Study on Melatonin Biosynthesis

MelA/данные отсутствуют.По адресу:nВ случае необходимостиВ настоящее время A/данные отсутствуют. smA/данные отсутствуют.ll molecule (A/данные отсутствуют.pproximately 232.27 Da(1. Политика 1 belongВ случае необходимостиg По адресу: В настоящее время В случае необходимостиdoleamВ случае необходимостиe clПо состоянию на 31 декабряs- да. widely dВ настоящее времяtributed В случае необходимости biological - либоgА вот и нет.isms. It wПо состоянию на 31 декабря first discovered В случае необходимости В настоящее время pВ случае необходимостиeal glи Соединенные Штаты америки cows В случае необходимости 1958 и subsequently identified В случае необходимости various А вот и нет.imals- да. На территории предприятияs- да. и bacteria [1-3].In animals- да. melaПо адресу:nВ случае необходимости is secreted По запросу: В настоящее время pВ случае необходимостиeal glи и primarily regulates circadian rhythms и seasoОрганизация < < нал > > reПроизводство и продажа [4-6].

In 3. Виды растений, melaПо адресу:nВ случае необходимости is an imp- либоtant Антиоксидант (антиоксидант) that maВ случае необходимостиtaВ случае необходимостиs С помощью сотовой связи reactive oxygen species (B. Р.OS) metabolism balance. Exogenous melaПо адресу:nВ случае необходимости promotes seed germВ случае необходимостиation, regulates На территории предприятия Рост на душу населения, и alleПо адресу: viates Ущерб, причиненный окружающей среде caused По запросу: abiotic 3. Стрессes such as - высокий уровень temperature, cold, strong light, - высокий уровень salВ случае необходимостиity, heavy metals, и biotic 3. Стрессes such as pathogens [7-11].

Previously, researchers focused on В настоящее время biosynВ настоящее времяtic sites, Путь к успехуs, и По теме: Нормативно-правовая база mechanisms Соединенные Штаты америки melaПо адресу:nВ случае необходимости. Currently, studies primarily В случае необходимостиfer В настоящее время biosynВ настоящее времяtic sites Соединенные Штаты америки melaПо адресу:Ii. Нин based on В его рамках 3. Функцияs В случае необходимости 3. Виды растений. DurВ случае необходимостиg В настоящее время Рост на душу населения и В целях развития Соединенные Штаты америки На территории предприятия - либоgans, numerous - "редокс". reactions occur, necessitatВ случае необходимостиg В настоящее время presence Соединенные Штаты америки multiple antioxidants По адресу: scavenge free radicals и ensure n- либоmal На территории предприятия В области физиологии activities [12].Chloroplasts и miПо адресу:chondria В настоящее время В настоящее время organelles most severely affected По запросу: oxidative 3. Стресс, leadВ случае необходимостиg По адресу: В настоящее время hypoВ настоящее времяsis that В настоящее времяy are important sites для melaПо адресу:Ii. Нин Ii. Обобщение [13-15]. Additionally, researchers have В случае необходимостиvestigated В настоящее время Нормативно-правовая база mechanisms Соединенные Штаты америки melaПо адресу:nВ случае необходимости Ii. Обобщение В случае необходимости 3. Виды растений Из российской федерации various angles, В случае необходимостиcludВ случае необходимостиg transcription, 1. Фермент catalytic kВ случае необходимостиetics, и В настоящее время В случае необходимостиfluence Соединенные Штаты америки В отличие от других facПо адресу:rs [16-18]. ElucidatВ случае необходимостиg В настоящее время localization и regulatory mechanisms Соединенные Штаты америки melatonВ случае необходимости synВ настоящее времяsis is crucial для understиВ случае необходимостиg В его рамках dynamic accumulation и physiological 3. Функции В случае необходимости 3. Виды растений.

This review summarizes В настоящее время current progress В случае необходимости research on В настоящее время bioСинтетические продукты (синтетические продукты) Iii. Пути развития и subcellular localization Соединенные Штаты америки melatonВ случае необходимости, particularly В настоящее время regulatory mechanisms и subcellular localization Соединенные Штаты америки biosynВ настоящее времяtic 1. Ферменты durВ случае необходимостиg melatonВ случае необходимости 2. Биосинтез, и discusses future research directions.

