Research Progress on the Mechanisms of Flavor and Aroma Metabolism Mediated by CCD in Higher Plant

doi:10.16420/j.issn.0513-353x.2026-0112

Acta Horticulturae Sinica ›› 2026, Vol. 53 ›› Issue (10): 3023-3048.doi: 10.16420/j.issn.0513-353x.2026-0112

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Research Progress on the Mechanisms of Flavor and Aroma Metabolism Mediated by CCD in Higher Plant

LIN Mengting¹, FU Xiaopeng¹, JIA Ruidong², ZHAO Xin², GE Hong², KOU Yaping²^,^*(), YANG Shuhua²^,^*()

¹ National Key Laboratory of Germplasm Innovation & Utilization of Horticultural Crops， College of Horticulture and Forestry Science，Huazhong Agricultural University， Wuhan 430070， China
² National Key Laboratory of Vegetable Biotechnology Breeding， Key Laboratory of Floriculture Biology and Germplasm Innovation (North)，Ministry of Agriculture and Rural Affairs，Institute of Vegetables and Flowers，Chinese Academy of Agricultural Sciences， Beijing 100081， China

Received:2026-03-31 Revised:2026-06-09 Online:2026-10-25 Published:2026-10-26
Contact: KOU Yaping, YANG Shuhua

Abstract

Abstract:

Carotenoids are widely distributed in nature，serving as important photosynthetic and chromogenic pigments. They play pivotal roles in regulating plant growth and development，improving flavor quality，and providing essential dietary nutrients for humans. This paper describes carotenoid categories and their roles in higher plant color and flavor，and systematically summarizes the metabolic regulatory network. Then，focusing on carotenoid cleavage dioxygenases（CCDs），this paper highlights the molecular basis of carotenoids effects on higher plant color and aroma metabolism，the molecular mechanisms underlying the coordinated regulation of these traits. This paper aims to provide a theoretical reference for the creation of novel elite germplasms with desirable color and aroma traits in higher plants， and offers guidance for the exploration and utilization of functional plants.

Key words: carotenoids, carotenoid cleavage dioxygenases, color and aroma metabolism

CLC Number:

Q 946

LIN Mengting, FU Xiaopeng, JIA Ruidong, ZHAO Xin, GE Hong, KOU Yaping, YANG Shuhua. Research Progress on the Mechanisms of Flavor and Aroma Metabolism Mediated by CCD in Higher Plant[J]. Acta Horticulturae Sinica, 2026, 53(10): 3023-3048.

