类胡萝卜素调控植物花色和果色机制研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2025-1235

摘要/Abstract

摘要：

类胡萝卜素是形成园艺植物花器官和果实色泽的重要色素，直接影响产品的观赏和经济价值。与叶片等光合器官不同，植物花、果发育着色过程中，类胡萝卜素的积累常伴随叶绿素降解及类黄酮等次生代谢物的显著变化，该过程受多层级调控网络协同调控。近年来，类胡萝卜素介导园艺植物花果着色的分子调控机制的研究发展迅速。本文全面总结该领域研究进展，明确PSY等类胡萝卜素合成关键酶基因和有色体分化与色素储存在代谢通路多样性中的核心作用。该代谢过程受到转录、转录后及表观遗传层面协同调控。在转录调控水平，MYB、bHLH、NAC、WRKY、AP2/ERF等转录因子构成复杂调控网络，通过靶向PSY、LCYB、CCD4等关键基因精准维持类胡萝卜素合成与降解的动态平衡。转录后调控层面，ORANGE蛋白对PSY的稳定作用、泛素化介导的蛋白降解、PPR蛋白调控以及miRNA介导的转录后沉默等多种机制，协同实现代谢通量的精细微调。表观遗传方面，DNA甲基化与组蛋白修饰动态调控关键基因染色质状态，影响色素积累的时空模式。同时，光照、温度等环境因子与乙烯、脱落酸等植物激素信号可整合至上述调控网络，共同决定花果颜色。本文全面梳理该多层面协同互作调控体系，深入揭示类胡萝卜素在花果着色中的合成与降解平衡机制，并基于此分析园艺产品色泽改良与分子育种实践中的关键问题及发展路径，为园艺产品色泽性状的精准调控与品质提升提供理论支撑和技术指导。

关键词: 类胡萝卜素, 花色, 果色, 转录调控, 表观遗传调控, 转录后调控, 代谢途径

Abstract:

Carotenoids are important pigments that determine the color of floral organs and fruits in horticultural plants, directly influencing their ornamental and economic value. Unlike that in photosynthetic organs such as leaves, the accumulation of carotenoids during the development and coloration of plant flowers and fruits is often accompanied by significant changes in chlorophyll degradation and levels of secondary metabolites such as flavonoids, a process that is coordinately regulated by a multi-level regulatory network. In recent years, research on the molecular regulatory mechanisms of carotenoid-mediated flower and fruit coloration in horticultural plants has developed rapidly. This paper comprehensively summarizes research progress in this field, clarifying the core roles of key enzymes encoded by carotenoid biosynthetic genes such as PSY and the processes of chromoplast differentiation and pigment storage in the diversity of metabolic pathways. This metabolic process is coordinately regulated at the transcriptional, post-transcriptional, and epigenetic levels. At the transcriptional level, transcription factors such as MYB, bHLH, NAC, WRKY, and AP2/ERF form complex regulatory networks, precisely maintaining the dynamic balance between carotenoid biosynthesis and degradation by targeting key genes including PSY, LCYB, and CCD4. At the post-transcriptional level, multiple mechanisms, including the stabilization of PSY by ORANGE proteins, ubiquitination-mediated protein degradation, PPR protein regulation, and miRNA-mediated post-transcriptional silencing, coordinately achieve fine-tuning of metabolic flux. Epigenetically, DNA methylation and histone modifications dynamically regulate the chromatin state of key genes, thereby affecting the spatiotemporal patterns of pigment accumulation. Meanwhile, environmental factors such as light and temperature, as well as phytohormone signals like ethylene and abscisic acid, can be integrated into the above regulatory network to jointly determine flower and fruit color. This paper comprehensively reviews this multi-level coordinated and interactive regulatory system, deeply reveals the balance mechanism of carotenoid biosynthesis and degradation in flower and fruit coloration, and based on this, analyzes key issues and development pathways for color improvement and molecular breeding of horticultural products, providing theoretical support and technical guidance for the precise regulation of color traits and quality improvement of horticultural products.

Key words: carotenoids, flower color, fruit color, transcriptional regulation, epigenetic regulation, post-transcriptional regulation, metabolic pathways

李富伟, 李宏宇. 类胡萝卜素调控植物花色和果色机制研究进展[J]. 生物技术通报, 2026, 42(10): 13-26.

LI Fu-wei, LI Hong-yu. Research Progress in the Mechanism of Carotenoids Regulating Plant Flower Color and Fruit Color[J]. Biotechnology Bulletin, 2026, 42(10): 13-26.

