苹果类胡萝卜素代谢调控及其抗逆相关机制研究进展

doi:10.16420/j.issn.0513-353x.2026-0307

摘要/Abstract

摘要：

苹果（Malus × domestica Borkh.）是全球重要的果树资源，果实品质与抗逆性协同改良是产业可持续发展的关键。类胡萝卜素作为决定果实色泽、营养品质及抗逆性的重要代谢物，其代谢调控机制是研究热点。本文中系统综述了苹果类胡萝卜素核心代谢途径及其关键酶的功能鉴定，从转录水平、激素信号及表观遗传层面梳理了其生物合成的多层次调控机制，探讨了类胡萝卜素通过抗氧化特性及脱落酸（ABA）信号通路介导抗逆响应的分子机制，总结了关键基因的遗传改良成效，以及多基因协同转化技术在苹果虾青素合成中的应用进展。未来应重点聚焦多组学整合解析、高效基因编辑体系构建及果实品质与抗逆协同改良等研究，以期为解析苹果果实品质形成机制及分子育种提供理论参考。

关键词: 苹果, 类胡萝卜素, 代谢调控, 抗逆, 遗传改良

Abstract:

Apple（Malus × domestica Borkh.）is an important fruit tree resource in the world，and the synergistic improvement of fruit quality and stress resistance is the key to the sustainable development of the industry. Carotenoids are crucial metabolites that determine fruit color，nutritional quality，and stress resistance，and their metabolic regulatory mechanisms are a research hotspot. In this paper，the main metabolic pathways of apple carotenoids and the functional identification of their key enzymes are systematically reviewed. The multi-level regulatory network underlying its biosynthesis is reviewed from the aspects of transcription factors，hormone signals，and epigenetics. The molecular mechanisms of carotenoids mediated stress response through antioxidant properties and abscisic acid（ABA）signaling pathway are discussed. The genetic improvement of key genes and the application progress of multi-gene co-transformation technology in apple astaxanthin synthesis are summarized. Future research focuses on the integrated analysis of multi-omics，the construction of efficient gene-editing system，and the synergistic improvement of fruit quality and stress resistance. These efforts will provide a theoretical basis for elucidating the regulatory mechanisms underlying fruit quality formation and facilitate molecular breeding in apple.

Key words: apple, carotenoid, metabolic regulation, stress resistance, genetic improvement

郭旭东, 王慧敏, 王馨苑, 李淳妤, 莫美华, 许艺淑, 韩晓雅, 原永兵, 贾东杰. 苹果类胡萝卜素代谢调控及其抗逆相关机制研究进展[J]. 园艺学报, 2026, 53(7): 1910-1928.

GUO Xudong, WANG Huimin, WANG Xinyuan, LI Chunyu, MO Meihua, XU Yishu, HAN Xiaoya, YUAN Yongbing, JIA Dongjie. Research Progress on Carotenoid Metabolism and its Role in Stress Tolerance in Apple[J]. Acta Horticulturae Sinica, 2026, 53(7): 1910-1928.