Figures/Tables 5

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分类 Type	物种 Species	表型 Phenotype	主要类胡萝卜素 Major carotenoids	主要挥发性降异戊二烯物质 Major volatile norisoprenoids		参考文献 Reference
花 Flower	矮牵牛Petunia hybrida‘Mitchell Diploid’	白色花瓣 White petal	/	β-紫罗兰酮β-ionone		Simkin et al.,2004
	百合Lilium spp.	橙/黄色花瓣Orange/yellow petal	花药黄质、紫黄质、辣椒红素、八氢番茄红素、玉米黄质、β-胡萝卜素Antheraxanthin，violaxanthin，capsanthin，phytoene，zeaxanthin，β-carotene	/		Yamagishi et al.,2010；王欢等,2021
	百日草 Zinnia elegans	黄/红色花瓣Yellow/red petal	β-胡萝卜素β-carotene	/		Qing et al.,2022
	大花蕙兰 Cymbidium hybrida	黄色花瓣Yellow petal	β-胡萝卜素β-carotene	/		Wang et al.,2014
	大马士革蔷薇 Rosa damascena	粉色花瓣 Pink petal	/	β-大马酮、β-大马烯酮、β-紫罗兰酮β-damascone，β-damascenone，β-ionone		Huang et al.,2009
	君子兰Clivia miniata	橙色花瓣Orange petal	黄体素、β-胡萝卜素、玉米黄质Lutein，β-carotene，zeaxanthin	6-甲基-5-庚烯-2-酮、β-紫罗兰酮 6-Methyl-5-hepte-2- one，β-ionone		Li et al.,2022
	滇牡丹Paeonia delavayi	黄色花瓣Yellow petal	黄体素、八氢番茄红素Lutein，phytoene	/		邹红竹等,2021
	番红花Crocus sativus	橙红色花柱 Orange-red stigma	藏红花素Crocin	藏红花醛、异佛尔酮、α-异佛尔酮、4-氧代异佛尔酮、2,6,6-三甲基-1,4-环己二烯-1-羧醛Safranal，isophorone，3,5,5-trimethyl-3-cyclohexen-1-one，2,6,6-trimethyl-2- cyclohexen-1,4-dione，2,6,6-trimethyl- 1,4-cyclo-hexadiene- 1-carboxaldehyde		Cadwallader,2002；Mirbakhsh et al.,2023
花 Flower	甘蓝型油菜Brassica napus	白/黄色花瓣 White/yellow petal	番茄红素、黄体素、β-胡萝卜素 Lycopene，lutein，β-carotene	α-紫罗兰酮α-ionone	曾德志,2013；Zhang et al.,2015
	鹤望兰 Strelitzia reginae	黄色花萼 Yellow sepal	β-胡萝卜素、β-隐黄质、黄体素 β-Carotene，β-cryptoxanthin，lutein	/	樊荣辉等,2016
	角堇Viola cornuta ‘Lutea Splendens’	黄色花瓣 Yellow petal	紫黄质、花药黄质Violaxanthin，antheraxanthin	/	Trajkovic et al.,2021
	金盏花Calendula officinalis	黄/橙色花瓣 Yellow/orange petal	黄体黄质、叶黄呋喃素、番茄红素 Lutein，flavoxanthin，lycopene	/	Kishimoto et al.,2007
	菊花Chrysanthemum morifolium	黄色花瓣 Yellow petal	黄体素、β-胡萝卜素 Lutein，β-carotene	/	Kishimoto & Ohmiya,2006
	香石竹Dianthus caryophyllus L.‘Club’	黄色花瓣 Yellow petal	新黄质、紫黄质、黄体素 Neoxanthin，violaxanthin，lutein	/	Ohmiya et al.,2020
	莲 Nelumbo nucifera	白/红/黄色花瓣 White/red/yellow petal	/	β-紫罗兰酮 β-ionone	王志文等,2023
	龙胆花 Gentiana lutea	黄/橙色花瓣 Yellow/orange petal	黄体素、β-胡萝卜素 Lutein，β-carotene	/	Berman et al.,2016
	美洲莲Nelumbo lutea（American lotus）	黄色花瓣 Yellow petal	β-胡萝卜素、ε-胡萝卜素、黄体素 β-carotene，ε-carotene，lutein	/	Zheng et al.,2022
	南非万寿菊 Osteospermum ecklonis	黄/橙色花瓣 Yellow/orange petal	番茄红素、(5′Z)-γ-胡萝卜素Lycopene，(5'Z)-γ-carotene	/	Kishimoto et al.,2007
	忍冬Lonicera japonica	黄色花瓣 Yellow petal	α-胡萝卜素、玉米黄质、紫黄质、γ-胡萝卜素α-Carotene，zeaxanthin，violaxanthin，γ-carotene	/	Xia et al.,2021
	石斛 Dendrobium nobile	黄色花瓣 Yellow petal	黄体素、α-隐黄质、玉米黄质、花药黄质Lutein，α-cryptoxanthin，zeaxanthin，antheraxanthin	/	黄昕蕾等,2019
	水仙Narcissus pseudonarcissus	白/黄/橙红色花瓣White/yellow/orange- red petal	β-胡萝卜素、黄体素 β-Carotene，lutein	/	Li et al.,2018
	水仙Narcissus tazetta var. chinensis‘Jinzhanyintai’	白色花被片 White tepal	黄体素 Lutein	/	Ren et al.,2017
	万寿菊 Tagetes erecta	黄/橙色花瓣 Yellow/orange petal	黄体素、玉米黄质、紫黄质、新黄质、花药黄质、α-胡萝卜素、β-胡萝卜素、番茄红素、辣椒红素Lutein，zeaxanthin，violaxanthin，neoxanthin，antheraxanthin，α-carotene，β-carotene，lycopene，capsanthin	/	Zhang et al.,2020
	文心兰Oncidium Gower Ramsey	黄色花瓣 Yellow petal	紫黄质、新黄质 Violaxanthin，neoxanthin	/	Hieber et al.,2005
	向日葵 Helianthus annuus	黄色花瓣Yellow petal	黄体素Lutein		Kumar & Gowda,2010
	向日葵 Helianthus annuus	橙色花粉Orange pollen	黄体素酯、α-胡萝卜素、β-胡萝卜素、隐黄质、叶黄呋喃素 Lutein esters，α-carotene，β-carotene，cryptoxanthin，flavoxanthin	/	Bertoli et al.,2011
	勋章菊属 Gazania spp.	黄/橙色花瓣 Yellow/orange petal	黄体素、(5′Z)-玉红黄质 Lutein，(5′Z)-rubixanthin	/	Kishimoto et al.,2007
	郁金香Tulipa gesneriana	黄/红色花瓣 Yellow/red petal	八氢番茄红素、肉豆蔻酸紫黄质酯、棕榈酸紫黄质酯 Phytoene，violaxanthin myristate，violaxanthin palmitate	/	Wang et al.,2022a
	月季Rosa hybrida ‘Yingge Yanwu’	黄色花瓣Yellow petal	β-胡萝卜素、紫黄质β-carotene，violaxanthin	/	Wang et al.,2025
	月季Rosa hybrida ‘YS Group’（‘Yunzheng Xiawei’×‘Sun City’）	黄色花瓣Yellow petal	肉豆蔻酸紫黄质酯 Violaxanthin myristate	/	Cheng et al.,2025
	月季Rosa hybrida ‘YS Group’（‘Yunzheng Xiawei’×‘Sun City’）	白色花瓣White petal	黄体素Lutein	/	Cheng et al.,2025
果实Fruit	西番莲Passiflora edulis	黄色果肉 Yellow flesh	顺式-ζ-胡萝卜素 cis-ζ-carotene	依杜兰异构体 Isomeric edulans	Winterhalter & Rouseff,2002；Wondracek et al.,2011
	番木瓜Carica papaya	黄/红色果肉 Yellow/red flesh	β-隐黄质、β-胡萝卜素、番茄红素 β-cryptoxanthin，β-carotene，lycopene	/	Schweiggert et al.,2011
