果实糖酸协同调控的研究进展

doi:10.16420/j.issn.0513-353x.2026-0447

摘要/Abstract

摘要：

糖酸协同调控机制是果实品质研究的热点。目前，果实糖酸协同调控涵盖了从代谢酶活性、跨膜转运到激素信号及表观遗传修饰等多个层次，并通过复杂的调控网络共同决定果实的糖酸平衡。本文中重点综述了果实中糖和有机酸在代谢及转运层面的偶联机制，深入阐述了近年来有关果实中糖酸协同调控的研究进展，重点讨论糖酸代谢、植物激素、转录因子及表观遗传修饰对果实糖酸协同调控的影响机制，旨在为深入解析水果糖酸协同调控机制及改良果实品质提供理论依据。

关键词: 果实, 糖, 有机酸, 协同调控

Abstract:

The synergistic regulation of sugar and acid is a popular themes in fruit quality. At present，the synergistic regulation of sugar and acid in fruit encompasses multiple levels，ranging from metabolic enzyme activity and transmembrane transport to hormonal signal transduction and epigenetic modification，collectively shaping fruit sugar-acid balance through a complex regulatory network. This review focuses on the coupling mechanisms between sugars and organic acids metabolism and transport in fruit，and provides an in-depth summary of recent advances in the synergistic regulation of fruit sugar and acid，with particular emphasis on the regulatory mechanisms of metabolic enzymes，phytohormones，transcription factors，and epigenetic modifications underlying this process. The aim is to provide a theoretical basis for further elucidation of the synergistic regulatory mechanism of sugar and acid in fresh fruits and improvement of fruit quality.

Key words: fruit, sugar, organic acids, synergistic regulation

焦田甜, 党素青, 祝令成, 李明军. 果实糖酸协同调控的研究进展[J]. 园艺学报, 2026, 53(7): 1891-1909.

JIAO Tiantian, DANG Suqing, ZHU Lingcheng, LI Mingjun. Research Progress on the Synergistic Regulation of Sugar and Acid in Fresh Fruits[J]. Acta Horticulturae Sinica, 2026, 53(7): 1891-1909.

图/表 3

表1 不同果实糖酸组成及发育过程中糖酸变化特征

Table 1 Sugar and organic acid composition and their dynamic changes during fruit development in different fruit species

水果 Fruit	糖分主导类型 Predominant sugar type	糖积累模式 Sugar accumulation pattern	有机酸主导类型 Predominant organic acid type	酸变化规律 Organic acid dynamics	参考文献 Reference
苹果 Apple	果糖积累型 Fructose-dominant	淀粉—糖中间积累型 “Starch-sugar” intermediate accumulation type	苹果酸型 Malate-dominant	先升后降 Increases initially and then decreases	Li et al.,2012；Ma et al.,2018
梨 Pear	果糖积累型 Fructose-dominant	淀粉—糖中间积累型 “Starch-sugar” intermediate accumulation type	苹果酸型 Malate-dominant	先升后降 Increases initially and then decreases	李甲明等， 2013；Wu et al.,2025
草莓 Strawberry	果糖积累型 Fructose-dominant	直接积累型 Direct sugar accumulation type	柠檬酸型 Citrate-dominant	先升后降 Increases initially and then decreases	郑丽静等， 2015；董书琦等,2023
葡萄 Grape	己糖积累型（葡萄糖+果糖） Hexose-dominant (glucose + fructose)	直接积累型 Direct sugar accumulation type	酒石酸型 Tartrate-dominant	先升后降 Increases initially and then decreases	Dai et al.,2009；Coelho et al.,2018
猕猴桃 Kiwifruit	果糖积累型 Fructose-dominant	淀粉转化型 Starch invert	奎宁酸型 Quinate-dominant	整体下降（奎宁酸降幅最大） Overall decline（greatest decrease during fruit development）	Wang et al.,2022
香蕉 Banana	己糖积累型（葡萄糖＋果糖） Hexose-dominant (glucose + fructose)	淀粉转化型 Starch invert	苹果酸型 Malate-dominant	先升后降 Increases initially and then decreases	Do Nascimento et al.,2006；Famiani et al.,2015
桃 Peach	蔗糖积累型 Sucrose-dominant	淀粉—糖中间积累型 “Starch-sugar” Intermediate accumulation type	苹果酸型 Malate-dominant	先升后降 Increases initially and then decreases	Wu et al.,2005；Zheng et al.,2021；李俊豪等,2025
杏 Apricot	蔗糖积累型 Sucrose-dominant	直接积累型 Direct sugar accumulation type	苹果酸/柠檬酸型 Malate/Citrate-dominant	先升后降 Increases initially and then decreases	Xi et al.,2016
柑橘 Citrus	蔗糖积累型 Sucrose-dominant	直接积累型 Direct sugar accumulation type	柠檬酸型 Citrate-dominant	先升后降 Increases initially and then decreases	Song et al.,2025
甜樱桃 Sweet cherry	葡萄糖积累型 Glucose-dominant	直接积累型 Direct sugar accumulation type	苹果酸型 Malate-dominant	持续积累（成熟后期部分品种略有下降） Continuous accumulation（a slight decline occurs in some cultivars at the late ripening stage）	李玉生等,2025

表1 不同果实糖酸组成及发育过程中糖酸变化特征

Table 1 Sugar and organic acid composition and their dynamic changes during fruit development in different fruit species