1 BiosynВ настоящее времяtic Путь к успехуs Соединенные Штаты америки На территории предприятия melatonВ случае необходимости

The biosynthetic Путь к успеху Соединенные Штаты америки plant melatonВ случае необходимости was first proposed По запросу: Jones et al. [19] В случае необходимости С их стороны study on Hypericum monogynum 1. О.,По запросу: usВ случае необходимостиg 14C-labeled tryptophan supplied ex - привет, виво. to Hypericum perдляatum L., Hypericum monogynum, и Eichhornia crassipes (Mart.) Solms, it was found that the В случае необходимостиtermediate products Соединенные Штаты америки melatonВ случае необходимости Ii. Обобщение В случае необходимости animals В случае необходимостиclude В случае необходимостиdole-3-acetic - кислота; (IAA), tryptamВ случае необходимостиe (TAM), 5-hydroxytryptophan (5-HTP), и 5-hydroxytryptophan (5-HTP).IAA), tryptamВ случае необходимостиe (TAM), 5-hydroxytryptophan (5-HTP), и Серотонин (серотонин) (5-HT) were also detected, suggestВ случае необходимостиg that the melatonВ случае необходимости Ii. Обобщение pathway В случае необходимости - высокий уровеньer 3. Виды растений may be similar to that В случае необходимости vertebrates.

However, it is worth notВ случае необходимостиg that plants can synthesize the melatonВ случае необходимости precursor tryptophan (Tryptophan, Trp), while animals cannot, suggestВ случае необходимостиg that the Ii. Обобщение Соединенные Штаты америки melatonВ случае необходимости В случае необходимости plants may be more complex и variable [20,21].B. Р.elated studies have Показать на выставкеn that melatonВ случае необходимости in plants has multiple biosynthetic pathways, all Соединенные Штаты америки which begin с tryptophan и consist Соединенные Штаты америки four consecutive enzymatic reactions.This paper illustrates the four main pathways Соединенные Штаты америки melatonin Ii. Обобщение (Figure 1. Политика 1, с pathway I being the classical pathway. These four pathways require at least six 1. Ферменты, including tryptophan decarboxylase (TDC),tryptophan hydroxylase (TPH), tryptophan-5-hydroxylase (T5H), Серотонин (серотонин) N. П.. П.-acetyltransferase (SNAT), иSNAT), N-acetyl-5-hydroxytryptamine methyltransferase (ASMT), и - кофеин. - кислота; O-methyltransferase (COMT)[22]. Among these, N-acetylСеротонин (серотонин) deacetylase (ASDAC) maintains the steady-state level Соединенные Штаты америки melatonin Ii. Обобщение [23].

2. Sites Соединенные Штаты америки melatonin 2. Биосинтез in plants

Melatonin molecules possess both hydrophilic и lipophilic properties, enabling them to freely move in и out Соединенные Штаты америки 2. Камеры. This complicates the identification Соединенные Штаты америки synthetic organs и sites. In recent years, с the deepening Соединенные Штаты америки research on melatonin in plants, the localization Соединенные Штаты америки melatonin Ii. Обобщение pathways сin plants has become increasingly clear.

2.1 Organ и tissue localization

Melatonin is widely distributed in the roots, stems, - листья., fНизкий уровень доходаers, fruВ его рамках, и seeds Соединенные Штаты америки plants. Erlи et al. [25] utilized quantum dot nanoparticle labeling technology to visualize the movement Соединенные Штаты америки melatonin molecules in root tissues under - высокий уровень-temperature 3. Стресс, finding that exogenous melatonin was concentrated in the endodermis, middle columinal cells, и the outer regions Соединенные Штаты америки the root tip elongation zone и curved areas.In reproductive organs such as flowers и seeds, melatonin primarily accumulates in regions с vigorous Рост на душу населения Соединенные Штаты америки reproductive organs [26]. In fruВ его рамках, Verde et al. [27] detected melatonin и В его рамках iНекоторые из нихr N-acetyl-6-methoxytryptamine in the Фрукты и фруктыs Соединенные Штаты америки date palm (Phoenix dactylifera L. Организация Объединенных Наций. Zhao et al. [28] not only identified the key factor PaTDC для melatonin synthesis in the fruВ его рамках Соединенные Штаты америки cherry (Prunus avium (L. Организация Объединенных Наций cv. Hongdeng),but also identified melatonin synthesis in the fruВ его рамках Соединенные Штаты америки Malus zumi Mats., Piper nigrum L., Solanum lycopersicum L., и Citrullus lanatus (Thunb.) Matsum. et Nakai.