	番茄Lycopersicon esculentum	红/黄色果肉 Red/yellow flesh	番茄红素、β-胡萝卜素、黄体素、八氢番茄红素、六氢番茄红素 Lycopene，β-carotene，lutein，phytoene，phytofluene	β-紫罗兰酮、β-大马烯酮、6-甲基-5-庚烯-2-酮β-ionone，β-damas- cenone，6-methyl- 5-heptene-2-one	Fraser et al.,2007；高媛等,2023
	凤梨Ananas comusus	黄色果肉 Yellow flesh	紫黄质、9-顺式紫黄质、β-胡萝卜素 Violaxanthin，9-cis-violaxanthin，β-carotene	/	Sugawara et al.,2019
	柑橘Citrus	红/黄色果肉/果皮 Red/yellow flesh/peel	β-隐黄质、紫黄质 β-cryptoxanthin，violaxanthin	/	Kato et al.,2004
	枸杞Lycium chinense	黄色果肉 Yellow flesh	β-胡萝卜素、β-隐黄质、玉米黄质、玉米黄素双棕榈酸酯β-carotene，β-cryptoxanthin，zeaxanthin，zeaxanthin dipalmitate	/	金波等,2025
	红毛丹Nephelium lappaceum	白色果肉 White flesh	/	β-大马烯酮 β-Damascenone	Friedrich & Acree,2002
	辣椒Capsicum annuum	红/橙/黄色果肉 Red/orange/yellow flesh	辣椒红素、玉米黄质、黄体素、紫黄质、β-胡萝卜素、β-隐黄质Capsanthin，zeaxanthin，lutein，violaxanthin，β-carotene，β-cryptoxanthin	/	段敏杰等,2024
	榴莲Durio zibethinus	黄色果肉 Yellow flesh	β-胡萝卜素、α-胡萝卜素、黄体素、玉米黄质β-carotene，α-carotene，lutein，zeaxanthin	/	Wisutiamonkul et al.,2015
	杧果Mangifera indica	黄色果肉/果皮 Yellow flesh/peel	α-胡萝卜素、β-胡萝卜素、番茄红素、β-隐黄质、玉米黄质、黄体素α-carotene，β-carotene，lycopene，β-cryptoxanthin，zeaxanthin，lutein	/	Liang et al.,2019
	猕猴桃Actinidia chinensis	黄/绿色果肉 Yellow/green flesh	β-胡萝卜素，黄体素、紫黄质 β-carotene，lutein，violaxanthin	/	Ampomah-Dwamena et al.,2009
	南瓜Cucurbita moschata	橙色果肉 Orange flesh	β-胡萝卜素、α-胡萝卜素、黄体素 β-carotene，α-carotene，lutein	/	王彬等,2017
	欧洲甜樱桃 Prunus avium	黄色果肉Yellow flesh	黄体素、β-胡萝卜素Lutein，β-carotene	/	Gross,1987
	苹果Malus × domestica	黄色果肉Yellow flesh	黄体素、八氢番茄红素、β-胡萝卜素 Lutein，phytoene，β-carotene	β-大马烯酮 β-damascenone	Friedrich & Acree,2002；Dang et al.,2021
	葡萄Vitis vinifera	黄绿色果肉 Yellow-green flesh	黄体素、β-胡萝卜素、紫黄质、新黄质、玉米黄质、花药黄质Lutein，β-carotene，violaxanthin，neoxanthin，zeaxanthin，antheraxanthin	/	Young et al.,2012
	桃Prunus persica	黄色果肉 Yellow flesh	γ-胡萝卜素、番茄红素、黄体素、α-胡萝卜素、(E/Z)-八氢番茄红素、β-胡萝卜素 γ-carotene，lycopene，lutein，α-carotene，(E/Z)-phytoene，β-carotene	β-紫罗兰酮、β-环柠檬醛、β-大马烯酮β-ionone，β-cyclocitral，β-damascenone	Liu et al.,2025
	甜瓜Cucumis melo	橙/黄色果肉 Orange/yellow flesh	β-胡萝卜素 β-carotene	β-大马烯酮 β-Damascenone	Esteras et al.,2018；Singh et al.,2022
	甜瓜Cucumis melo L. var. makuwa	黄色果肉 Yellow flesh	黄体素、β-胡萝卜素 Lutein，β-Carotene	/	Tuan et al.,2019
	西瓜Citrullus lanatus	红/橙/黄色果肉 Red/orange/yellow flesh	番茄红素、花药黄质、玉米黄质、紫黄质、 β-胡萝卜素、黄体素、β-隐黄质、八氢番茄红素 Lycopene，antheraxanthin，zeaxanthin，violaxanthin，β-carotene，lutein，β-cryptoxanthin，phytoene	香叶基丙酮、β-紫罗兰酮 Geranylacetone，β-ionone	Liu et al.,2012；Fang et al.,2020
	香蕉Musa paradisiaca	黄色果肉/果皮 Yellow flesh/peel	黄体素、α-胡萝卜素、β-胡萝卜素 Lutein，α-carotene，β-carotene	/	Gross,1987
果实Fruit	杏Prunus armeniaca	橙色果肉 Orange flesh	β-胡萝卜素、α-胡萝卜素、八氢番茄红素、 β-隐黄质、黄体素 β-carotene，α-carotene，phytoene，β-cryptoxanthin，lutein	β-紫罗兰酮、二氢-β-紫罗兰酮、6-甲基-5-庚烯-2-酮、β-大马烯酮β-ionone，dihydro- β-ionone，6-methyl-5-hepten- 2-one，β-damascenone	Xi et al.,2020；Zhou et al.,2021
	杨桃Averrhoa carambola	黄色果肉 Yellow flesh	/	β-大马烯酮、大柱-4,6,8-三烯异构体、大柱-4,6, 8-三烯-3-酮、大柱-5,8-二烯-4-酮β-damascenone，isomeric megastigma-4,6,8- trienes，megastigma-4,6,8- triene-3-ones，megastigma-5,8- dien-4-one	Winterhalter & Rouseff,2002
	柚Citrus maxima	红色至橙色果肉 Red to orange flesh	番茄红素、β-胡萝卜素 Lycopene，β-carotene	/	Zhang et al.,2025a
	栀子Gardenia jasminoides	黄色至橙红色果肉Yellow to orange-red flesh	藏红花素 Crocin	/	Pham et al.,2000
种子 Seed	芡欧鼠尾草（RS产区）Salvia Hispanica （grown in Rio Grande do Sul）	棕色种皮 Brown seed coat	黄体素、玉米黄质 Lutein，zeaxanthin	/	Da Silva et al.,2017
	玉米Zea mays	黄色种子Yellow seeds	甜玉米：玉米黄质、黄体素、α-隐黄质 Sweet corn：zeaxanthin，lutein，α-cryptoxanthin	/	陈洁琼等,2015
	玉米Zea mays	黄色种子Yellow seeds	糯玉米：玉米黄质、新黄质、花药黄质 Waxy corn：zeaxanthin，neoxanthin，antheraxanthin	/	陈洁琼等,2015
块根 Tube- rous	甘薯Ipomoea batatas	橙色块根 Orange tuber	β-胡萝卜素、β-隐黄质 β-carotene，β-cryptoxanthin	/	Wei et al.,2025
块根 Tube- rous	胡萝卜Daucus carota	红/橙/黄色块根 Red/orange/yellow tuber	番茄红素、β-胡萝卜素、α-胡萝卜素、黄体素Lycopene，β-carotene，α-carotene，lutein	/	Rodriguez-Concepcion & Stange,2013
块茎 Tuber	马铃薯Solanum tuberosum	白/黄色块茎 White/yellow tuber	紫黄质、花药黄质、黄体素、玉米黄质 Violaxanthin，antheraxanthin，lutein，zeaxanthin	/	Morris et al.,2004
叶Leaf	白菜Brassica rapa L. spp. pekinensis	橙色叶片 Orange leaf	α-胡萝卜素、β-胡萝卜素、番茄红素α-carotene，β-carotene，lycopene	/	马帅等,2021
	菠菜Spinacia oleracea	绿色叶片 Green leaf	黄体素、β-胡萝卜素、紫黄质、新黄质Lutein，β-carotene，violaxanthin，neoxanthin	呋喃衍生物 Furan derivative	Winterhalter & Rouseff,2002；Bunea et al.,2008
	茶Camellia sinensis	绿色叶片 Green leaf	β-胡萝卜素、黄体素β-carotene，lutein	/	Wang et al.,2009
	茶Camellia sinensis	加工叶片 Processed leaf	/	β-紫罗兰酮、β-大马烯酮、茶螺烯酮β-ionone，β-damascenone，theaspirone	陈丽等,2019