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酶 Enzyme	基因 Gene	验证体系 Validation system	主要功能与表型 Main function and phenotype	参考文献 Reference
八氢番茄红素合成酶 Phytoene synthase	MdPSY1，MdPSY2	番茄、苹果愈伤、大肠杆菌 Tomato，apple callus，Escherichia coli	催化GGPP转化为15-顺式-八氢番茄红素；过表达MdPSY1增加类胡萝卜素积累并加速叶绿体向有色体转化；过表达MdPSY2增加八氢番茄红素和β-胡萝卜素的积累量 Catalyzes GGPP to 15-cis-phytoene；overexpression of MdPSY1 increases carotenoid accumulation and accelerates chloroplast-to-chromoplast transition to enhance carotenoid storage capacity；overexpression of MdPSY2 promotes accumulation of phytoene and β-carotene	Ampomah-Dwamena et al.,2015,2022；Dang et al.,2021
八氢番茄红素脱氢酶 Phytoene desaturase	MdPDS	苹果砧木‘JM2’、苹果愈伤 Apple rootstock ‘JM2’，apple callus	催化15-顺式-八氢番茄红素转化为9,15,9’-三-顺式- ζ-胡萝卜素；CRISPR/Cas9敲除MdPDS导致白化株系（单靶点31.8%，双靶点84%）；miRNA沉默MdPDS获得71%白化株系 Catalyzes 15-cis-phytoene to 9,15,9’-tri-cis-ζ-carotene；CRISPR/Cas9-mediated knockout of MdPDS results albino lines（31.8% for single target，84% for dual targets）；miRNA-mediated silencing of MdPDS results in 71% albino lines	Nishitani et al.,2016；Charrier et al.,2019a,2019b
15-顺式-ζ-胡萝卜素异构酶 15-cis-ζ-Carotene isomerase	MdZISO	苹果果皮（表达分析） Apple skin （expression analysis）	催化9,15,9’-三-顺式-ζ-胡萝卜素转化为9,9’-二-顺式-ζ-胡萝卜素；MdZISO的表达水平与果实最终类胡萝卜素含量高度正相关，在盛花后20 d表达量最高，且该时期的表达量对后期类胡萝卜素的积累具有预测价值 Catalyzes 9,15,9’-tri-cis-ζ-carotene to 9,9’-di-cis-ζ-carotene；MdZISO expression is highly positively correlated with final carotenoid content，peaking at 20 days after full bloom，and its expression level at this stage predicts subsequent accumulation	Ampomah-Dwamena et al.,2012；Cho et al.,2020,2021
ζ-胡萝卜素脱氢酶 ζ-Carotene desaturase	MdZDS	苹果愈伤、拟南芥、烟草 Apple callus，Arabidopsis，tobacco	催化9,9’-二-顺式-ζ-胡萝卜素转化7,9,9’,7’-四顺式-番茄红素；过表达MdZDS提高类胡萝卜素含量，增强耐盐碱和抗氧化能力 Catalyzes 9,9’-di-cis-ζ-carotene to 7,9,9’,7’-tetra-cis-lycopene；overexpression of MdZDS increases carotenoid content and enhances salt-alkali tolerance and antioxidant capacity	Wang et al.,2023
类胡萝卜素异构酶 Carotene isomerase	MdCRTISO	苹果愈伤、苹果果皮 Apple callus，apple skin	催化7,9,9’,7’-四顺式-番茄红素转化为全反式-番茄红素；过表达MdCRTISO使总类胡萝卜素提高1.4倍，提高ABA积累水平 Catalyzes 7,9,9’,7’-tetra-cis-lycopene to all-trans-lycopene；overexpression of MdCRTISO increases total carotenoids by 1.4-fold and promotes ABA accumulation	Wang et al.,2023；魏佳琦等,2024
番茄红素ε-环化酶 Lycopene ε-cyclase	MdLCYE	苹果愈伤、苹果果皮 Apple callus，apple skin	催化全反式-番茄红素转化为α-胡萝卜素；过表达MdLCYE促进总类胡萝卜素和ABA的积累，干扰其表达则结果相反 Catalyzes all-trans-lycopene to α-carotene；overexpression of MdLCYE increases total carotenoid and ABA accumulation，while silencing it produces the opposite effect	Wang et al.,2023
