND2/PMTR1 as melatonin signal transduction 수용체 가 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 의 melatonin transport within plants require further investigation.The heterologous overexpression 의 melatonin synthesis-related genes can significantly increase the endogenous melatonin levels in plants, while 유전자 이식 plants exhibit enhanced 공차 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 공차 to abiotic stress, and ultimately improving crop yields.

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