分类 Type	物种 Species	表型 Phenotype	主要类胡萝卜素 Major carotenoids	主要挥发性降异戊二烯物质 Major volatile norisoprenoids		参考文献 Reference
花 Flower	矮牵牛Petunia hybrida‘Mitchell Diploid’	白色花瓣 White petal	/	β-紫罗兰酮β-ionone		Simkin et al.,2004
	百合Lilium spp.	橙/黄色花瓣Orange/yellow petal	花药黄质、紫黄质、辣椒红素、八氢番茄红素、玉米黄质、β-胡萝卜素Antheraxanthin，violaxanthin，capsanthin，phytoene，zeaxanthin，β-carotene	/		Yamagishi et al.,2010；王欢等,2021
	百日草 Zinnia elegans	黄/红色花瓣Yellow/red petal	β-胡萝卜素β-carotene	/		Qing et al.,2022
	大花蕙兰 Cymbidium hybrida	黄色花瓣Yellow petal	β-胡萝卜素β-carotene	/		Wang et al.,2014
	大马士革蔷薇 Rosa damascena	粉色花瓣 Pink petal	/	β-大马酮、β-大马烯酮、β-紫罗兰酮β-damascone，β-damascenone，β-ionone		Huang et al.,2009
	君子兰Clivia miniata	橙色花瓣Orange petal	黄体素、β-胡萝卜素、玉米黄质Lutein，β-carotene，zeaxanthin	6-甲基-5-庚烯-2-酮、β-紫罗兰酮 6-Methyl-5-hepte-2- one，β-ionone		Li et al.,2022
	滇牡丹Paeonia delavayi	黄色花瓣Yellow petal	黄体素、八氢番茄红素Lutein，phytoene	/		邹红竹等,2021
	番红花Crocus sativus	橙红色花柱 Orange-red stigma	藏红花素Crocin	藏红花醛、异佛尔酮、α-异佛尔酮、4-氧代异佛尔酮、2,6,6-三甲基-1,4-环己二烯-1-羧醛Safranal，isophorone，3,5,5-trimethyl-3-cyclohexen-1-one，2,6,6-trimethyl-2- cyclohexen-1,4-dione，2,6,6-trimethyl- 1,4-cyclo-hexadiene- 1-carboxaldehyde		Cadwallader,2002；Mirbakhsh et al.,2023
花 Flower	甘蓝型油菜Brassica napus	白/黄色花瓣 White/yellow petal	番茄红素、黄体素、β-胡萝卜素 Lycopene，lutein，β-carotene	α-紫罗兰酮α-ionone	曾德志,2013；Zhang et al.,2015
	鹤望兰 Strelitzia reginae	黄色花萼 Yellow sepal	β-胡萝卜素、β-隐黄质、黄体素 β-Carotene，β-cryptoxanthin，lutein	/	樊荣辉等,2016
	角堇Viola cornuta ‘Lutea Splendens’	黄色花瓣 Yellow petal	紫黄质、花药黄质Violaxanthin，antheraxanthin	/	Trajkovic et al.,2021
	金盏花Calendula officinalis	黄/橙色花瓣 Yellow/orange petal	黄体黄质、叶黄呋喃素、番茄红素 Lutein，flavoxanthin，lycopene	/	Kishimoto et al.,2007
	菊花Chrysanthemum morifolium	黄色花瓣 Yellow petal	黄体素、β-胡萝卜素 Lutein，β-carotene	/	Kishimoto & Ohmiya,2006
	香石竹Dianthus caryophyllus L.‘Club’	黄色花瓣 Yellow petal	新黄质、紫黄质、黄体素 Neoxanthin，violaxanthin，lutein	/	Ohmiya et al.,2020
	莲 Nelumbo nucifera	白/红/黄色花瓣 White/red/yellow petal	/	β-紫罗兰酮 β-ionone	王志文等,2023
	龙胆花 Gentiana lutea	黄/橙色花瓣 Yellow/orange petal	黄体素、β-胡萝卜素 Lutein，β-carotene	/	Berman et al.,2016
	美洲莲Nelumbo lutea（American lotus）	黄色花瓣 Yellow petal	β-胡萝卜素、ε-胡萝卜素、黄体素 β-carotene，ε-carotene，lutein	/	Zheng et al.,2022
	南非万寿菊 Osteospermum ecklonis	黄/橙色花瓣 Yellow/orange petal	番茄红素、(5′Z)-γ-胡萝卜素Lycopene，(5'Z)-γ-carotene	/	Kishimoto et al.,2007
	忍冬Lonicera japonica	黄色花瓣 Yellow petal	α-胡萝卜素、玉米黄质、紫黄质、γ-胡萝卜素α-Carotene，zeaxanthin，violaxanthin，γ-carotene	/	Xia et al.,2021
	石斛 Dendrobium nobile	黄色花瓣 Yellow petal	黄体素、α-隐黄质、玉米黄质、花药黄质Lutein，α-cryptoxanthin，zeaxanthin，antheraxanthin	/	黄昕蕾等,2019
	水仙Narcissus pseudonarcissus	白/黄/橙红色花瓣White/yellow/orange- red petal	β-胡萝卜素、黄体素 β-Carotene，lutein	/	Li et al.,2018
	水仙Narcissus tazetta var. chinensis‘Jinzhanyintai’	白色花被片 White tepal	黄体素 Lutein	/	Ren et al.,2017
	万寿菊 Tagetes erecta	黄/橙色花瓣 Yellow/orange petal	黄体素、玉米黄质、紫黄质、新黄质、花药黄质、α-胡萝卜素、β-胡萝卜素、番茄红素、辣椒红素Lutein，zeaxanthin，violaxanthin，neoxanthin，antheraxanthin，α-carotene，β-carotene，lycopene，capsanthin	/	Zhang et al.,2020
	文心兰Oncidium Gower Ramsey	黄色花瓣 Yellow petal	紫黄质、新黄质 Violaxanthin，neoxanthin	/	Hieber et al.,2005
	向日葵 Helianthus annuus	黄色花瓣Yellow petal	黄体素Lutein		Kumar & Gowda,2010
	向日葵 Helianthus annuus	橙色花粉Orange pollen	黄体素酯、α-胡萝卜素、β-胡萝卜素、隐黄质、叶黄呋喃素 Lutein esters，α-carotene，β-carotene，cryptoxanthin，flavoxanthin	/	Bertoli et al.,2011
	勋章菊属 Gazania spp.	黄/橙色花瓣 Yellow/orange petal	黄体素、(5′Z)-玉红黄质 Lutein，(5′Z)-rubixanthin	/	Kishimoto et al.,2007
	郁金香Tulipa gesneriana	黄/红色花瓣 Yellow/red petal	八氢番茄红素、肉豆蔻酸紫黄质酯、棕榈酸紫黄质酯 Phytoene，violaxanthin myristate，violaxanthin palmitate	/	Wang et al.,2022a
	月季Rosa hybrida ‘Yingge Yanwu’	黄色花瓣Yellow petal	β-胡萝卜素、紫黄质β-carotene，violaxanthin	/	Wang et al.,2025
	月季Rosa hybrida ‘YS Group’（‘Yunzheng Xiawei’×‘Sun City’）	黄色花瓣Yellow petal	肉豆蔻酸紫黄质酯 Violaxanthin myristate	/	Cheng et al.,2025
	月季Rosa hybrida ‘YS Group’（‘Yunzheng Xiawei’×‘Sun City’）	白色花瓣White petal	黄体素Lutein	/	Cheng et al.,2025
果实Fruit	西番莲Passiflora edulis	黄色果肉 Yellow flesh	顺式-ζ-胡萝卜素 cis-ζ-carotene	依杜兰异构体 Isomeric edulans	Winterhalter & Rouseff,2002；Wondracek et al.,2011
	番木瓜Carica papaya	黄/红色果肉 Yellow/red flesh	β-隐黄质、β-胡萝卜素、番茄红素 β-cryptoxanthin，β-carotene，lycopene	/	Schweiggert et al.,2011
	番茄Lycopersicon esculentum	红/黄色果肉 Red/yellow flesh	番茄红素、β-胡萝卜素、黄体素、八氢番茄红素、六氢番茄红素 Lycopene，β-carotene，lutein，phytoene，phytofluene	β-紫罗兰酮、β-大马烯酮、6-甲基-5-庚烯-2-酮β-ionone，β-damas- cenone，6-methyl- 5-heptene-2-one	Fraser et al.,2007；高媛等,2023
