园艺植物苦味研究进展

doi:10.16420/j.issn.0513-353x.2024-0067

摘要/Abstract

摘要：

园艺植物食用部位中的苦味对其风味影响较大。本文就苦味的呈味机理、常见园艺植物中苦味物质、苦味评价及苦味代谢物筛选方法等进行了综述，旨在为不同应用目标的园艺植物培育及苦味物质的综合开发利用提供参考。

Abstract:

The bitter taste in the edible parts of horticultural plants significantly impacts their flavor. In this paper，the mechanisms of bitter taste，common bitter substances in horticultural plants，evaluation methods，and screening techniques for bitter metabolites are reviewed. The aim is to serve as a reference for the comprehensive development and utilization of bitter substances in horticultural plant breeding for various application goals.

Key words: horticultural plant, bitter taste, taste mechanism, evaluation methods, bitter metabolites

郑传奇, 唐玉超, 杨盼盼, 彭富海, 徐雷锋, 唐乐, 支永明, 明军. 园艺植物苦味研究进展[J]. 园艺学报, 2024, 51(7): 1529-1546.

ZHENG Chuanqi, TANG Yuchao, YANG Panpan, PENG Fuhai, XU Leifeng, TANG Le, ZHI Yongming, MING Jun. Advances in Bitter Taste of the Horticultural Plants[J]. Acta Horticulturae Sinica, 2024, 51(7): 1529-1546.

