番茄风味和功能性成分研究进展

doi:10.16420/j.issn.0513-353x.2023-0315

摘要/Abstract

摘要：

番茄（Solanum lycopersicum）风味物质主要由可溶性糖（葡萄糖、果糖等）、有机酸（柠檬酸、苹果酸等）和挥发性物质（醇类、酯类等）等构成。番茄还含有多种功能性成分，包括抗坏血酸、番茄红素、γ-氨基丁酸、维生素D和维生素E等，赋予番茄果实重要的保健功能。本文中概述了番茄风味物质和功能性成分的组成、合成途径及调控机制等方面的研究进展，以期为创制富含风味物质和功能性成分的番茄种质并培育风味好的功能性番茄奠定基础。

关键词: 番茄, 风味, 功能性成分, 合成途径, 调控机制, 综述

Abstract:

The flavor substances of tomato（Solanum lycopersicum）in tomatoes are mainly composed of soluble sugars（such as glucose and fructose），organic acids（citric acid and malic acid，etc.） and volatile substances（alcohols and esters，etc.）. Tomato also contains various functional components，such as ascorbic acid，lycopene，γ-aminobutyric acid，vitamin D and vitamin E. These components endow tomato fruit with important health functions. In this review，the composition，synthetic pathway and regulatory mechanism of flavor and functional components have been analyzed in tomato to create tomato germplasm that is rich in flavor substances and functional components through gene editing in the future. This lays the groundwork for cultivating functional tomato with good flavor.

Key words: tomato, flavor, functional components, synthesis pathway, regulatory mechanism, review

沈衡, 王琳, 李骞, 袁守娟, 郑伟, 王涛涛, 叶志彪, 杨长宪. 番茄风味和功能性成分研究进展[J]. 园艺学报, 2024, 51(2): 423-438.

SHEN Heng, WANG Lin, LI Qian, YUAN Shoujuan, ZHENG Wei, WANG Taotao, YE Zhibiao, YANG Changxian. Analyzing Flavor and Functional Components in Tomatoes：a Review[J]. Acta Horticulturae Sinica, 2024, 51(2): 423-438.

图/表 3

表1 番茄21个主要香气成分

Table 1 Overview of the 21 main components of tomato aroma

挥发性化合物Volatile compound	气味描述Odor descriptor
异戊醛3-Methylbutanal	清爽、花香、巧克力、桃、肥腻 Ethereal，aldehydic，chocolate，peach，fatty
2-甲基丁醛2-Methylbutanal	霉味、可可粉、咖啡、坚果 Musty，cocoa，coffee，nutty
3-甲基丁醇3-Methylbutanol	杂醇、油、酒精、威士忌酒、果香、香蕉 Fusel，oil，alcoholic，whiskey，fruity，banana
2-异丁基噻唑2-Isobutylthiazole	清新、芥末、水蜡、番茄、叶片、泥土、青菜、金属 Green，wasabi，privet，tomato，leaf，earthy，vegetable，metallic
1-戊烯-3-酮1-Penten-3-one	辛辣、胡椒、荠菜、大蒜、洋葱 Pungent，peppery，mustard，garlic，onion
顺式-3-乙烯醇 (Z)-3-Hexenal	清新、肥腻、青草、杂草、果味、苹果 Green，fatty，grassy，weedy，fruity，apple
己醛Hexanal	甘甜、杏仁、果味、清新、树叶、苹果、李、青菜 Sweet，almond，fruity，green，leafy，apple，plum，vegetable
反式-2-乙烯醛 (E)-2-Hexenal	略苦、新鲜、树叶、清新、清爽、果味、麻辣、草本 Sharp，fresh，leafy，green，clean，fruity，spicy，herbal
反式-2-庚烯醛 (E)-2-Heptenal	辛辣、清新、青菜、新鲜、肥腻 Pungent，green，vegetable，fresh，fatty
反式-2,4-癸二烯醛 (E, E)-2,4-Decadienal	油性、黄瓜、甜瓜、柑橘、南瓜、坚果、肉 Oily，cucumber，melon，citrus，pumpkin，nut，meat
苯乙醛Phenylacetaldehyde	清新、甘甜、花香、风信子、三叶草、蜂蜜、可可 Green，sweet，floral，hyacinth，clover，honey，cocoa
愈创木酚Guaiacol	酚醛、烟熏、香料、香草、木质 Phenolic，smoke，spice，vanilla，woody
2-苯乙醇2-Phenylethanol	甜味、花香、新鲜、面包、玫瑰、蜂蜜 Sweet，floral，fresh，bready，rose，honey
水杨酸甲酯Methyl salicylate	冬青、薄荷 Wintergreen，mint
B-硝基苯乙烷1-Nitro-2-phenylethane	花朵、香料 Flower，spice
丁香酚Eugenol	甘甜、辛辣、丁香、木质 Sweet，spicy，clove，woody
6-甲基-5庚烯-2酮6-Methyl-5-hepten-2-one	柑橘、清新、霉臭、柠檬、苹果Citrus，green，musty，lemongrass，apple
柠檬醛Geranial	辛辣、柠檬、甘甜 Sharp，lemon，sweet
大马士酮β-Damascenone	苹果、玫瑰、蜂蜜、烟草、甘甜 Apple，rose，honey，tobacco，sweet
香叶基丙酮Geranylacetone	新鲜、清新、果味、蜡质、玫瑰、木质、木兰花、浓香的热带水果Fresh，green，fruity，waxy，rose，woody，magnolia，tropical
β-紫罗兰酮β-Ionone	花香、木质、甜味、果味、浆果、浓香的热带水果、蜂蜡 Floral，woody，sweet，fruity，berry，tropical，beeswax