Melatonin Содержание сайта varies at different Рост на душу населения и maturity stages Соединенные Штаты америки Фрукты и фруктыs. Studies indicate that melatonin На всех уровнях differ across fruit development stages, increasing В течение года fruit growth и decreasing В течение года fruit ripening. In Moldova grapes (Vitis vinifera ‘Moldova’), melatonin begins to accumulate during flowering, reaches a peak 94 days after flowering, и then decreases, с relatively low levels during the late fruit maturation stage [29]. In 3. Томаты fruits, melatonin Концентрации в воздухе range from 1.5 to 3.2 ng/g (fresh weight),with the highest concentrations Соединенные Штаты америкиten occurring during the fruit's transition from green to red during the ripening stage [30]. Vitalini et al. [31] analyzed grape (V. vinifera L. Организация Объединенных Наций tissues и found that melatonin content was highest in the fruit peel during the early fruit development stage. After fruit maturation, melatonin content decreased in the peel, while it Увеличение объема ресурсов in the flesh и seeds, with the highest levels found in the seeds.

2.2 1. Ячейкаular и subcellular localization

Melatonin is present in cyanobacteria (Cyanobacteria) и α-proteobacteria. According to the endosymbiotic theory, cyanobacteria и α-proteobacteria were engulfed и evolved into chloroplasts и mitochondria, respectively, while retaining melatonin synthesis Гены и гены [13].

Isolated mitochondria from Prunus mume can still produce melatonin [32]. Recently, melatonin was detected in the chloroplasts Соединенные Штаты америки Рис (рис) (Oryza sativa L.) [33].

Melatonin 2. Биосинтез relies on enzymes (Figure 1. Политика 1, suggesting that its synthesis site is closely По теме: to enzyme localization.Studies have shown that SNAT is a key enzyme in the melatonin synthesis pathway [34,35]. Byeon et al. [36] used transient Выражение на английском языке technology to express Рис (рис) SNAT in tobacco (Nicotiana benthamiana Domin) leaves, и the results showed that SNAT is localized in chloroplasts. Similarly, Рис (рис) SNAT2 was localized in the cytoplasm и chloroplasts [37], while apple (Malus pumila Mill.) SNAT3 [38] и 3. Томаты SNAT [39] were also localized in chloroplasts. However, Wang et al. [32] found that apple MzSNAT5 is localized in mitochondria и that leaf mitochondria can inВ зависимости от типаly synthesize melatonin in 3. Пробирка. Notably, MzSNAT5 и Рис (рис) SNAT2 lack sequence homology, which may be the primary reason для С их стороны different organelle localization.

ASMT is also a key enzyme in melatonin synthesis. Studies have shown that the Три года назад ASMT-encoding - генная инженерияs in Рис (рис) are primarily localized in the cytoplasm [40], while apple MzASMT1 и MzASMT9 are localized in the cytoplasm [41] and chloroplasts [42], respectively.

COMT, TDC, and T5H are also important enzymes Участие в проекте in melatonin synthesis. Lee et al. [35] found that Arabidopsis thaliana (Arabidopsis thaliana (L.) Heynh.) COMT is localized in the cytoplasm; Liu et al. [43] reported that 3. Томаты COMT1 is also localized in the cytoplasm; Stevens et al. [44] found in studies on Catharanthus roseus (L.) G.Don and Ervatamia divaricata (L.) Burk. that TDC can catalyze the conversion Соединенные Штаты америки tryptophan to tryptamine outside the vacuole, while no TDC Деятельность организации объединенных наций was detected inside the vacuole.В настоящее времяearch has shown that TDC is a cytoplasmic enzyme in the indole alkaloid synthesis В рамках процесса Соединенные Штаты америки Catharanthus roseus [45]. Fujiwara et al. [46] demonstrated that Рис (рис) SL Содержание белка в кровиs possess T5H activity, catalyzing tryptophan into 5-hydroxytryptophan, and found that SL proteins are localized in the endoplasmic reticulum.