图/表 4

参考文献 129

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植物名 Plant name		主要苦味化合物 Main bitter compound	参考文献 Reference
蜜柚、柑橘、脐橙、柠檬Honey pomelo，citrus，navel orange，lemon		柚皮苷、新橙皮苷、枸杞苷、柠檬苦素、宜昌素、诺米林酸、诺米林、奥巴叩酮及配糖体Naringin，neohesperidin，goyaglycoside，limonin，ichangensin，nomilin acid，nomilin，obacunone and glycosome	毕静莹，2019；Huang et al.，2021；余歆等，2021
苹果Apple		根皮素、儿茶素和原花青素Phloretin，catechins and proanthocyanidins	乜兰春和孙建设，2005
桃、李、杏仁 Peach，plum，almond		表儿茶素、槲皮素、绿原酸、新绿原酸、苦杏仁甙、野黑樱苷和洋李甙 Epicatechin，quercetin，chlorogenic acid，neochlorogenic acid，amygdalin，prunasin and prunin	Bolarinwa et al.，2014；Thodberg et al.，2018；张俊环等，2023
红枣、骏枣Red dates，Junzao		没食子酸、原儿茶酸、对羟基苯甲酸、儿茶素、表儿茶素、迷迭酸、鞣花酸、咖啡酸、阿魏酸、香豆酸、香豆素Gallic acid，protocatechuic acid，p-hydroxybenzoic acid，catechin，epicatechin，rosmarinic acid，ellagic acid，caffeic acid，ferulic acid，coumaric acid，coumarin	Xie et al.，2017；蒲云峰，2019
拟南芥、花椰菜、青花菜Arabidopsis thaliana，cauliflower，broccoli	硫代葡萄糖苷、甲状腺肿素、前致甲状腺肿素、芸薹葡糖硫苷、新葡萄糖芸薹素、黑介子苷Glucosinolate，gottrin，progoitrin，glucobrassicin，neoglucobrassicin，sinigrin			Zabaras et al.，2013
莴苣Lettuce	莴苣素、山莴苣苦素Lactucin，lactucopicrin			火国涛等，2023
蒲公英Dandelion	木犀草素、芦丁、槲皮素、萜类成分、蒲公英甾、咖啡酸、绿原酸 Luteolin，rutin，quercetin，terpenoid component，taraxasterol，caffeic acid，chlorogenic acid			石爱文等，2024
苦丁菜 Lxeris denticulata	生物碱、萜类、黄酮类、木犀草素、槲皮素、糖苷类 Alkaloids，terpene，flavanoids，luteolin，quercetin，glycoside			孙为刚，2023
苦瓜、黄瓜、甜瓜 Momordica charantia，cucumber，muskmelon	奎宁葫芦素A、B、C、D和E（其中葫芦素B、D和E在葫芦科植物中普遍存在，葫芦素A和C只在黄瓜属中发现，Momordicoside K、L与Momordicines Ⅰ、Ⅱ在苦瓜中分离得到） Quinine，Cucurbitins A，B，C，D and E（The cucurbitaceae family widely contains cucurbitins B，D，and E，while cucumber exclusively possesses cucurbitins A and C. Additionally，momordicoside K，L along with momordicinesⅠand Ⅱ are specifically isolated from the genus Momordicines.）			Chen，2005；Shang et al.，2014；崔竣杰等，2015；付金玉等，2019；顾兴芳等，2000
马铃薯Potato	糖基生物碱（a-龙葵碱、a-卡茄碱和β2-卡茄碱）Glycoalkaloids（a-Solanine，a-carsolanine and β2-carsolanine）			Zarzecka et al.，2013；叶广继等，2022
番茄Tomato	糖基生物碱（番茄苷）Glycoalkaloids（tomatine）			Tohge et al.，2014
青花椒 Zanthoxylum schinifolium	熊果苷、表儿茶素、金丝桃苷、异槲皮苷、伞形酮、番石榴苷、槲皮素、槲皮苷、山奈酚、异鼠李素、原花青素B2 Arbutin，epicatechin，hyperoside，isoquercitrin，umbelliferone，guajavarin，quercetin，quercitrin，kaempferol，isorhamnetin，proanthocyanidin b2			黄山，2023