表1 番茄21个主要香气成分

Table 1 Overview of the 21 main components of tomato aroma

挥发性化合物Volatile compound	气味描述Odor descriptor
异戊醛3-Methylbutanal	清爽、花香、巧克力、桃、肥腻 Ethereal，aldehydic，chocolate，peach，fatty
2-甲基丁醛2-Methylbutanal	霉味、可可粉、咖啡、坚果 Musty，cocoa，coffee，nutty
3-甲基丁醇3-Methylbutanol	杂醇、油、酒精、威士忌酒、果香、香蕉 Fusel，oil，alcoholic，whiskey，fruity，banana
2-异丁基噻唑2-Isobutylthiazole	清新、芥末、水蜡、番茄、叶片、泥土、青菜、金属 Green，wasabi，privet，tomato，leaf，earthy，vegetable，metallic
1-戊烯-3-酮1-Penten-3-one	辛辣、胡椒、荠菜、大蒜、洋葱 Pungent，peppery，mustard，garlic，onion
顺式-3-乙烯醇 (Z)-3-Hexenal	清新、肥腻、青草、杂草、果味、苹果 Green，fatty，grassy，weedy，fruity，apple
己醛Hexanal	甘甜、杏仁、果味、清新、树叶、苹果、李、青菜 Sweet，almond，fruity，green，leafy，apple，plum，vegetable
反式-2-乙烯醛 (E)-2-Hexenal	略苦、新鲜、树叶、清新、清爽、果味、麻辣、草本 Sharp，fresh，leafy，green，clean，fruity，spicy，herbal
反式-2-庚烯醛 (E)-2-Heptenal	辛辣、清新、青菜、新鲜、肥腻 Pungent，green，vegetable，fresh，fatty
反式-2,4-癸二烯醛 (E, E)-2,4-Decadienal	油性、黄瓜、甜瓜、柑橘、南瓜、坚果、肉 Oily，cucumber，melon，citrus，pumpkin，nut，meat
苯乙醛Phenylacetaldehyde	清新、甘甜、花香、风信子、三叶草、蜂蜜、可可 Green，sweet，floral，hyacinth，clover，honey，cocoa
愈创木酚Guaiacol	酚醛、烟熏、香料、香草、木质 Phenolic，smoke，spice，vanilla，woody
2-苯乙醇2-Phenylethanol	甜味、花香、新鲜、面包、玫瑰、蜂蜜 Sweet，floral，fresh，bready，rose，honey
水杨酸甲酯Methyl salicylate	冬青、薄荷 Wintergreen，mint
B-硝基苯乙烷1-Nitro-2-phenylethane	花朵、香料 Flower，spice
丁香酚Eugenol	甘甜、辛辣、丁香、木质 Sweet，spicy，clove，woody
6-甲基-5庚烯-2酮6-Methyl-5-hepten-2-one	柑橘、清新、霉臭、柠檬、苹果Citrus，green，musty，lemongrass，apple
柠檬醛Geranial	辛辣、柠檬、甘甜 Sharp，lemon，sweet
大马士酮β-Damascenone	苹果、玫瑰、蜂蜜、烟草、甘甜 Apple，rose，honey，tobacco，sweet
香叶基丙酮Geranylacetone	新鲜、清新、果味、蜡质、玫瑰、木质、木兰花、浓香的热带水果Fresh，green，fruity，waxy，rose，woody，magnolia，tropical
β-紫罗兰酮β-Ionone	花香、木质、甜味、果味、浆果、浓香的热带水果、蜂蜡 Floral，woody，sweet，fruity，berry，tropical，beeswax