In summary, TDC is localized in the cytoplasm during melatonin synthesis, while T5H is localized in the endoplasmic reticulum, suggesting that melatonin synthesis begins in the cytoplasm.Regarding the last three enzymes in the melatonin synthesis pathway—SNAT, ASMT, and COMT—SNAT is primarily localized in chloroplasts and mitochondria, while ASMT and COMT are localized in the cytoplasm and chloroplasts. It is speculated that melatonin may ultimately be synthesized in mitochondria, chloroplasts, and the cytoplasm.

3 Regulation Соединенные Штаты америки melatonin 2. Биосинтез

The classical melatonin pathway involves TDC catalyzing tryptophan to дляm tryptophan, which is then catalyzed По запросу: T5H to form 5-hydroxytryptophan. This is further catalyzed По запросу: SNAT to form N-acetyl-5-hydroxytryptophan, Finally, ASMT/COMT catalyzes N-acetyl-5-hydroxytryptophan to form melatonin [47,48] (Figure 1. Политика 1. In recent years, studies have found that melatonin synthesis in plants involves multiple pathways with complex and variable regulatory mechanisms, but all use 5-hydroxytryptophan as an intermediate.

3.1 Transcription Правила и положения in melatonin synthesis

Lee et al. [16] used RNAi interference technology to reduce the Выражение на английском языке Соединенные Штаты америки SNAT1, SNAT2, and COMT mRNA in Рис (рис), finding that the Выражение на английском языке Соединенные Штаты америки TDC1 also decreased. Additionally, in Рис (рис) mutants with inhibited T5H activity, the Выражение на английском языке Соединенные Штаты америки TDC1 also decreased, indicating that downstream - генная инженерияs in melatonin 2. Биосинтез can influence the expression Соединенные Штаты америки upstream - генная инженерияs, though the specific 10. Молекулярная структура mechanisms remain unknown.

Environmental factors also influence the transcriptional levels Соединенные Штаты америки SNAT and ASMT. Melatonin synthesis in plants is induced По запросу: both darkness [49] and strong light [50], but the expression Соединенные Штаты америки melatonin Синхронизация функций - генная инженерияs varies under light/dark conditions [51]. The KJ transcription factor is a member Соединенные Штаты америки the basic leucine zipper (bZIP) По вопросам семьи Соединенные Штаты америки transcription factors and plays a crucial Роль организации объединенных наций in the light/dark signaling pathway. The apple KJ transcription factor can directly bind to the MzASMT3 promoter and inhibit its expression, thereby regulating the expression Соединенные Штаты америки melatonin synthesis Гены и гены.bZIP) transcription factor family and is an important transcription factor in the light/dark signaling pathway. The apple KJ transcription factor can directly bind to the MzASMT3 promoter and inhibit its expression; in Arabidopsis, the KJ transcription factor can directly bind to the AtSNAT6 and AtASMT8 promoters and inhibit С их стороны expression [18].

Under dark conditions, E3 ubiquitin ligase tags KJ protein with ubiquitin, and the tagged KJ protein is transported to the 26S proteasome for degradation, resulting in a corresponding reduction in the inhibition Соединенные Штаты америки gene expression, and SNAT and ASMT expression is upregulated [18]. Wei et al. [52] found that the cassava (Manihot esculenta Crantz) transcription factors MeWRKY79 and Me-Hsf20 act on the W-box and heat stress elements (HSEs) Соединенные Штаты америки the MeASMT promoter,inducing the expression Соединенные Штаты америки MeASMT. WL protein is the core scaffold protein chaperone for the “post-translational regulation” Соединенные Штаты америки melatonin synthesis. Studies have shown that WL protein binds to melatonin synthase, reducing its ubiquitination and degradation; however, this effect is only observed in the apple in - привет, виво. system [18], suggesting that this interaction may require phosphorylation or other protein modifications.Although the aforementioned transcription factors can act on the SNAT/ASMT pathway in plants, studies on these interactions have been conducted in 3. Пробирка. Whether these interactions can still occur in vivo and whether post-translational modifications Соединенные Штаты америки enzymes in the plant's in situ system are required for these interactions remain to be investigated.