红花椒 Pericarpium zanthoxyli bungeani	苯丙氨酸、缬氨酸、异亮氨酸、zanthoamide A、木兰花碱、小檗碱、槲皮素、槲皮苷、槲皮素-3-O-α-L-阿拉伯糖苷、异鼠李素、芥子酸、水杨酸、熊果苷和腺苷 Phenprobamate，valine，l-isoleucine，zanthoamide A，magnolflorine，berbine，quercetin，quercitrin，guajavarin，isorhamnetin，sinapic acid，salicylic acid，arbutin and adenosine			梅小飞，2020
苦茶 Bitter tea	可可碱、缬氨酸、苦茶碱、儿茶素、苦味肽、咖啡因 Theobromine，valine，tetramethyluric acid，catechin，bitter peptide，caffeine theobromine			Yu & Yang，2020；魏沙沙等，2021
绿茶 Green tea	γ-氨基丁酸、咖啡碱、表没食子儿茶素、表没食子儿茶素没食子酸酯、表儿茶素没食子酸酯Caffeine，epigallocatechin，epigallocatechin gallate，epicatechin gallate 4-aminobutyric acid			黄藩等，2023
燕麦 Oat	Avenacoside A、Avenacoside B、燕麦蒽酰胺2c、燕麦蒽酰胺2p、燕麦蒽酰胺1p、9-羟基-顺,反-10,12-十八碳二烯-1-单甘油酯 Avenacoside A，avenacoside B，avenanthramide 2c，avenanthramide 2p，avenanthramide 1p，9-hydroxy-cis,trans-10,12-octadecadiene-1-monoglycerol ester			金文苑，2018；皇甫文倩等，2019
黄连 Coptis chinensis	黄连素Berberine			陈维萍等，2023
厚朴 Mangnolia officinalis	厚朴酚与和厚朴酚Magnolol and honokiol			荆文光等，2022
穿心莲Andrographis paniculata	穿心莲内酯、新穿心莲内酯、14-脱氧穿心莲内酯、脱氢穿心莲内酯 Andrographolide，neoandrographolide，14-deoxyandrographolide，andrographolide			Zhang et al.，2018
桔梗Platycodon grandiflorus	去芹糖桔梗皂苷D、桔梗皂苷D2、桔梗皂苷D、远志皂苷D Desapioplatycodin D，platycodin D2，platycodin D，polygalacin-D3			陆海洋等，2018；张玲等，2019
百合Lily	百合皂苷、儿茶素、表儿茶素、飞燕草色素、咖啡酸、哌啶、根皮素、木犀草素、槲皮素 Lilium saponin，catechin，epicatechin，delphinidin，caffeic acid，piperidine，phloretin，luteolin，quercetin			代丽凤等，2020
吴茱萸 Fructus evodiae	吴茱萸苦素、茱萸苦素乙酸酯、Calodendrolide，6β-乙酰氧基-5-表柠檬苦素、加洁茉里苦素、石虎柠檬素、吴茱萸内酯醇、黄柏酮、evodirutaenin A、格罗苦素甲Rutaevin，rutaevine acetate，calodendrolide，6β-acetoxy-5-epillimonin，jangomolide，limonexic acid，evodol，obacunon，evodirutaenin A，agropicrin			孔奕丹等，2023
氨基酸及其苦味肽 Amino acids and their bitter peptides	人体中必需氨基酸均具有苦味，并以苯丙氨酸和色氨酸的苦味最强；肽分子质量在3 kDa以下时才具有苦味，且苦味强度与疏水氨基酸的含量呈正相关 All essential amino acids in the human body have a bitter taste，with phenylalanine and tryptophan exhibiting the strongest bitterness. Peptides with a molecular weight below 3 kDa are perceived as bitter，and the intensity of bitterness is directly related to the presence of hydrophobic amino acids			彭静和孙威江，2017；田霄艳等，2022