图/表 3

图1 类胡萝卜素生物合成途径类胡萝卜素在质体中合成，经GGPPS催化生成前体GGPP，再经PSY缩合及多步脱氢异构反应生成番茄红素，随后分为β，β和β，ε两个分支。β，β分支中，LCYB催化生成β-胡萝卜素，进一步衍生出玉米黄质、紫黄质、新黄质及脱落酸等；β，ε分支中，LCYE与LCYB协同催化生成α-胡萝卜素并最终形成叶黄素。箭头代表酶促代谢反应方向，黑色箭头代表直接调控，蓝色箭头代表间接调控。PSY：八氢番茄红素合酶；PDS：八氢番茄红素脱氢酶；Z-ISO：ζ-胡萝卜素异构酶；ZDS：ζ-胡萝卜素脱氢酶；CRTISO：类胡萝卜素异构酶；LCYE：番茄红素ε-环化酶；LCYB：番茄红素β-环化酶；CYP97A/C：细胞色素P450羟化酶；BCH：β-胡萝卜素羟化酶；CCS：辣椒红素/辣椒玉红素合酶；VDE：紫黄质脱环氧化酶；ZEP：玉米黄质环氧化酶；NXS：新黄质合酶；CCD：类胡萝卜素裂解双加氧酶；NCED：9-顺式环氧类胡萝卜素双加氧酶。蓝色为前体MEP途径，橘红色为上游主干路径，绿色为β，ε‑支路，黄色为β，β‑支路，右侧为β，β‑分支衍生物合成通路

Fig. 1 Carotenoid biosynthesis pathwaysCarotenoids are synthesized in plastids, where GGPPS catalyzes the formation of the precursor GGPP, which is subsequently converted to lycopene via PSY-mediated condensation and multi-step desaturation and isomerization reactions. The pathway then diverges into two branches: in the β,β-branch, LCYB catalyzes the cyclization of lycopene to β-carotene, which is further hydroxylated and epoxidized to yield zeaxanthin, violaxanthin, neoxanthin, and abscisic acid; in the β,ε-branch, LCYE and LCYB act cooperatively to produce α-carotene, which is ultimately converted to lutein. Arrows indicate the direction of enzymatic metabolic reactions; black arrows represent direct regulation, and blue arrows represent indirect regulation. PSY: phytoene synthase; PDS: phytoene desaturase; Z-ISO: ζ-carotene isomerase; ZDS: ζ-carotene desaturase; CRTISO: carotenoid isomerase; LCYE: lycopene ε-cyclase; LCYB: lycopene β-cyclase; CYP97A/C: cytochrome P450 hydroxylase; BCH: β-carotene hydroxylase; CCS: capsanthin/capsorubin synthase; VDE: violaxanthin de-epoxidase; ZEP: zeaxanthin epoxidase; NXS: neoxanthin synthase; CCD: carotenoid cleavage dioxygenase; NCED: 9-cis-epoxycarotenoid dioxygenase. Blue represents the MEP pathway for carotenoid precursor synthesis; orange indicates the upstream core backbone pathway; green indicates the β,ε-branch; yellow indicates the β,β-branch; the right side shows derivative synthesis from the β,β-branch

表1 影响类胡萝卜素生物合成的关键转录因子

Table 1 Key transcription factors affecting carotenoid biosynthesis

转录因子 Transcription factors	物种 Species	作用机制 Regulatory mechanism	对色泽的作用 Impact on color		参考文献 Reference
转录因子 Transcription factors	物种 Species	作用机制 Regulatory mechanism	正向 Positive	负向 Negative	参考文献 Reference
MYB	桂花、辣椒、猴面花、鹅掌楸、柑橘、蒺藜苜蓿	直接结合启动子激活结构基因；少数成员抑制结构基因或招募抑制复合物	促进黄、橙、红色素积累	改变色素组成，导致留绿或褪色	［43-46， 84-85］
NAC	番茄、木瓜、柑橘、辣椒	部分成员直接激活合成基因，部分成员抑制限速步骤，部分成员调控叶绿素降解间接促进着色	促进果实转红或黄色积累	抑制番茄红素合成，导致黄/橙色果	［47-55］
bHLH	番茄、木瓜、柑橘、芒果	响应光照调控代谢流分支，部分成员直接激活/抑制结构基因，部分成员通过负反馈精确控制着色	促进类胡萝卜素合成与着色	淡绿色未成熟果实和黄化成熟果实	［56-58， 61-63， 87］
AP2/ERF	番茄、柑橘、津巴多橘	受乙烯诱导，广泛激活类胡萝卜素合成途径上游前体到下游修饰环节多个关键基因	上调柑橘类胡萝卜素，增强果皮橙红色泽与着色	下调番茄总类胡萝卜素和番茄红素含量，果色偏橙	［64-67］
WRKY	番茄、柑橘	激活MEP途径；激活合成基因；协同调控叶绿素降解	上调类胡萝卜素积累，驱动果实由绿色转为黄色或红色	—	［68-70］
BBX	辣椒、番茄、芒果	光形态建成调控因子，直接激活合成基因；部分成员协同调控花青素与类胡萝卜素通路	上调类胡萝卜素/花青素，促进辣椒、番茄、芒果着色	—	［74-76］
MADS	辣椒、柑橘、榴莲	直接激活合成结构基因启动子；部分成员同时激活叶绿素降解基因，协同调控色素代谢	促进类胡萝卜素积累，调控辣椒、柑橘、榴莲果实着色	—	［77-80］