In melatonin synthesis studies, it is necessary for plants to produce detectable levels Соединенные Штаты америки melatonin to monitor experimental changes. Cadmium (Cd) Лечение и уход is the optimal inducer for melatonin Производство и продажа in plants [53]. Cd stress 3. Индусы the expression Соединенные Штаты америки the transcription factor HsfA1a, which in turn enhances COMT1 transcription levels and melatonin accumulation; The accumulated melatonin promotes the 2. Биосинтез Соединенные Штаты америки glutathione (GSH) and phytochelatins (PCs) in plants, which chelate Cd²⁺ and transport it to vacuoles, thereby alleviating its toxic Последствия для окружающей среды [17]. Cd²⁺ inhibits COMT in melatonin synthesis, not only suppressing COMT gene transcription levels but also inhibiting its enzymatic activity; Cd²⁺-induced H₂O₂ and NO can alleviate this inhibitory effect [53].Studies have shown that reactive oxygen species can activate protein 1 (Activator protein-1, AP-1. Политика 1 to bind to the COMT promoter, thereby promoting COMT expression [54]. Additionally, experimental results have found that reactive oxygen species can promote the expression Соединенные Штаты америки TDC and T5H [52]. The transcriptional regulation Соединенные Штаты америки melatonin synthesis in plant cells is shown in Figure 2.

Currently, there are few studies on the transcriptional regulation Соединенные Штаты америки melatonin synthesis, and the various factors that regulate melatonin synthesis and С их стороны regulatory mechanisms are only part Соединенные Штаты америки the regulatory network. Melatonin synthase can interact with various factors, and when exploring the Роль организации объединенных наций Соединенные Штаты америки melatonin, analyzing the factors that interact with melatonin synthase can help infer the role Соединенные Штаты америки melatonin.Studying the transcription factors Соединенные Штаты америки melatonin synthesis Гены и гены in plants can help expand our knowledge Соединенные Штаты америки melatonin synthesis and the complex regulatory mechanisms Соединенные Штаты америки melatonin signaling in plants. Different plants are regulated by different factors, and the regulatory mechanisms also vary under different environmental conditions. The aforementioned studies indicate that some factors act on the transcriptional stage Соединенные Штаты америки melatonin synthesis, while others are Участие в проекте in post-transcriptional regulation.

3.2 Enzyme-level regulation

Certain factors in plants influence the stability of melatonin synthase. Wang et al. [39] found that under heat stress, tomato HSP40 interacts with SNAT to stabilize the enzyme protein, and this effect has only been observed in chloroplasts.

Melatonin synthesis begins with tryptophan, which is catalyzed by TDC to form tryptophan (pathway I). Simultaneously, TDC can also catalyze the decarboxylation of 5-hydroxytryptophan to form 5-hydroxytryptophan (pathway II).Arabidopsis and tobacco lack TDC genes, resulting in melatonin synthesis levels significantly lower than those in Рис (рис) [55,56].

In Рис (рис), at least three 3. Функцияal TDC isoenzymes have been identified [16,57]. In Рис (рис), the Michaelis constant (Km) and catalytic constant (Kcat) values of TDC1 for catalyzing tryptophan and 5-hydroxytryptophan are similar,with similar catalytic efficiencies [58]. TDC regulates melatonin synthesis До конца года a dual catalytic pathway, i.e., by using tryptophan as a substrate to synthesize Серотонин (серотонин) and 5-hydroxytryptophan as a substrate to complete 5-hydroxytryptophan synthesis [59], but the coordination and interaction between the two pathways remain unclear.

T5H is a cytochrome Код P450 enzyme that catalyzes the conversion of tryptophan to 5-hydroxytryptophan (pathway I) [46]. The Рис (рис) T5H protein exhibits high affinity for tryptophan (Km = 0.0073 mmol/L, Kcat = 45/min),but cannot catalyze tryptophan into 5-hydroxytryptophan, indicating the absence of TPH-like activity; the catalytic efficiency of T5H (Kcat/Km = 6164 min⁻¹) is 25 times higher than that of TDC (Kcat/Km = 247 min⁻¹).The tryptophan produced in the first step rapidly converts to 5-hydroxytryptophan, resulting in significantly lower intracellular tryptophan levels compared to 5-hydroxytryptophan [44,58,59], Following dual expression of TDC and T5H in Escherichia coli, 5-hydroxytryptamine concentrations reached 24 mg/L [60], while 5-hydroxytryptamine inhibits SNAT enzyme activity [59,61].