植物名 Plant name		主要苦味化合物 Main bitter compound	参考文献 Reference
蜜柚、柑橘、脐橙、柠檬Honey pomelo，citrus，navel orange，lemon		柚皮苷、新橙皮苷、枸杞苷、柠檬苦素、宜昌素、诺米林酸、诺米林、奥巴叩酮及配糖体Naringin，neohesperidin，goyaglycoside，limonin，ichangensin，nomilin acid，nomilin，obacunone and glycosome	毕静莹，2019；Huang et al.，2021；余歆等，2021
苹果Apple		根皮素、儿茶素和原花青素Phloretin，catechins and proanthocyanidins	乜兰春和孙建设，2005
桃、李、杏仁 Peach，plum，almond		表儿茶素、槲皮素、绿原酸、新绿原酸、苦杏仁甙、野黑樱苷和洋李甙 Epicatechin，quercetin，chlorogenic acid，neochlorogenic acid，amygdalin，prunasin and prunin	Bolarinwa et al.，2014；Thodberg et al.，2018；张俊环等，2023
红枣、骏枣Red dates，Junzao		没食子酸、原儿茶酸、对羟基苯甲酸、儿茶素、表儿茶素、迷迭酸、鞣花酸、咖啡酸、阿魏酸、香豆酸、香豆素Gallic acid，protocatechuic acid，p-hydroxybenzoic acid，catechin，epicatechin，rosmarinic acid，ellagic acid，caffeic acid，ferulic acid，coumaric acid，coumarin	Xie et al.，2017；蒲云峰，2019
拟南芥、花椰菜、青花菜Arabidopsis thaliana，cauliflower，broccoli	硫代葡萄糖苷、甲状腺肿素、前致甲状腺肿素、芸薹葡糖硫苷、新葡萄糖芸薹素、黑介子苷Glucosinolate，gottrin，progoitrin，glucobrassicin，neoglucobrassicin，sinigrin			Zabaras et al.，2013
莴苣Lettuce	莴苣素、山莴苣苦素Lactucin，lactucopicrin			火国涛等，2023
蒲公英Dandelion	木犀草素、芦丁、槲皮素、萜类成分、蒲公英甾、咖啡酸、绿原酸 Luteolin，rutin，quercetin，terpenoid component，taraxasterol，caffeic acid，chlorogenic acid			石爱文等，2024
苦丁菜 Lxeris denticulata	生物碱、萜类、黄酮类、木犀草素、槲皮素、糖苷类 Alkaloids，terpene，flavanoids，luteolin，quercetin，glycoside			孙为刚，2023
苦瓜、黄瓜、甜瓜 Momordica charantia，cucumber，muskmelon	奎宁葫芦素A、B、C、D和E（其中葫芦素B、D和E在葫芦科植物中普遍存在，葫芦素A和C只在黄瓜属中发现，Momordicoside K、L与Momordicines Ⅰ、Ⅱ在苦瓜中分离得到） Quinine，Cucurbitins A，B，C，D and E（The cucurbitaceae family widely contains cucurbitins B，D，and E，while cucumber exclusively possesses cucurbitins A and C. Additionally，momordicoside K，L along with momordicinesⅠand Ⅱ are specifically isolated from the genus Momordicines.）			Chen，2005；Shang et al.，2014；崔竣杰等，2015；付金玉等，2019；顾兴芳等，2000
马铃薯Potato	糖基生物碱（a-龙葵碱、a-卡茄碱和β2-卡茄碱）Glycoalkaloids（a-Solanine，a-carsolanine and β2-carsolanine）			Zarzecka et al.，2013；叶广继等，2022
番茄Tomato	糖基生物碱（番茄苷）Glycoalkaloids（tomatine）			Tohge et al.，2014
青花椒 Zanthoxylum schinifolium	熊果苷、表儿茶素、金丝桃苷、异槲皮苷、伞形酮、番石榴苷、槲皮素、槲皮苷、山奈酚、异鼠李素、原花青素B2 Arbutin，epicatechin，hyperoside，isoquercitrin，umbelliferone，guajavarin，quercetin，quercitrin，kaempferol，isorhamnetin，proanthocyanidin b2			黄山，2023
红花椒 Pericarpium zanthoxyli bungeani	苯丙氨酸、缬氨酸、异亮氨酸、zanthoamide A、木兰花碱、小檗碱、槲皮素、槲皮苷、槲皮素-3-O-α-L-阿拉伯糖苷、异鼠李素、芥子酸、水杨酸、熊果苷和腺苷 Phenprobamate，valine，l-isoleucine，zanthoamide A，magnolflorine，berbine，quercetin，quercitrin，guajavarin，isorhamnetin，sinapic acid，salicylic acid，arbutin and adenosine			梅小飞，2020
苦茶 Bitter tea	可可碱、缬氨酸、苦茶碱、儿茶素、苦味肽、咖啡因 Theobromine，valine，tetramethyluric acid，catechin，bitter peptide，caffeine theobromine			Yu & Yang，2020；魏沙沙等，2021
绿茶 Green tea	γ-氨基丁酸、咖啡碱、表没食子儿茶素、表没食子儿茶素没食子酸酯、表儿茶素没食子酸酯Caffeine，epigallocatechin，epigallocatechin gallate，epicatechin gallate 4-aminobutyric acid			黄藩等，2023
燕麦 Oat	Avenacoside A、Avenacoside B、燕麦蒽酰胺2c、燕麦蒽酰胺2p、燕麦蒽酰胺1p、9-羟基-顺,反-10,12-十八碳二烯-1-单甘油酯 Avenacoside A，avenacoside B，avenanthramide 2c，avenanthramide 2p，avenanthramide 1p，9-hydroxy-cis,trans-10,12-octadecadiene-1-monoglycerol ester			金文苑，2018；皇甫文倩等，2019
黄连 Coptis chinensis	黄连素Berberine			陈维萍等，2023
厚朴 Mangnolia officinalis	厚朴酚与和厚朴酚Magnolol and honokiol			荆文光等，2022
穿心莲Andrographis paniculata	穿心莲内酯、新穿心莲内酯、14-脱氧穿心莲内酯、脱氢穿心莲内酯 Andrographolide，neoandrographolide，14-deoxyandrographolide，andrographolide			Zhang et al.，2018
桔梗Platycodon grandiflorus	去芹糖桔梗皂苷D、桔梗皂苷D2、桔梗皂苷D、远志皂苷D Desapioplatycodin D，platycodin D2，platycodin D，polygalacin-D3			陆海洋等，2018；张玲等，2019
百合Lily	百合皂苷、儿茶素、表儿茶素、飞燕草色素、咖啡酸、哌啶、根皮素、木犀草素、槲皮素 Lilium saponin，catechin，epicatechin，delphinidin，caffeic acid，piperidine，phloretin，luteolin，quercetin			代丽凤等，2020
吴茱萸 Fructus evodiae	吴茱萸苦素、茱萸苦素乙酸酯、Calodendrolide，6β-乙酰氧基-5-表柠檬苦素、加洁茉里苦素、石虎柠檬素、吴茱萸内酯醇、黄柏酮、evodirutaenin A、格罗苦素甲Rutaevin，rutaevine acetate，calodendrolide，6β-acetoxy-5-epillimonin，jangomolide，limonexic acid，evodol，obacunon，evodirutaenin A，agropicrin			孔奕丹等，2023
氨基酸及其苦味肽 Amino acids and their bitter peptides	人体中必需氨基酸均具有苦味，并以苯丙氨酸和色氨酸的苦味最强；肽分子质量在3 kDa以下时才具有苦味，且苦味强度与疏水氨基酸的含量呈正相关 All essential amino acids in the human body have a bitter taste，with phenylalanine and tryptophan exhibiting the strongest bitterness. Peptides with a molecular weight below 3 kDa are perceived as bitter，and the intensity of bitterness is directly related to the presence of hydrophobic amino acids			彭静和孙威江，2017；田霄艳等，2022

