果实花色苷酶促降解机制研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2026-0164

摘要/Abstract

摘要：

花色苷是一类广泛存在于植物体内的天然水溶性色素，作为决定果实外观色泽与营养品质的关键组分，其积累水平直接影响果实商品经济价值。然而，部分研究表明花色苷过量积累会影响某些果实香气和口感，因此维持其积累动态平衡可为果实品质改良提供新思路。花色苷较不稳定，易受光照、温度等外界环境因子的影响而降解。花色苷降解对维持其代谢稳态起着关键作用，酶促降解是介导植物体内花色苷降解的核心途径。本文系统综述了花色苷降解相关酶类及其作用机制的研究进展，重点概述了β-葡萄糖苷酶、过氧化物酶及多酚氧化酶3类关键酶介导的花色苷降解模型。β-葡萄糖苷酶可特异性水解花色苷糖苷键，释放稳定性较低的花色素苷元并自发裂解，促进其降解；花色素苷元亦可被过氧化物酶进一步氧化降解。过氧化物酶与多酚氧化酶/漆酶可将酚类底物催化为强氧化活性的醌类中间体，并与花色苷发生耦合氧化反应介导其降解。此外，多酚氧化酶/漆酶在体外条件下可直接降解花色苷。上述酶类在液泡中协同介导花色苷降解，参与果实色泽品质形成，但其调控的分子机制仍有待深入解析。本文围绕降解酶催化机制、多酶协同模式、合成与降解互作关系及环境响应机制等关键科学问题与研究挑战展开论述，旨在为精准调控花色苷积累的动态平衡、多维度改良果实品质提供理论支撑。

关键词: 果实, 花色苷, 酶促降解, β-葡萄糖苷酶, 过氧化物酶, 多酚氧化酶, 漆酶, 分子机制

Abstract:

Anthocyanins are a class of natural water-soluble pigments widely distributed in plants. As major determinants of fruit coloration and nutritional quality, the accumulation directly influences fruit marketability and economic value. Studies have shown that excessive anthocyanin accumulation can adversely affect fruit aroma and flavor to some extent, underscoring the importance of maintaining a dynamic balance in anthocyanin levels for optimal fruit quality. Anthocyanins are inherently unstable and susceptible to degradation under environmental factors such as light and temperature. Anthocyanin degradation is therefore essential for maintaining metabolic homeostasis, with enzymatic degradation constituting the primary pathway in plants. This review systematically summarizes current advances in enzymes involved in anthocyanin degradation and their degradation mechanisms, with particular emphasis on degradation models mediated by three major enzyme classes such as β-glucosidase, peroxidase, and polyphenol oxidase. β-glucosidases specifically hydrolyze the glycosidic bonds of anthocyanins, releasing unstable anthocyanidins undergo subsequently spontaneous degradation and can be further oxidized by peroxidases. Peroxidases and polyphenol oxidases/laccases catalyze phenolic substrates into highly reactive quinone intermediates, which mediate anthocyanin degradation through coupled oxidation reactions. In addition, polyphenol oxidases/laccases can directly degrade anthocyanins in vitro. Collectively, these enzymes synergistically function within the vacuole to modulate anthocyanin degradation and contribute to fruit coloration and quality formation, although the regulatory mechanisms remain further analysis. This review further discusses key scientific questions and challenges in the field, including the catalytic mechanisms of degradation enzymes, multi-enzyme coordination patterns, the interplay between biosynthesis and degradation and environmental response mechanisms, aiming to provide a theoretical framework for the precise regulation of anthocyanin homeostasis and the multidimensional improvement of fruit quality.

Key words: fruit, anthocyanin, enzymatic degradation, β-glucosidase, peroxidase, polyphenol oxidase, laccase, molecular mechanism

彭楚, 孙娟利, 郑蓓蓓, 张若西, 韩月彭, 赵云. 果实花色苷酶促降解机制研究进展[J]. 生物技术通报, doi: 10.13560/j.cnki.biotech.bull.1985.2026-0164.

PENG Chu, SUN Juan-li, ZHENG Bei-bei, ZHANG Ruo-xi, HAN Yue-peng, ZHAO Yun. Research Progress in the Enzymatic Degradation Mechanism of Anthocyanins in Fruits[J]. Biotechnology Bulletin, doi: 10.13560/j.cnki.biotech.bull.1985.2026-0164.