	凤梨Ananas comusus	黄色果肉 Yellow flesh	紫黄质、9-顺式紫黄质、β-胡萝卜素 Violaxanthin，9-cis-violaxanthin，β-carotene	/	Sugawara et al.,2019
	柑橘Citrus	红/黄色果肉/果皮 Red/yellow flesh/peel	β-隐黄质、紫黄质 β-cryptoxanthin，violaxanthin	/	Kato et al.,2004
	枸杞Lycium chinense	黄色果肉 Yellow flesh	β-胡萝卜素、β-隐黄质、玉米黄质、玉米黄素双棕榈酸酯β-carotene，β-cryptoxanthin，zeaxanthin，zeaxanthin dipalmitate	/	金波等,2025
	红毛丹Nephelium lappaceum	白色果肉 White flesh	/	β-大马烯酮 β-Damascenone	Friedrich & Acree,2002
	辣椒Capsicum annuum	红/橙/黄色果肉 Red/orange/yellow flesh	辣椒红素、玉米黄质、黄体素、紫黄质、β-胡萝卜素、β-隐黄质Capsanthin，zeaxanthin，lutein，violaxanthin，β-carotene，β-cryptoxanthin	/	段敏杰等,2024
	榴莲Durio zibethinus	黄色果肉 Yellow flesh	β-胡萝卜素、α-胡萝卜素、黄体素、玉米黄质β-carotene，α-carotene，lutein，zeaxanthin	/	Wisutiamonkul et al.,2015
	杧果Mangifera indica	黄色果肉/果皮 Yellow flesh/peel	α-胡萝卜素、β-胡萝卜素、番茄红素、β-隐黄质、玉米黄质、黄体素α-carotene，β-carotene，lycopene，β-cryptoxanthin，zeaxanthin，lutein	/	Liang et al.,2019
	猕猴桃Actinidia chinensis	黄/绿色果肉 Yellow/green flesh	β-胡萝卜素，黄体素、紫黄质 β-carotene，lutein，violaxanthin	/	Ampomah-Dwamena et al.,2009
	南瓜Cucurbita moschata	橙色果肉 Orange flesh	β-胡萝卜素、α-胡萝卜素、黄体素 β-carotene，α-carotene，lutein	/	王彬等,2017
	欧洲甜樱桃 Prunus avium	黄色果肉Yellow flesh	黄体素、β-胡萝卜素Lutein，β-carotene	/	Gross,1987
	苹果Malus × domestica	黄色果肉Yellow flesh	黄体素、八氢番茄红素、β-胡萝卜素 Lutein，phytoene，β-carotene	β-大马烯酮 β-damascenone	Friedrich & Acree,2002；Dang et al.,2021
	葡萄Vitis vinifera	黄绿色果肉 Yellow-green flesh	黄体素、β-胡萝卜素、紫黄质、新黄质、玉米黄质、花药黄质Lutein，β-carotene，violaxanthin，neoxanthin，zeaxanthin，antheraxanthin	/	Young et al.,2012
	桃Prunus persica	黄色果肉 Yellow flesh	γ-胡萝卜素、番茄红素、黄体素、α-胡萝卜素、(E/Z)-八氢番茄红素、β-胡萝卜素 γ-carotene，lycopene，lutein，α-carotene，(E/Z)-phytoene，β-carotene	β-紫罗兰酮、β-环柠檬醛、β-大马烯酮β-ionone，β-cyclocitral，β-damascenone	Liu et al.,2025
	甜瓜Cucumis melo	橙/黄色果肉 Orange/yellow flesh	β-胡萝卜素 β-carotene	β-大马烯酮 β-Damascenone	Esteras et al.,2018；Singh et al.,2022
	甜瓜Cucumis melo L. var. makuwa	黄色果肉 Yellow flesh	黄体素、β-胡萝卜素 Lutein，β-Carotene	/	Tuan et al.,2019
	西瓜Citrullus lanatus	红/橙/黄色果肉 Red/orange/yellow flesh	番茄红素、花药黄质、玉米黄质、紫黄质、 β-胡萝卜素、黄体素、β-隐黄质、八氢番茄红素 Lycopene，antheraxanthin，zeaxanthin，violaxanthin，β-carotene，lutein，β-cryptoxanthin，phytoene	香叶基丙酮、β-紫罗兰酮 Geranylacetone，β-ionone	Liu et al.,2012；Fang et al.,2020
	香蕉Musa paradisiaca	黄色果肉/果皮 Yellow flesh/peel	黄体素、α-胡萝卜素、β-胡萝卜素 Lutein，α-carotene，β-carotene	/	Gross,1987
果实Fruit	杏Prunus armeniaca	橙色果肉 Orange flesh	β-胡萝卜素、α-胡萝卜素、八氢番茄红素、 β-隐黄质、黄体素 β-carotene，α-carotene，phytoene，β-cryptoxanthin，lutein	β-紫罗兰酮、二氢-β-紫罗兰酮、6-甲基-5-庚烯-2-酮、β-大马烯酮β-ionone，dihydro- β-ionone，6-methyl-5-hepten- 2-one，β-damascenone	Xi et al.,2020；Zhou et al.,2021
	杨桃Averrhoa carambola	黄色果肉 Yellow flesh	/	β-大马烯酮、大柱-4,6,8-三烯异构体、大柱-4,6, 8-三烯-3-酮、大柱-5,8-二烯-4-酮β-damascenone，isomeric megastigma-4,6,8- trienes，megastigma-4,6,8- triene-3-ones，megastigma-5,8- dien-4-one	Winterhalter & Rouseff,2002
	柚Citrus maxima	红色至橙色果肉 Red to orange flesh	番茄红素、β-胡萝卜素 Lycopene，β-carotene	/	Zhang et al.,2025a
	栀子Gardenia jasminoides	黄色至橙红色果肉Yellow to orange-red flesh	藏红花素 Crocin	/	Pham et al.,2000
种子 Seed	芡欧鼠尾草（RS产区）Salvia Hispanica （grown in Rio Grande do Sul）	棕色种皮 Brown seed coat	黄体素、玉米黄质 Lutein，zeaxanthin	/	Da Silva et al.,2017
	玉米Zea mays	黄色种子Yellow seeds	甜玉米：玉米黄质、黄体素、α-隐黄质 Sweet corn：zeaxanthin，lutein，α-cryptoxanthin	/	陈洁琼等,2015
	玉米Zea mays	黄色种子Yellow seeds	糯玉米：玉米黄质、新黄质、花药黄质 Waxy corn：zeaxanthin，neoxanthin，antheraxanthin	/	陈洁琼等,2015
块根 Tube- rous	甘薯Ipomoea batatas	橙色块根 Orange tuber	β-胡萝卜素、β-隐黄质 β-carotene，β-cryptoxanthin	/	Wei et al.,2025
块根 Tube- rous	胡萝卜Daucus carota	红/橙/黄色块根 Red/orange/yellow tuber	番茄红素、β-胡萝卜素、α-胡萝卜素、黄体素Lycopene，β-carotene，α-carotene，lutein	/	Rodriguez-Concepcion & Stange,2013
块茎 Tuber	马铃薯Solanum tuberosum	白/黄色块茎 White/yellow tuber	紫黄质、花药黄质、黄体素、玉米黄质 Violaxanthin，antheraxanthin，lutein，zeaxanthin	/	Morris et al.,2004
叶Leaf	白菜Brassica rapa L. spp. pekinensis	橙色叶片 Orange leaf	α-胡萝卜素、β-胡萝卜素、番茄红素α-carotene，β-carotene，lycopene	/	马帅等,2021
	菠菜Spinacia oleracea	绿色叶片 Green leaf	黄体素、β-胡萝卜素、紫黄质、新黄质Lutein，β-carotene，violaxanthin，neoxanthin	呋喃衍生物 Furan derivative	Winterhalter & Rouseff,2002；Bunea et al.,2008
	茶Camellia sinensis	绿色叶片 Green leaf	β-胡萝卜素、黄体素β-carotene，lutein	/	Wang et al.,2009
	茶Camellia sinensis	加工叶片 Processed leaf	/	β-紫罗兰酮、β-大马烯酮、茶螺烯酮β-ionone，β-damascenone，theaspirone	陈丽等,2019