番茄红素β-环化酶 Lycopene β-cyclase	MdLCYb	苹果愈伤 Apple callus	催化全反式-番茄红素转化为β-胡萝卜素；过表达MdLCYb提升β-胡萝卜素、紫黄质及新黄质含量，并降低八氢番茄红素含量 Catalyzes all-trans-lycopene to β-carotene；overexpression of MdLCYb increases the contents of β-carotene，violaxanthin，and neoxanthin，yet reduces phytoene content	Jia et al.,2024
细胞色素P450羟化酶 Cytochrome P450 carotene hydroxylase	MdCYP97A3	苹果愈伤 Apple callus	催化α-胡萝卜素转化为叶黄素；过表达MdCYP97A3提高叶黄素含量，沉默则降低 Catalyzes α-carotene to lutein；overexpression of MdCYP97A3 increases lutein content，while silencing decreases it	Huang et al.,2024
β-胡萝卜素羟化酶 β-Carotene hydroxylase	MdBCH1，MdBCH2	拟南芥、苹果愈伤 Arabidopsis，apple callus	催化β-胡萝卜素转化为玉米黄质；过表达MdBCH1提高玉米黄质含量，增强干旱和渗透胁迫抗性 Catalyzes β-carotene to zeaxanthin；overexpression of MdBCH1 increases zeaxanthin content and improves tolerance to drought and osmotic stress	田雨晨,2025
类胡萝卜素裂解双加氧酶 Carotenoid cleavage dioxygenase	MdCCD4	大肠杆菌、苹果愈伤 Escherichia coli，apple callus	裂解β-胡萝卜素生成β-紫罗兰酮；过表达MdCCD4降低类胡萝卜素含量，沉默则升高 Cleaves β-carotene to form β-ionone；overexpression of MdCCD4 decreases carotenoid content，whereas silencing increases it	Huang et al.,2009,2024
类胡萝卜素裂解双加氧酶 Carotenoid cleavage dioxygenase	MdCCD7，MdCCD8	拟南芥、苹果植株 Arabidopsis，apple plant	裂解β-胡萝卜素，参与独脚金内酯合成；MdCCD7和MdCCD8可分别恢复拟南芥max3（CCD7缺陷）和max4（CCD8缺陷）突变体的分枝缺陷；且沉默MdCCD7侧枝数量显著增加，沉默MdCCD8表型轻微存在功能冗余 Cleaves β-carotene and participates in strigolactone biosynthesis；MdCCD7 and MdCCD8 can rescue the branching defects of Arabidopsis max3 and max4 mutants，respectively；silencing MdCCD7 markedly increases lateral branch number，while silencing MdCCD8 shows mild phenotypes due to functional redundancy	Foster et al.,2018
9-顺式-环氧类胡萝卜素双加氧酶 9-cis-Epoxycarotenoid dioxygenase	MdNCED1，MdNCED3，MpNCED1，MpNCED2，MhNCED3	拟南芥 Arabidopsis	裂解紫黄质和新黄质生成ABA前体；MdNCED3表达与ABA含量正相关；MpNCED2在果实发育、干旱、高温和低温胁迫下持续表达，而MpNCED1仅响应高温和低温胁迫；过表达MhNCED3提高ABA含量，增强对渗透胁迫、Cd²⁺胁迫和Cl^-胁迫的抗性 Cleaves violaxanthin and neoxanthin to produce ABA precursors；MdNCED3 expression is positively correlated with ABA content；MpNCED2 is continuously expressed during fruit development and under drought，high and low temperature stresses，while MpNCED1 only responds to high and low temperature stresses；overexpression of MhNCED3 elevates ABA content and enhances resistance to osmotic，Cd²⁺ and Cl^- stresses	姜志昂等,2014；Xia et al.,2014；Zhang et al.,2014；张玮玮,2015；Li et al.,2020；李嘉伟等,2025