图1 糖、酸和挥发性化合物合成通路（1）γ-氨基丁酸；（2）谷氨酸；（3）α-酮戊二酸；（4）异柠檬酸；（5）柠檬酸钠；（6）草酰乙酸；（7）琥珀酰硫激酶；（8）琥珀酸；（9）延胡索酸；（10）苹果酸；（11）辅酶A；（12）乙酰辅酶A；（13）乌头酸；（14）乙醛酸乙酯；（15）脂肪酸；（16）丙酮酸激酶；（17）蔗糖；（18）果糖；（19）鸟苷二磷酸葡萄糖；（20）果糖-6-磷酸；（21）二磷酸葡萄糖腺苷二钠；（22）D-葡萄糖-6-磷酸；（23）葡萄糖-1-磷酸；（24）6-磷酸葡萄糖酸；（25）淀粉；（26）戊糖磷酸途径；（27）磷酸丙糖；（28）糖酵解；（29）果糖-1,6-双磷酸；（30）淀粉合成酶；（31）磷酸葡萄糖变位酶；（32）蔗糖磷酸合成酶；（33）蔗糖二磷酸酯酶；（34）蔗糖合成酶；（35）可溶性酸化转化酶；（36）细胞壁结合酸性转化酶；（37）卡尔文循环；（38）类胡萝卜素；（39）八氢番茄红素；（40）牻牛儿基牻牛儿基焦磷酸；（41）异戊烯基焦磷酸；（42）二甲基焦磷酸烯丙基；（43）邻苯二甲基酸单乙酯；（44）二叔丁基过氧化物；（45）GPT酶激活蛋白；（46）磷酸烯醇式丙酮酸；（47）1-脱氧木酮糖-5-磷酸合成酶；（48）异戊二烯基二磷酸；（49）加氢脱硫；（50）DNA解旋酶；（51）八氢番茄红素合成酶；（52）八氢番茄红素脱氢酶；（53）ζ-胡萝卜素脱氢酶；（54）脱镁甲基化酶；（55）支链醇类；（56）支链酯类；（57）支链醛；（58）亮氨酸；（59）缬氨酸；（60）异亮氨酸；（61）氧代异戊酸脂；（62）2,3-二羟基异戊酸脂；（63）乙酰肌酸酯；（64）双对苯二甲酸酯；（65）苏氨酸；（66）脯氨酸羟化酶；（67）乙酰乳酸合成酶；（68）支链氨基酸转氨酶；（69）醇酰基转移酶；（70）羧酸酯酶；（71）苯丙氨酸；（72）苯乙胺；（73）E-肉桂酸；（74）醇类；（75）醛类；（76）酰基列胶；（77、78）亚油酸；（79）长链脂质过氧化氢；（80）氢过氧化物的外消旋混合物；（81）酸类；（82）顺-2-戊烯-1-醇；（83）1-辛烯-3-醇；（84）2-戊烯醛；（85）1-戊烯-3-酮；（86）3-己烯醛；（87）2-庚烯醛；（88）己醛；（89）2-壬烯醛；（90）2,4-癸二烯醛；（91）酯类；（92）9-赖氨酸氧化酶；（93）13-赖氨酸氧化酶；（94）脂肪酶；（95）酯氢过氧化物裂解酶；（96）乙醇脱氢酶；（97）β-裂变；（98）自氧化；（99）醛基脱氢酶；（100）谷氨酸循环；（101）三羧酸循环；（102）间类异戊二烯。