分类Type	物种 Species	蛋白功能 Function of protein					参考文献Reference
分类Type	物种 Species	基因名称 Gene name		作用底物 Substrate	裂解位点 Cleavage site	裂解产物 Cleavage product	参考文献Reference
花 Flower	甘蓝型油菜 Brassica napus	BnaC3.CCD4		α-胡萝卜素和/或β-胡萝卜素α-carotene and/or β-carotene	/	α-紫罗兰酮α-Ionone	Zhang et al.,2015；梁波,2023
		Bna.CCD4		β-胡萝卜素β-Carotene	9,10	C14双醛 C14 dialdehyde
		Bna.CCD4		8′-阿朴-β-胡萝卜醛8′-apo-β-carotenal	9,10	C17双醛 C17 dialdehyde
	君子兰 Clivia miniata	CmCCD1a、CmCCD1b		番茄红素 Lycopene	5,6/5′,6′	6-甲基-5-庚烯-2-酮 6-Methyl-5-heptene-2-one	Li et al.,2022
	君子兰 Clivia miniata	CmCCD1a、CmCCD1b		β-胡萝卜素 β-carotene	9,10/9′,10′	β-紫罗兰酮 β-Ionone	Li et al.,2022
	莲Nelumbo nucifera	NnCCD4		β-胡萝卜素β-Carotene	/	β-紫罗兰酮 β-Ionone	王志文等,2023
	忍冬 Lonicera japonica	LjCCD4		黄体素、β-胡萝卜素 Lutein，β-carotene	9,10/9′,10′	10′-阿朴-β-胡萝卜醛、3-羟基-10′-阿朴-β-胡萝卜醛、3-羟基-10′-阿朴-α-胡萝卜醛10'-apo-β-carotenal, 3-hydroxy-10'-apo-β-carotenal, 3-hydroxy-10'-apo-α-carotenal	Pu et al.,2020
		LjCCD1b		黄体素、β-胡萝卜素 Lutein，β-carotene	9,10/9′,10′	C14双醛 C14 dialdehyde
		LjCCD1b		10′-阿朴-β-胡萝卜醛10′-apo-β-carotenal	9,10	C14双醛 C14 dialdehyde
	铁皮石斛 Dendrobium officinale	DoCCD1		番茄红素Lycopene	5,6/5′,6′	6-甲基-5-庚烯-2-酮 6-Methyl-5-heptene-2-one	Wang et al., 2022b
		DoCCD1		番茄红素Lycopene	9,10/9′,10′	假紫罗兰酮Pseudoionone
		DoCCD1		β-胡萝卜素β-Carotene	9,10/9′,10′	β-紫罗兰酮β-Ionone
	向日葵 Helianthus annuus	HaCCD1		β-胡萝卜素β-Carotene	9,10/9′,10′	β-紫罗兰酮β-Ionone	Qi et al.,2022
		HaCCD1		玉米黄质Zeaxanthin	9,10/9′,10′	3-羟基-β-紫罗兰酮 3-Hydroxy-β-ionone
		HaCCD1		虾青素Astaxanthin	9,10/9′,10′	3-羟基-4-氧代-β-紫罗兰酮3-Hydroxy-4-oxo-β-ionone
		HaCCD1		番茄红素Lycopene	9,10/9′,10′	假紫罗兰酮Pseudoionone
		HaCCD1		黄体素Lutein	9,10/9′,10′	3-Hydroxy-β-ionone
	香雪兰 Freesia hybrida	FhCCD2		玉米黄质 Zeaxanthin	7,8/7′,8′	3-羟基-β-环柠檬醛、藏花酸二醛3-Hydroxy-β-cyclocitral, crocetin dialdehyde	Fang et al.,2020
	月季Rosa chinensis	RcCCD4		β-胡萝卜素 β-Carotene	/	二氢-β-紫罗兰酮 Dihydro-β-ionone	付琪等,2025
果实 Fruit	大豆Glycine max	GmCCD4		β-胡萝卜素/α-胡萝卜素β-Carotene/α-carotene	9,10/9′,10′	β-紫罗兰酮 β-Ionone	Gao et al.,2021
	多毛番茄 Solanum habrochaites	ShCCD4b		ζ-胡萝卜素ζ-carotene	9,10/9′,10′	香叶酮Geranylacetone	Yoo et al.,2023
				β-胡萝卜素β-carotene	9,10/9′,10′	β-紫罗兰酮β-Ionone
	黄果枸杞 Lycium barbarum	LbCCD1		玉米黄质Zeaxanthin	9,10/9′,10′	3-羟基-β-紫罗兰酮 3-Hydroxy-β-ionone	Ke et al.,2024
				虾青素Astaxanthin	9,10/9′,10′	3-羟基-4-氧代-β-紫罗兰酮3-Hydroxy-4-oxo-β-ionone
				8′-阿朴-β-胡萝卜醛8'-apo-β-carotenal	9,10/9′,10′	β-紫罗兰酮 β-Ionone
	枸杞Lycium chinense	LcCCD1		β-胡萝卜素β-Carotene	/	β-紫罗兰酮β-Ionone	田小卫,2016
				番茄红素Lycopene	/	假紫罗兰酮Pseudoionone
		LcCCD4		β-胡萝卜素β-Carotene	/	β-紫罗兰酮β-Ionone
	辣椒 Capsicum annuum	CaCCD1A、CaCCD1B		β-胡萝卜素β-Carotene	9,10/9′,10′	α-紫罗兰酮α-Ionone	Cheng et al.,2021
	粟Setaria italica	SiCCD1		玉米黄质Zeaxanthin	/	/	He et al.,2022
果实 Fruit	无花果Ficus carica	FcCCD1A	环状β-胡萝卜素 Cyclic β-carotene		9,10/9′,10′	α-紫罗兰酮、β-紫罗兰酮 α-Iionone，β-ionone	Nawade et al.,2019
		FcCCD1A	δ-胡萝卜素δ-Carotene		9,10	α-紫罗兰酮 α-Ionone
		FcCCD1B	番茄红素、δ-胡萝卜素中的无环片段 Lycopene，an acyclic moiety of δ-carotene		5,6/5′,6′	6-甲基-5-庚烯-2-酮 6-Methyl-5-heptene-2-one
	小果野芭蕉 Musa acuminata‘Rasthali’	RAS-CCD4	β-胡萝卜素 β-Carotene		9,10	β-紫罗兰酮、10′-阿朴-β-胡萝卜醛β-Ionone，10′-apo-β-carotenal	Awasthi et al.,2022
		RAS-CCD4	10′-阿朴-β-胡萝卜醛 10’-apo-β-carotenal		9′,10′	β-紫罗兰酮、C14双醛 β-Ionone，C14 dialdehyde
	小麦Triticum spp.	TaCCD1	8′-阿朴-β-胡萝卜醛 8'-apo-β-carotenal		9,10/9′,10′	β-紫罗兰酮、10,8′-阿朴- β-胡萝卜二醛 β-Ionone，10,8′-apo-β-carotenedial	Qin et al.,2016
		TaCCD1	β-胡萝卜素β-Carotene		9,10/9′,10′	β-紫罗兰酮 β-Ionone
		TaCCD1	黄体素Lutein		9,10/9′,10′	3-羟基-α-紫罗兰酮、3-羟基-β-紫罗兰酮3-Hydroxy-α- ionone，3-hydroxy-β-ionone
		TaCCD1	玉米黄质Zeaxanthin		9,10/9′,10′	3-羟基-β-紫罗兰酮 3-Hydroxy-β-ionone
	栀子Gardenia jasminoides	GjCCD4a	玉米黄质 Zeaxanthin		7,8/7′,8′	3-羟基-β-环柠檬醛、藏花酸二醛3-Hydroxy-β-cyclocitral，crocetin dialdehyde	Zheng et al., 2022
		GjCCD4a	β-胡萝卜素β-Carotene		7,8/7′,8′	β-环柠檬醛、藏花酸二醛 β-cyclocitral, crocetin dialdehyde
块根Tuberous	胡萝卜 Daucus carota	DcCCD4	α-胡萝卜素 α-Carotene		9,10/9′,10′	α-紫罗兰酮、β-紫罗兰酮 α-Ionone，β-ionone	Li et al.,2021
块根Tuberous	胡萝卜 Daucus carota	DcCCD4	β-胡萝卜素β-Carotene		9,10/9′,10′	β-紫罗兰酮 β-Ionone
叶 Leaf	茶Camellia sinensis	CsCCD1a	β-胡萝卜素β-Carotene		9,10/9′,10′	β-紫罗兰酮 β-Ionone	Wang et al.,2020
	稻Oryza sativa	AtCCD4	β-胡萝卜素β-Carotene		9,10/9′,10′	β-紫罗兰酮 β-Ionone	Song et al.,2016
					7,8/7′,8′	β-环柠檬醛β-Cyclocitral	Song et al.,2016
		OsCCD4b	β-胡萝卜素、玉米黄质 β-Carotene，zeaxanthin		7′,8′	C30、C10类胡萝卜素裂解物 C30，C10 apocarotenoids	Zhang et al.,2025b
		ZAS	10′-阿朴-玉米黄质醛10′-apo-zeaxanthinal		13,14	扎西诺酮 Zaxinone	Wang et al.,2019
	大叶醉鱼草 Buddleja davidii	BdCCD4	玉米黄质Zeaxanthin		7,8/7′,8′	/	Ahrazem et al.,2017
	烟草Nicotiana tabacum	NtCCD1	β-胡萝卜素β-Carotene		/	β-紫罗兰酮β-Ionone	都菲,2021；Li et al.,2022
		NtCCD10	八氢番茄红素Phytoene		9,10/9′,10′	香叶基丙酮Geranylacetone
		NtCCD10	β-胡萝卜素β-Carotene		9,10/9′,10′	β-紫罗兰酮β-Ionone
		NtCCD10	番茄红素Lycopene		5,6/5′,6′	6-甲基-5-庚烯-2-酮 6-Methyl-5-hepten-2-one