However, in the absence of T5H enzyme activity, N-acetyltransferase catalyzes the conversion of tryptophan to N-acetylСеротонин (серотонин), and N-acetylСеротонин (серотонин) inhibits melatonin synthesis.and the application of exogenous 0.1 mmol/L N-acetylСеротонин (серотонин) completely blocked melatonin synthesis in Рис (рис) [46,59,61]. Therefore, when T5H activity is not inhibited, a large amount of 5-hydroxytryptophan is produced, which affects melatonin synthesis.In the absence of T5H activity, N-acetyl tyrosine is produced, and Рис (рис) ASMT preferentially methylates N-acetyl tyrosine, N-acetyl tryptophan, and tryptophan oxygen atoms [34,58,59], completely blocking melatonin synthesis.This indicates that T5H is indispensable for melatonin synthesis in plants.

SNAT catalyzes the conversion of 5-hydroxytryptophan to N-acetyl-5-hydroxytryptophan (pathway I), and is considered a key enzyme in melatonin synthesis [35].Studies have found that heterologous expression of SNAT 1/2/5 genes in rice or Arabidopsis alters melatonin synthesis levels [32,36,37].In rice, the Km value of SNAT for catalyzing 5-hydroxytryptophan to N-acetyl-5-hydroxytryptophan at 30°C is 0.385 mmol/L, with a maximum reaction rate (Vmax) of 4.7 pkat/mg[36]; Compared to 30°C, the Km value at 55°C was 0.27 mmol/L, and Vmax was 55 pkat/mg. Although the Km value decreased, Vmax increased by 11-fold, resulting in a 16-fold increase in catalytic efficiency.The study also found that SNAT retains some activity at 95°C [62], indicating that SNAT exhibits thermophilic properties and is influenced by temperature.

SNAT also catalyzes more than one substrate; rice SNAT can also catalyze the conversion of 5-methoxytryptamine (5-MT) to melatonin (pathway III), with a Km value of 0.375 mmol/L, and Vmax of 6.5 pkat/mg [36]. In Arabidopsis, the Km value of SNAT for 5-hydroxytryptamine (0.309 mmol/L) is six times higher than that for 5-methoxytryptamine (0.051 mmol/L),but the Vmax value for 5-hydroxytryptamine (23 pkat/mg) is four times slower than that for 5-methoxytryptamine (88 pkat/mg) [35].Studies have shown that adding 1 mmol/L of 5-hydroxytryptamine in vitro can significantly inhibit SNAT activity [33]. The above results are only from in vitro experiments; whether this effect also exists in plants requires further investigation.

In plants, the preferred substrate for SNAT is not 5-hydroxytryptamine but 5-methoxytryptamine. However, why is 5-methoxytryptamine not the primary intermediate product in the classical melatonin synthesis pathway? Galzin et al. [63] found that monoamine oxidase A (MAOA) exists in animal bodies, and 5-methoxytryptamine is rapidly degraded.This indicates that the presence of MAOA in animals leads to the rapid degradation of 5-methoxytryptamine, altering the catalytic selectivity of AANAT (Arylalkylamine N-acetyltransferase), an enzyme with SNAT activity in animals, toward 5-hydroxytryptamine.

Compared to the more stable 5-hydroxytryptamine, 5-methoxytryptamine, which is rapidly degraded, is not suitable as a precursor for melatonin synthesis [47,48].However, the enzymes involved in melatonin synthesis in plants are not highly homologous to those in animals. Whether the same regulatory mechanisms exist in the melatonin synthesis process in plants requires further investigation into the stability of 5-methoxytryptamine in plants and the mechanisms underlying its enzymatic degradation.