酶 Enzyme	基因 Gene	验证体系 Validation system	主要功能与表型 Main function and phenotype	参考文献 Reference
八氢番茄红素合成酶 Phytoene synthase	MdPSY1，MdPSY2	番茄、苹果愈伤、大肠杆菌 Tomato，apple callus，Escherichia coli	催化GGPP转化为15-顺式-八氢番茄红素；过表达MdPSY1增加类胡萝卜素积累并加速叶绿体向有色体转化；过表达MdPSY2增加八氢番茄红素和β-胡萝卜素的积累量 Catalyzes GGPP to 15-cis-phytoene；overexpression of MdPSY1 increases carotenoid accumulation and accelerates chloroplast-to-chromoplast transition to enhance carotenoid storage capacity；overexpression of MdPSY2 promotes accumulation of phytoene and β-carotene	Ampomah-Dwamena et al.,2015,2022；Dang et al.,2021
八氢番茄红素脱氢酶 Phytoene desaturase	MdPDS	苹果砧木‘JM2’、苹果愈伤 Apple rootstock ‘JM2’，apple callus	催化15-顺式-八氢番茄红素转化为9,15,9’-三-顺式- ζ-胡萝卜素；CRISPR/Cas9敲除MdPDS导致白化株系（单靶点31.8%，双靶点84%）；miRNA沉默MdPDS获得71%白化株系 Catalyzes 15-cis-phytoene to 9,15,9’-tri-cis-ζ-carotene；CRISPR/Cas9-mediated knockout of MdPDS results albino lines（31.8% for single target，84% for dual targets）；miRNA-mediated silencing of MdPDS results in 71% albino lines	Nishitani et al.,2016；Charrier et al.,2019a,2019b
15-顺式-ζ-胡萝卜素异构酶 15-cis-ζ-Carotene isomerase	MdZISO	苹果果皮（表达分析） Apple skin （expression analysis）	催化9,15,9’-三-顺式-ζ-胡萝卜素转化为9,9’-二-顺式-ζ-胡萝卜素；MdZISO的表达水平与果实最终类胡萝卜素含量高度正相关，在盛花后20 d表达量最高，且该时期的表达量对后期类胡萝卜素的积累具有预测价值 Catalyzes 9,15,9’-tri-cis-ζ-carotene to 9,9’-di-cis-ζ-carotene；MdZISO expression is highly positively correlated with final carotenoid content，peaking at 20 days after full bloom，and its expression level at this stage predicts subsequent accumulation	Ampomah-Dwamena et al.,2012；Cho et al.,2020,2021
ζ-胡萝卜素脱氢酶 ζ-Carotene desaturase	MdZDS	苹果愈伤、拟南芥、烟草 Apple callus，Arabidopsis，tobacco	催化9,9’-二-顺式-ζ-胡萝卜素转化7,9,9’,7’-四顺式-番茄红素；过表达MdZDS提高类胡萝卜素含量，增强耐盐碱和抗氧化能力 Catalyzes 9,9’-di-cis-ζ-carotene to 7,9,9’,7’-tetra-cis-lycopene；overexpression of MdZDS increases carotenoid content and enhances salt-alkali tolerance and antioxidant capacity	Wang et al.,2023
类胡萝卜素异构酶 Carotene isomerase	MdCRTISO	苹果愈伤、苹果果皮 Apple callus，apple skin	催化7,9,9’,7’-四顺式-番茄红素转化为全反式-番茄红素；过表达MdCRTISO使总类胡萝卜素提高1.4倍，提高ABA积累水平 Catalyzes 7,9,9’,7’-tetra-cis-lycopene to all-trans-lycopene；overexpression of MdCRTISO increases total carotenoids by 1.4-fold and promotes ABA accumulation	Wang et al.,2023；魏佳琦等,2024
番茄红素ε-环化酶 Lycopene ε-cyclase	MdLCYE	苹果愈伤、苹果果皮 Apple callus，apple skin	催化全反式-番茄红素转化为α-胡萝卜素；过表达MdLCYE促进总类胡萝卜素和ABA的积累，干扰其表达则结果相反 Catalyzes all-trans-lycopene to α-carotene；overexpression of MdLCYE increases total carotenoid and ABA accumulation，while silencing it produces the opposite effect	Wang et al.,2023
番茄红素β-环化酶 Lycopene β-cyclase	MdLCYb	苹果愈伤 Apple callus	催化全反式-番茄红素转化为β-胡萝卜素；过表达MdLCYb提升β-胡萝卜素、紫黄质及新黄质含量，并降低八氢番茄红素含量 Catalyzes all-trans-lycopene to β-carotene；overexpression of MdLCYb increases the contents of β-carotene，violaxanthin，and neoxanthin，yet reduces phytoene content	Jia et al.,2024
细胞色素P450羟化酶 Cytochrome P450 carotene hydroxylase	MdCYP97A3	苹果愈伤 Apple callus	催化α-胡萝卜素转化为叶黄素；过表达MdCYP97A3提高叶黄素含量，沉默则降低 Catalyzes α-carotene to lutein；overexpression of MdCYP97A3 increases lutein content，while silencing decreases it	Huang et al.,2024
β-胡萝卜素羟化酶 β-Carotene hydroxylase	MdBCH1，MdBCH2	拟南芥、苹果愈伤 Arabidopsis，apple callus	催化β-胡萝卜素转化为玉米黄质；过表达MdBCH1提高玉米黄质含量，增强干旱和渗透胁迫抗性 Catalyzes β-carotene to zeaxanthin；overexpression of MdBCH1 increases zeaxanthin content and improves tolerance to drought and osmotic stress	田雨晨,2025
类胡萝卜素裂解双加氧酶 Carotenoid cleavage dioxygenase	MdCCD4	大肠杆菌、苹果愈伤 Escherichia coli，apple callus	裂解β-胡萝卜素生成β-紫罗兰酮；过表达MdCCD4降低类胡萝卜素含量，沉默则升高 Cleaves β-carotene to form β-ionone；overexpression of MdCCD4 decreases carotenoid content，whereas silencing increases it	Huang et al.,2009,2024
类胡萝卜素裂解双加氧酶 Carotenoid cleavage dioxygenase	MdCCD7，MdCCD8	拟南芥、苹果植株 Arabidopsis，apple plant	裂解β-胡萝卜素，参与独脚金内酯合成；MdCCD7和MdCCD8可分别恢复拟南芥max3（CCD7缺陷）和max4（CCD8缺陷）突变体的分枝缺陷；且沉默MdCCD7侧枝数量显著增加，沉默MdCCD8表型轻微存在功能冗余 Cleaves β-carotene and participates in strigolactone biosynthesis；MdCCD7 and MdCCD8 can rescue the branching defects of Arabidopsis max3 and max4 mutants，respectively；silencing MdCCD7 markedly increases lateral branch number，while silencing MdCCD8 shows mild phenotypes due to functional redundancy	Foster et al.,2018
9-顺式-环氧类胡萝卜素双加氧酶 9-cis-Epoxycarotenoid dioxygenase	MdNCED1，MdNCED3，MpNCED1，MpNCED2，MhNCED3	拟南芥 Arabidopsis	裂解紫黄质和新黄质生成ABA前体；MdNCED3表达与ABA含量正相关；MpNCED2在果实发育、干旱、高温和低温胁迫下持续表达，而MpNCED1仅响应高温和低温胁迫；过表达MhNCED3提高ABA含量，增强对渗透胁迫、Cd²⁺胁迫和Cl^-胁迫的抗性 Cleaves violaxanthin and neoxanthin to produce ABA precursors；MdNCED3 expression is positively correlated with ABA content；MpNCED2 is continuously expressed during fruit development and under drought，high and low temperature stresses，while MpNCED1 only responds to high and low temperature stresses；overexpression of MhNCED3 elevates ABA content and enhances resistance to osmotic，Cd²⁺ and Cl^- stresses	姜志昂等,2014；Xia et al.,2014；Zhang et al.,2014；张玮玮,2015；Li et al.,2020；李嘉伟等,2025