分类Type	物种 Species	蛋白功能 Function of protein					参考文献Reference
分类Type	物种 Species	基因名称 Gene name		作用底物 Substrate	裂解位点 Cleavage site	裂解产物 Cleavage product	参考文献Reference
花 Flower	甘蓝型油菜 Brassica napus	BnaC3.CCD4		α-胡萝卜素和/或β-胡萝卜素α-carotene and/or β-carotene	/	α-紫罗兰酮α-Ionone	Zhang et al.,2015；梁波,2023
		Bna.CCD4		β-胡萝卜素β-Carotene	9,10	C14双醛 C14 dialdehyde
		Bna.CCD4		8′-阿朴-β-胡萝卜醛8′-apo-β-carotenal	9,10	C17双醛 C17 dialdehyde
	君子兰 Clivia miniata	CmCCD1a、CmCCD1b		番茄红素 Lycopene	5,6/5′,6′	6-甲基-5-庚烯-2-酮 6-Methyl-5-heptene-2-one	Li et al.,2022
	君子兰 Clivia miniata	CmCCD1a、CmCCD1b		β-胡萝卜素 β-carotene	9,10/9′,10′	β-紫罗兰酮 β-Ionone	Li et al.,2022
	莲Nelumbo nucifera	NnCCD4		β-胡萝卜素β-Carotene	/	β-紫罗兰酮 β-Ionone	王志文等,2023
	忍冬 Lonicera japonica	LjCCD4		黄体素、β-胡萝卜素 Lutein，β-carotene	9,10/9′,10′	10′-阿朴-β-胡萝卜醛、3-羟基-10′-阿朴-β-胡萝卜醛、3-羟基-10′-阿朴-α-胡萝卜醛10'-apo-β-carotenal, 3-hydroxy-10'-apo-β-carotenal, 3-hydroxy-10'-apo-α-carotenal	Pu et al.,2020
		LjCCD1b		黄体素、β-胡萝卜素 Lutein，β-carotene	9,10/9′,10′	C14双醛 C14 dialdehyde
		LjCCD1b		10′-阿朴-β-胡萝卜醛10′-apo-β-carotenal	9,10	C14双醛 C14 dialdehyde
	铁皮石斛 Dendrobium officinale	DoCCD1		番茄红素Lycopene	5,6/5′,6′	6-甲基-5-庚烯-2-酮 6-Methyl-5-heptene-2-one	Wang et al., 2022b
		DoCCD1		番茄红素Lycopene	9,10/9′,10′	假紫罗兰酮Pseudoionone
		DoCCD1		β-胡萝卜素β-Carotene	9,10/9′,10′	β-紫罗兰酮β-Ionone
	向日葵 Helianthus annuus	HaCCD1		β-胡萝卜素β-Carotene	9,10/9′,10′	β-紫罗兰酮β-Ionone	Qi et al.,2022
		HaCCD1		玉米黄质Zeaxanthin	9,10/9′,10′	3-羟基-β-紫罗兰酮 3-Hydroxy-β-ionone
		HaCCD1		虾青素Astaxanthin	9,10/9′,10′	3-羟基-4-氧代-β-紫罗兰酮3-Hydroxy-4-oxo-β-ionone
		HaCCD1		番茄红素Lycopene	9,10/9′,10′	假紫罗兰酮Pseudoionone
		HaCCD1		黄体素Lutein	9,10/9′,10′	3-Hydroxy-β-ionone
	香雪兰 Freesia hybrida	FhCCD2		玉米黄质 Zeaxanthin	7,8/7′,8′	3-羟基-β-环柠檬醛、藏花酸二醛3-Hydroxy-β-cyclocitral, crocetin dialdehyde	Fang et al.,2020
	月季Rosa chinensis	RcCCD4		β-胡萝卜素 β-Carotene	/	二氢-β-紫罗兰酮 Dihydro-β-ionone	付琪等,2025
果实 Fruit	大豆Glycine max	GmCCD4		β-胡萝卜素/α-胡萝卜素β-Carotene/α-carotene	9,10/9′,10′	β-紫罗兰酮 β-Ionone	Gao et al.,2021
	多毛番茄 Solanum habrochaites	ShCCD4b		ζ-胡萝卜素ζ-carotene	9,10/9′,10′	香叶酮Geranylacetone	Yoo et al.,2023
				β-胡萝卜素β-carotene	9,10/9′,10′	β-紫罗兰酮β-Ionone
	黄果枸杞 Lycium barbarum	LbCCD1		玉米黄质Zeaxanthin	9,10/9′,10′	3-羟基-β-紫罗兰酮 3-Hydroxy-β-ionone	Ke et al.,2024
				虾青素Astaxanthin	9,10/9′,10′	3-羟基-4-氧代-β-紫罗兰酮3-Hydroxy-4-oxo-β-ionone
				8′-阿朴-β-胡萝卜醛8'-apo-β-carotenal	9,10/9′,10′	β-紫罗兰酮 β-Ionone
	枸杞Lycium chinense	LcCCD1		β-胡萝卜素β-Carotene	/	β-紫罗兰酮β-Ionone	田小卫,2016
				番茄红素Lycopene	/	假紫罗兰酮Pseudoionone