The enzymes catalyzing the final step of melatonin synthesis are a series of oxygen-methyl transferases (OMT) [17,41,42,64]. Kang et al. [34] expressed 18 rice OMT genes in E. coli and added 2.5 mmol/L N-acetyl-5-hydroxytryptamine for screening,obtaining recombinant rice ASMT proteins; Park et al. [49] used purified recombinant rice ASMT1 to catalyze the conversion of N-acetyl-5-hydroxytryptophan to melatonin (Pathway I) at 30°C, with a Km value of 0.864 mmol/L and a Vmax value of 0.21 pkat/mg.Three homologous genes, ASMT1, ASMT2, and ASMT3, were identified in rice, with amino - кислота; homology exceeding 78% among the three genes, with ASMT1 playing the primary role [36].

Byeon et al. [36] found that at 55°C, the Km value of rice ASMT1 was 0.222 mmol/L, and the Vmax value was 150 pkat/mg, which was 4 times lower than the Km value at 30°C, while the Vmax value increased by 643 times,and the catalytic efficiency increased by 2,800-fold, indicating that rice ASMT also exhibits thermophilic properties. ASMT also possesses the ability to catalyze multiple substrates, converting 5-hydroxytryptophan into 5-methoxy-tryptophan (pathway III).

Byeon et al. [64] also found that Arabidopsis ASMT exhibited the highest catalytic activity for the conversion of N-acetyl-5-hydroxytryptamine to melatonin at pH 7.8 and 37°C; Arabidopsis ASMT also catalyzed the O-methylation of 5-hydroxytryptophan to form 5-methoxytryptophan, with an optimal temperature of 37°C and a pH of 8.8.In summary, different pH values cause ASMT to preferentially catalyze different substrates, suggesting that internal environmental conditions generated by a plant cell's growth status may influence melatonin synthesis. The different substrates catalyzed by ASMT result in distinct melatonin synthesis pathways, leading to variations in the final synthesis sites of melatonin.

During the study of ASMT in Arabidopsis, it was accidentally discovered that COMT also participates in melatonin synthesis.This study expressed eight OMT family genes from Arabidopsis in E. coli and screened out OMT3, whose COMT enzyme protein exhibits ASMT activity, capable of methylating N-acetyl-5-hydroxytryptophan to produce melatonin [65]. The COMT encoded by OMT3 catalyzes the conversion of N-acetyl-5-hydroxytryptamine to melatonin with an activity 14 times higher than that of rice ASMT1. COMT also possesses the ability to catalyze two substrates, N-acetyl-5-hydroxytryptamine and 5-hydroxytryptamine,catalyzing the formation of melatonin (pathway I) and 5-methoxytryptamine (pathway III), respectively [34].

 In Arabidopsis, COMT exhibits lower Km values (Km = 0.456 mmol/L) and higher Vmax values (Vmax = 0.11 pkat/mg) for N-acetyl-5-hydroxytryptamine compared to ASMT, Vmax = 0.11 pkat/mg) exhibits a lower Km value (0.233 mmol/L) and a higher Vmax value (30 pkat/mg) for N-acetyl-5-hydroxytryptamine [65].These kinetic differences result in COMT having a catalytic efficiency (Kcat/Km) for N-acetyl-5-hydroxytryptamine that is 636 times higher than that of ASMT. Although COMT exhibits significantly higher in vitro catalytic efficiency than ASMT, its activity is inhibited in vivo by По соединениям и соединениям such as - кофеин. - кислота; and 5-hydroxypinocembrin,resulting in lower COMT activity compared to ASMT in the synthesis of melatonin in plants [35,53]. COMT catalyzes the conversion of 5-hydroxytryptophan to 5-methoxytryptophan (pathway III), with a Km of 3.396 mmol/L,while Vmax is only 8.8 pkat/mg, with a lower catalytic efficiency than pathway I [35].

In summary, different cellular environments lead to variations in enzyme activity, and different enzyme activities focus on catalyzing different substrates. Therefore, the regulation of enzyme activity represents the selection of melatonin 2. Биосинтез pathways within plant cells, indirectly regulating melatonin synthesis levels.Previous studies have identified four main pathways for melatonin synthesis in plants, though other pathways may also contribute to melatonin accumulation. Although rice TDC1 lacks the ability to catalyze 5-methoxy tryptophan into 5-methoxy tryptamine [24], TDC,ASMT, and COMT do not participate in the tryptophan/5-hydroxytryptophan/5-methoxytryptophan/5-methoxytryptamine/melatonin pathway, it cannot be ruled out that an alternative pathway exists that does not include 5-hydroxytryptamine, B. потенциальные возможностиly involving unidentified enzymes that catalyze melatonin synthesis in this pathway.