表1 影响类胡萝卜素生物合成的关键转录因子

Table 1 Key transcription factors affecting carotenoid biosynthesis

转录因子 Transcription factors	物种 Species	作用机制 Regulatory mechanism	对色泽的作用 Impact on color		参考文献 Reference
转录因子 Transcription factors	物种 Species	作用机制 Regulatory mechanism	正向 Positive	负向 Negative	参考文献 Reference
MYB	桂花、辣椒、猴面花、鹅掌楸、柑橘、蒺藜苜蓿	直接结合启动子激活结构基因；少数成员抑制结构基因或招募抑制复合物	促进黄、橙、红色素积累	改变色素组成，导致留绿或褪色	［43-46， 84-85］
NAC	番茄、木瓜、柑橘、辣椒	部分成员直接激活合成基因，部分成员抑制限速步骤，部分成员调控叶绿素降解间接促进着色	促进果实转红或黄色积累	抑制番茄红素合成，导致黄/橙色果	［47-55］
bHLH	番茄、木瓜、柑橘、芒果	响应光照调控代谢流分支，部分成员直接激活/抑制结构基因，部分成员通过负反馈精确控制着色	促进类胡萝卜素合成与着色	淡绿色未成熟果实和黄化成熟果实	［56-58， 61-63， 87］
AP2/ERF	番茄、柑橘、津巴多橘	受乙烯诱导，广泛激活类胡萝卜素合成途径上游前体到下游修饰环节多个关键基因	上调柑橘类胡萝卜素，增强果皮橙红色泽与着色	下调番茄总类胡萝卜素和番茄红素含量，果色偏橙	［64-67］
WRKY	番茄、柑橘	激活MEP途径；激活合成基因；协同调控叶绿素降解	上调类胡萝卜素积累，驱动果实由绿色转为黄色或红色	—	［68-70］
BBX	辣椒、番茄、芒果	光形态建成调控因子，直接激活合成基因；部分成员协同调控花青素与类胡萝卜素通路	上调类胡萝卜素/花青素，促进辣椒、番茄、芒果着色	—	［74-76］
MADS	辣椒、柑橘、榴莲	直接激活合成结构基因启动子；部分成员同时激活叶绿素降解基因，协同调控色素代谢	促进类胡萝卜素积累，调控辣椒、柑橘、榴莲果实着色	—	［77-80］

图2 植物花果色泽形成中的类胡萝卜素代谢多层次调控机制类胡萝卜素代谢受转录、转录后、翻译后及表观遗传水平的协同调控。在转录水平，MYB、NAC、bHLH、WRKY、AP2/ERF、MADS等转录因子发挥关键作用；其中，MYB、BBX和bHLH可协同调控类胡萝卜素与类黄酮代谢。PSY作为关键限速酶，LCYE/LCYB作为代谢分支点，均受到多层次调控。在转录后水平，ORANGE蛋白对PSY的稳定作用与PPSR1介导的降解作用形成互补的双向调控网络。在表观遗传层面，类胡萝卜素代谢受DNA甲基化与组蛋白乙酰化的动态变化的共同调控。橙色箭头和字体表示正向调控/促进作用；蓝色箭头和字体表示负向调控/抑制作用。浅绿色模块为转录因子家族；深粉色模块为转录后/翻译后调控；黄色模块为表观遗传修饰；蓝色模块为类黄酮与类胡萝卜素合成协同调控

Fig. 2 Multi-level regulatory mechanism of carotenoid metabolism on color formation in plant flowers and fruitsCarotenoid metabolism is coordinately regulated at the transcriptional, post-transcriptional, post-translational, and epigenetic levels. At the transcriptional level, transcription factors such as MYB, NAC, bHLH, WRKY, AP2/ERF, and MADS play key roles; among them, MYB, BBX, and bHLH cooperatively regulate the biosynthesis of both carotenoids and flavonoids. PSY, as the key rate-limiting enzyme, and LCYE/LCYB, as metabolic branch points, are subject to multi-level regulation. At the post-transcriptional level, the stabilizing effect of ORANGE protein on PSY and the degradation mediated by PPSR1 form a complementary bidirectional regulatory network. At the epigenetic level, dynamic changes in DNA methylation and histone acetylation co-regulate the expression of key genes involved in carotenoid metabolism. Orange arrows and fonts indicate positive regulation/promotion; blue arrows and fonts indicate negative regulation/inhibition. Light green module is the transcription factor family; the dark pink module is post-transcriptional/post-translational regulation; the yellow module is epigenetic modification; the blue module is the synergistic regulation of flavonoid and carotenoid synthesis

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