		LcCCD4		β-胡萝卜素β-Carotene	/	β-紫罗兰酮β-Ionone
	辣椒 Capsicum annuum	CaCCD1A、CaCCD1B		β-胡萝卜素β-Carotene	9,10/9′,10′	α-紫罗兰酮α-Ionone	Cheng et al.,2021
	粟Setaria italica	SiCCD1		玉米黄质Zeaxanthin	/	/	He et al.,2022
果实 Fruit	无花果Ficus carica	FcCCD1A	环状β-胡萝卜素 Cyclic β-carotene		9,10/9′,10′	α-紫罗兰酮、β-紫罗兰酮 α-Iionone，β-ionone	Nawade et al.,2019
		FcCCD1A	δ-胡萝卜素δ-Carotene		9,10	α-紫罗兰酮 α-Ionone
		FcCCD1B	番茄红素、δ-胡萝卜素中的无环片段 Lycopene，an acyclic moiety of δ-carotene		5,6/5′,6′	6-甲基-5-庚烯-2-酮 6-Methyl-5-heptene-2-one
	小果野芭蕉 Musa acuminata‘Rasthali’	RAS-CCD4	β-胡萝卜素 β-Carotene		9,10	β-紫罗兰酮、10′-阿朴-β-胡萝卜醛β-Ionone，10′-apo-β-carotenal	Awasthi et al.,2022
		RAS-CCD4	10′-阿朴-β-胡萝卜醛 10’-apo-β-carotenal		9′,10′	β-紫罗兰酮、C14双醛 β-Ionone，C14 dialdehyde
	小麦Triticum spp.	TaCCD1	8′-阿朴-β-胡萝卜醛 8'-apo-β-carotenal		9,10/9′,10′	β-紫罗兰酮、10,8′-阿朴- β-胡萝卜二醛 β-Ionone，10,8′-apo-β-carotenedial	Qin et al.,2016
		TaCCD1	β-胡萝卜素β-Carotene		9,10/9′,10′	β-紫罗兰酮 β-Ionone
		TaCCD1	黄体素Lutein		9,10/9′,10′	3-羟基-α-紫罗兰酮、3-羟基-β-紫罗兰酮3-Hydroxy-α- ionone，3-hydroxy-β-ionone
		TaCCD1	玉米黄质Zeaxanthin		9,10/9′,10′	3-羟基-β-紫罗兰酮 3-Hydroxy-β-ionone
	栀子Gardenia jasminoides	GjCCD4a	玉米黄质 Zeaxanthin		7,8/7′,8′	3-羟基-β-环柠檬醛、藏花酸二醛3-Hydroxy-β-cyclocitral，crocetin dialdehyde	Zheng et al., 2022
		GjCCD4a	β-胡萝卜素β-Carotene		7,8/7′,8′	β-环柠檬醛、藏花酸二醛 β-cyclocitral, crocetin dialdehyde
块根Tuberous	胡萝卜 Daucus carota	DcCCD4	α-胡萝卜素 α-Carotene		9,10/9′,10′	α-紫罗兰酮、β-紫罗兰酮 α-Ionone，β-ionone	Li et al.,2021
块根Tuberous	胡萝卜 Daucus carota	DcCCD4	β-胡萝卜素β-Carotene		9,10/9′,10′	β-紫罗兰酮 β-Ionone
叶 Leaf	茶Camellia sinensis	CsCCD1a	β-胡萝卜素β-Carotene		9,10/9′,10′	β-紫罗兰酮 β-Ionone	Wang et al.,2020
	稻Oryza sativa	AtCCD4	β-胡萝卜素β-Carotene		9,10/9′,10′	β-紫罗兰酮 β-Ionone	Song et al.,2016
					7,8/7′,8′	β-环柠檬醛β-Cyclocitral	Song et al.,2016
		OsCCD4b	β-胡萝卜素、玉米黄质 β-Carotene，zeaxanthin		7′,8′	C30、C10类胡萝卜素裂解物 C30，C10 apocarotenoids	Zhang et al.,2025b
		ZAS	10′-阿朴-玉米黄质醛10′-apo-zeaxanthinal		13,14	扎西诺酮 Zaxinone	Wang et al.,2019
	大叶醉鱼草 Buddleja davidii	BdCCD4	玉米黄质Zeaxanthin		7,8/7′,8′	/	Ahrazem et al.,2017
	烟草Nicotiana tabacum	NtCCD1	β-胡萝卜素β-Carotene		/	β-紫罗兰酮β-Ionone	都菲,2021；Li et al.,2022
		NtCCD10	八氢番茄红素Phytoene		9,10/9′,10′	香叶基丙酮Geranylacetone
		NtCCD10	β-胡萝卜素β-Carotene		9,10/9′,10′	β-紫罗兰酮β-Ionone
		NtCCD10	番茄红素Lycopene		5,6/5′,6′	6-甲基-5-庚烯-2-酮 6-Methyl-5-hepten-2-one

Research Progress on the Mechanisms of Flavor and Aroma Metabolism Mediated by CCD in Higher Plant

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