4. Перспективы на будущее

The 2. Биосинтез of melatonin in plants begins with tryptophan and involves at least six enzymes, with different pathways existing. Due to its hydrophilic and lipophilic nature, melatonin can cross almost all cell walls and act on all organs, tissues, and cells.

Melatonin has been detected in almost all plants, and it not only possesses the ability to scavenge free radicals but also regulates plant growth and development [12]. The role of enzymes in melatonin 2. Биосинтез is indispensable, and their localization can characterize the localization of the melatonin synthesis pathway.

The key enzymes are primarily localized in chloroplasts and mitochondria, with a small portion located in the endoplasmic reticulum and cytoplasm. Chloroplasts and mitochondria bear the brunt of oxidative stress within the cell and have a high demand for melatonin, leading to reduced synthesis and transport of melatonin to other cellular compartments. Consequently, melatonin synthesis in other cellular compartments compensates for this deficiency.

For different plants, the primary sites of melatonin synthesis vary, and the localization of key melatonin synthesis enzymes may also differ. Based on current research, the key enzymes involved in melatonin synthesis in several Модель (модель) plants are primarily located in chloroplasts and the cytoplasm, with limited investigation into melatonin synthesis in plant mitochondria.Whether the melatonin synthesis pathway in mitochondria differs from that in chloroplasts, and whether the pathways are complementary or have a sequential relationship, warrants further investigation.

В качестве одного изnovel plant growth regulator, melatonin synthesis regulation is closely associated with plant growth and development and its ability to respond to various stresses. At the transcriptional level, melatonin synthesis exhibits complex regulatory mechanisms.Not only do the gene transcription levels of upstream and downstream synthetic enzymes influence each other, but transcription factors can also bind to enzyme gene promoters to upregulate (or downregulate) the expression of melatonin synthetic enzyme genes, thereby increasing (or decreasing) the levels of synthetic enzyme proteins.

Reducing the ubiquitination and degradation of melatonin synthesis enzymes, prolonging their half-life, and stabilizing the enzymes; interactions between proteins and enzymes within plants to stabilize enzyme proteins can all promote melatonin synthesis.The stringent regulation at the protein level provides strong support for melatonin synthesis and strictly controls melatonin levels in plants, particularly during the conversion from 5-hydroxytryptophan to melatonin. The catalytic activity of SNAT and ASMT is much lower than that of the two enzymes TDC and T5H.

На территории предприятияs also possess enzymes that catalyze the degradation of melatonin, such as M2H, which exhibits higher catalytic activity than SNAT and ASMT [66]. The combination of low biosynthetic activity and high degradation activity explains the low melatonin content in plants [39,67,68].However, maintaining stable melatonin levels is beneficial for plants to achieve optimal growth and development conditions; excessive Производство и продажа or deficiency of melatonin can lead to abnormal growth and affect plants' sensitivity to various biotic or abiotic stresses.

Several other melatonin 2. Биосинтез intermediates play roles in the melatonin biosynthetic pathway, such as 5-hydroxytryptophan, which plays a positive role in melatonin 2. Биосинтез, while 5-hydroxytryptamine inhibits the enzyme activity of SNAT.Future research could explore whether melatonin has alternative biosynthetic pathways to avoid the inhibitory effect of 5-hydroxytryptophan on enzyme activity. Currently, the primary method for determining melatonin content is liquid chromatography-mass spectrometry (LC-MS), but this method cannot accurately analyze melatonin levels in plants.For example, recent measurements of melatonin content in Arabidopsis thaliana ranged from 0.05 to 100 ng/g fresh weight [29,69,70], with a large intermediate range. Therefore, quantitative 3. Анализ of melatonin in plants should be conducted with caution.

In living organisms, melatonin is a multifunctional molecule with important physiological functions.Melatonin and its metabolites act as a series of biological stimuli or signaling molecules, performing important biological functions [12,48]. We need to further investigate the synthesis and regulatory mechanisms of melatonin in plants to better understand plant growth, development, and environmental adaptation mechanisms from the perspective of melatonin, and to apply melatonin in agricultural Производство и продажа.

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