奎宁/（× 10^-6mol · L^-1）Quinine	苦味描述 Bitter taste description	苦味值 Bitter tastevalue	盐酸小檗碱/（mmol · L^-1）Berberine hydrochloride	苦味描述 Bitter taste description	苦味等级 Bitterness rating
3	不苦 Imperceptible	0	0	不苦Imperceptible	Ⅰ
6	微苦Slight	1	0.027	微苦Slight	Ⅱ
12	中等苦Moderate	2	0.134	中等苦度Moderate	Ⅲ
24	较苦Quite	3	0.269	很苦，但仍可接受 Great，but acceptable	Ⅳ
48	苦Great	4	1.344	极苦，几乎不能接受 Extreme，almost unacceptable	Ⅴ
96	很苦Extreme	5	—	—	—

奎宁/（× 10^-6mol · L^-1）Quinine	苦味描述 Bitter taste description	苦味值 Bitter tastevalue	盐酸小檗碱/（mmol · L^-1）Berberine hydrochloride	苦味描述 Bitter taste description	苦味等级 Bitterness rating
3	不苦 Imperceptible	0	0	不苦Imperceptible	Ⅰ
6	微苦Slight	1	0.027	微苦Slight	Ⅱ
12	中等苦Moderate	2	0.134	中等苦度Moderate	Ⅲ
24	较苦Quite	3	0.269	很苦，但仍可接受 Great，but acceptable	Ⅳ
48	苦Great	4	1.344	极苦，几乎不能接受 Extreme，almost unacceptable	Ⅴ
96	很苦Extreme	5	—	—	—

分类 Classification	目的 Purpose	优点 Advantage	缺点 Defect
非靶向代谢组学 Untargeted metabolomics	代谢物检测（主要是初级代谢物），寻找潜在生物标记物 Metabolite detection（primarily primary metabolites），the aim of identifying potential biomarkers	无偏向性，覆盖率高，操作简单，可以发现新的标记物 Impartiality，extensive scope，simplicity，and discovery of new markers	灵敏度低，低丰度代谢物检测低，需依赖公共数据库，准确鉴定代谢物难，验证困难 Low sensitivity and abundance，reliance on public databases，accurate identification of metabolites is challenging，and verification is difficult
靶向代谢组学 Targeted metabolomics	少量目标代谢物检测，关注目标通路代谢物，可以进行临床标记物验证，研究调控机制 The detection of a small amount of target metabolites，focusing on the metabolites of the target pathway，can validate them as clinical markers and study the regulatory mechanism	灵敏度高，绝对定性定量 Sensitivity，absolute qualitative，and quantitative	需要标准品，需构建标准曲线Standards are required，and standard curves are constructed
广泛靶向代谢组学 Broadly targeted metabolomics	初生与次生代谢物检测，寻找潜在的生物标记物，关注目标代谢物通路，研究其调控机制Primary and secondary metabolite detection，searching for potential biomarkers，focusing on target metabolite pathways，and studying their regulatory mechanisms	灵敏度高，覆盖率高 High sensitivity and high coverage	相对定性定量，需要数据库，准确鉴定代谢物难 Relatively qualitative and quantitative，reliance on public databases，accurate identification of metabolites is difficult