Fig. 1 Overview of the sugar，acid and volatile compound VOC pathway （Adapted from Tieman et al.，2012；Martina et al.，2021；陈同强等，2022）

图2 功能性成分合成通路（1）乙酰辅酶A；（2）丙酮酸；（3）3-磷酸甘油醛；（4）异戊烯基焦磷酸，二甲烯丙基焦磷酸；（5）牻牛儿基牻牛儿基焦磷酸；（6）八氢番茄红素；（7）顺式-ζ-胡萝卜素；（8）番茄红素原；（9）番茄红素；（10）β-胡萝卜素；（11）α-胡萝卜素；（12）玉米黄质；（13）叶黄素；（14）草酰乙酸酯；（15）琥珀酸；（16）琥珀酰辅酶A；（17）柠檬酸；（18）α-酮戊二酸；（19）L-谷氨酸；（20）三聚戊酸脱氢酶或转氨酶；（21）二氧化碳；（22）谷氨酸脱羧酶；（23）钙调结构域；（24）γ-氨基丁酸；（25）γ-氨基丁酸转氨酶；（26）琥珀酸半醛；（27）琥珀酸半醛脱氢酶，（28）1-脱氧-D-木醇糖；（29）甲基-D-赤藓醇磷酸盐；（30）异戊烯基焦磷酸；（31）烯丙基磷酸二甲酯；（32）牻牛儿牻牛儿基焦磷酸合成酶；（33）牻牛儿牻牛儿基焦磷酸；（34）植基二磷酸；（35）2-甲基-6-叶绿基-1,4-苯醌；（36）2,3-二甲基-6-叶绿基-1,4-苯醌；（37）均化剂；（38）δ-生育酚；（39）γ-生育酚；（40）β-生育酚；（41）α-生育酚；（42）4-羟基苯基丙酮酸酯；（43）酪氨酸；（44）莽草酸；（45）莽草酸酯；（46）3-脱氢喹酸酯；（47）3-脱氧-D-阿拉伯庚糖酮酸-7-磷酸，（48）膦烯醇丙酮酸；（49）赤藓糖-4-磷酸；（50）2-甲基-6-叶绿基-1,4-苯烯醌；（51）2,3-二甲基-6-叶绿基-1,4-苯烯醌，（52）δ-生育三烯酚；（53）β-生育三烯酚；（54）γ-生育三烯酚；（55）α-生育三烯酚；（56）2-甲基-6-叶绿基-1,4-苯醌甲基转移酶基因，（57）法尼基焦磷酸，（58）牻牛儿牻牛儿基焦磷酸酯；（59）叶绿醌，维生素K；（60）类胡萝卜素，维生素A原；（61）生育色满酚，维生素E，α-生育酚；（62）α-生育酚自由基；（63）单极氢抗坏血酸酯；（64）脱氢抗坏血酸；（65）L-抗坏血酸；（66）还原型辅酶Ⅱ，（67）烟酰胺腺嘌呤二核苷磷酸；（68）L-半乳糖基-1,4-内酯；（69）L-半乳糖；（70）L-半乳糖-1-磷酸；（71）半乳糖磷酸酶，（72）GDP-L-半乳糖磷酸；（73）GDP-甘露糖-3’,5’-异构酶；（74）GDP-D-甘露糖，（75）GDP-甘露糖焦磷酸化酶，（76）D-甘露糖-1-磷酸；（77）D-葡萄糖-6-磷酸；（78）环蒿烯醇；（79）芸苔素类固醇，植物甾醇；（80）维生素D3原，7-脱氢胆固醇；（81）紫外线；（82）7-脱氢胆固醇还原酶，（83）胆固醇；（84）维生素前酯D3；（85）维生素D3，胆钙化醇。

Fig. 2 Functional component synthesis pathway （Adapted from Liu et al.，2015；Munoz & Munne-Bosch，2019；Sarasa et al.，2020；Mellidou et al.，2021；Straeten & Strobbe，2022）

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