图/表 3

图1 组织损伤条件下细胞质中花色苷降解途径示意图A：植物受各类胁迫作用时，其细胞膜结构完整性受损，导致膜体系解体，质体中的PPO/CO被释放至细胞质；B：释放的PPO/CO可氧化游离酚类底物生成醌类物质，后者与花色苷形成耦合物，该耦合物经水解、聚合反应导致组织褪色；此外，该过程中生成的醌类物质可被快速还原为酚类底物，形成循环反应，高效降解细胞质中的花色苷；PPO/CO：多酚氧化酶家族中儿茶酚氧化酶类型

Fig. 1 Schematic representation of cytoplasmic anthocyanin degradation triggered by tissue injuryA: Under various stress conditions, the disruption of cellular membrane integrity leads to the breakdown of membrane systems, resulting in the release of plastid-localized PPO/CO into the cytoplasm. B: The released PPO/CO catalyzes the oxidation of free phenolic substrates to generate reactive quinone intermediates, which form conjugates with anthocyanins. These conjugates subsequently undergo hydrolysis and polymerization, ultimately causing tissue discoloration. Moreover, the quinone intermediates can be rapidly reduced back to phenolic substrates, establishing a redox cycle that facilitates efficient anthocyanin degradation in the cytoplasm. PPO/CO: Catechol oxidase belonging to the polyphenol oxidase family

图2 正常生理条件下液泡内花色苷降解途径示意图正常生理条件下液泡内花色苷酶促降解途径。（1）花色苷经BGLU水解生成花色素苷元，后者可一定程度上发生自发裂解；（2）BGLU水解产生的花色素苷元被PRX氧化为醌类物质，再与花色苷耦合形成醌类共轭物，经水解或聚合导致花色苷降解；（3）（4）液泡中游离酚类物质分别被PRX和PPO/LAC氧化为醌类，进而通过与途径（2）相似的耦合反应介导花色苷降解；（5）花色苷可被PPO/LAC直接降解。BGLU：β-葡萄糖苷酶；PRX：过氧化物酶；PPO/LAC：多酚氧化酶家族中漆酶类型

Fig. 2 Proposed schematic model illustrating enzymatic degradation pathways of anthocyanins in the vacuole under normal physiological conditionsUnder normal physiological conditions, anthocyanins sequestered in the vacuole undergo enzymatic degradation through multiple degradation routes. 1) Anthocyanins are hydrolyzed by BGLU to generate anthocyanidins, which are inherently unstable and prone to spontaneous decomposition. 2) Anthocyanidins generated by BGLU are further oxidized to quinones by PRX, which then couple with anthocyanins to form quinone conjugates, leading to anthocyanin degradation through hydrolysis or polymerization. 3) 4) Free phenol compounds present in the vacuole are oxidized to quinones by PRX and PPO/LAC, which then mediate anthocyanin degradation by a coupling reaction similar to the pathway 2). 5) Anthocyanins are directly oxidatively degraded by PPO/LAC. BGLU: β-glucosidase; PRX: Peroxidase; PPO/LAC: the laccase belonging to the polyphenol oxidase family

图3 环境因子调控果实花色苷代谢及育种应用示意图光照、温度作为影响果实花色苷积累的关键环境因子，可通过协同调控花色苷生物合成通路与降解途径中关键基因的表达水平，精准调控果实花色苷积累量。其中，DFR、ANS和UFGT等花色苷生物合成关键结构基因，其表达量上调可显著促进花色苷积累；而BGLU、PRX和PPO等花色苷降解酶相关基因，其表达量上调则会加速花色苷分解代谢进程。花色苷合成与降解代谢过程在转录调控层面协同响应光照、温度等环境信号的变化，二者之间的动态平衡共同维持果实花色苷代谢稳态。解析环境因子协同调控花色苷合成与降解的分子机制，不仅可为研发以色泽改良为核心的品质调控技术提供理论依据，也可为果实品质性状的遗传改良提供新思路。DFR：二氢黄酮醇4-还原酶；ANS：花色素合成酶；UFGT：UDP葡萄糖-类黄酮3-O-葡糖基转移酶；BGLU：β-葡萄糖苷酶；PRX：过氧化物酶；PPO：多酚氧化酶

Fig. 3 Schematic diagram illustrating environmental regulation of anthocyanin metabolism in fruit and its applications in breeding programsEnvironmental factors such as light and temperature regulate anthocyanin accumulation in fruit by coordinately modulating the expressions of key genes involved in both biosynthesis and degradation. Enhanced expressions of key biosynthetic genes, including DFR, ANS and UFGT, promotes anthocyanin accumulation, whereas the upregulation of degradation-associated genes such as BGLU, PRX and PPO accelerates anthocyanin turnover. Anthocyanin biosynthesis and degradation respond coordinately to changes in environmental cues at the transcriptional level, and their coordinated interplay maintains anthocyanin metabolic homeostasis in fruit. Deciphering the molecular mechanism by which environmental factors coordinately regulate anthocyanin biosynthesis and degradation will provide a foundation for coloration quality improvement strategies and genetic enhancement of fruit quality traits. DFR: Dihydroflavonol 4-reductase; ANS: Anthocyanidin synthase; UFGT: UDP-glucose: flavonoid 3-O-glucosyltransferase; BGLU: β-glucosidase; PRX: peroxidase; PPO: polyphenol oxidase

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