水果 Fruit	糖分主导类型 Predominant sugar type	糖积累模式 Sugar accumulation pattern	有机酸主导类型 Predominant organic acid type	酸变化规律 Organic acid dynamics	参考文献 Reference
苹果 Apple	果糖积累型 Fructose-dominant	淀粉—糖中间积累型 “Starch-sugar” intermediate accumulation type	苹果酸型 Malate-dominant	先升后降 Increases initially and then decreases	Li et al.,2012；Ma et al.,2018
梨 Pear	果糖积累型 Fructose-dominant	淀粉—糖中间积累型 “Starch-sugar” intermediate accumulation type	苹果酸型 Malate-dominant	先升后降 Increases initially and then decreases	李甲明等， 2013；Wu et al.,2025
草莓 Strawberry	果糖积累型 Fructose-dominant	直接积累型 Direct sugar accumulation type	柠檬酸型 Citrate-dominant	先升后降 Increases initially and then decreases	郑丽静等， 2015；董书琦等,2023
葡萄 Grape	己糖积累型（葡萄糖+果糖） Hexose-dominant (glucose + fructose)	直接积累型 Direct sugar accumulation type	酒石酸型 Tartrate-dominant	先升后降 Increases initially and then decreases	Dai et al.,2009；Coelho et al.,2018
猕猴桃 Kiwifruit	果糖积累型 Fructose-dominant	淀粉转化型 Starch invert	奎宁酸型 Quinate-dominant	整体下降（奎宁酸降幅最大） Overall decline（greatest decrease during fruit development）	Wang et al.,2022
香蕉 Banana	己糖积累型（葡萄糖＋果糖） Hexose-dominant (glucose + fructose)	淀粉转化型 Starch invert	苹果酸型 Malate-dominant	先升后降 Increases initially and then decreases	Do Nascimento et al.,2006；Famiani et al.,2015
桃 Peach	蔗糖积累型 Sucrose-dominant	淀粉—糖中间积累型 “Starch-sugar” Intermediate accumulation type	苹果酸型 Malate-dominant	先升后降 Increases initially and then decreases	Wu et al.,2005；Zheng et al.,2021；李俊豪等,2025
杏 Apricot	蔗糖积累型 Sucrose-dominant	直接积累型 Direct sugar accumulation type	苹果酸/柠檬酸型 Malate/Citrate-dominant	先升后降 Increases initially and then decreases	Xi et al.,2016
柑橘 Citrus	蔗糖积累型 Sucrose-dominant	直接积累型 Direct sugar accumulation type	柠檬酸型 Citrate-dominant	先升后降 Increases initially and then decreases	Song et al.,2025
甜樱桃 Sweet cherry	葡萄糖积累型 Glucose-dominant	直接积累型 Direct sugar accumulation type	苹果酸型 Malate-dominant	持续积累（成熟后期部分品种略有下降） Continuous accumulation（a slight decline occurs in some cultivars at the late ripening stage）	李玉生等,2025

图1 果实糖酸代谢及转运关系图 HK：己糖激酶；PFK：磷酸果糖激酶；PK：丙酮酸激酶；G6Pase：葡萄糖-6-磷酸酶：FBPase：果糖-1,6-二磷酸酶；PEPCK：磷酸烯醇式丙酮酸；MDH：苹果酸脱氢酶。紫色图标代表糖/H+反向转运蛋白；橙色图标代表糖/H+同向转运蛋白；灰色图标代表苹果酸/柠檬酸转运蛋白；蓝色图标代表质子泵。V-ATPase：液泡H+-腺苷三磷酸酶；V-PPase：液泡H+-焦磷酸酶；P-ATPase：液泡P型腺苷三磷酸酶；VGT：液泡葡萄糖转运蛋白；TMT：液泡膜单糖转运蛋白；TST：液泡膜糖转运蛋白；SUT：蔗糖转运蛋白；ERDL6：早期响应干旱类似蛋白；INT：肌醇转运蛋白；Cit1：柠檬酸转运蛋白1；tDT：液泡膜二羧酸转运蛋白；ALMT：铝激活型苹果酸转运通道；glc：葡萄糖；fru：果糖；suc：蔗糖；Mal2-：苹果酸二价阴离子；Cit3-：柠檬酸三价阴离子；HCit2-/（H2Cit-）：柠檬酸二价阴离子/（柠檬酸单价阴离子）

Fig. 1 Schematic diagram of sugar and organic acid metabolism and transport in fruit HK：Hexokinase；PFK：Phosphofructokinase；PK：Pyruvate kinase；G6Pase：Glucose-6-phosphatase；FBPase：Fructose-1,6-bisphosphatase；PEPCK：Phosphoenolpyruvate carboxykinase；MDH：Malate dehydrogenase. Purple icons represent sugar/H+ antiporters；Orange icons represent sugar/H+ symporters；Gray icons represent malate/citrate transporters；Blue icons represent proton pumps. V-ATPase：Vacuolar H+-adenosine triphosphatase；V-PPase：Vacuolar H+-Pyrophosphatase；P-ATPase：P-type adenosine triphosphatase；VGT：Vacuolar glucose transporter；TMT：Tonoplast monosaccharide transporter；TST：Tonoplast sugar transporter；SUT：Sucrose transporter；ERDL6：Early response to dehydration 6-like transporter；INT：Inositol transporter；Cit1：Citrate transporter 1；tDT：Tonoplast dicarboxylate transporter；ALMT：Aluminum-activated malate transporter；glc：Glucose；fru：Fructose；suc：Sucrose；Mal2-：Dianion malate；Cit3-：Trianion citrate；HCit2-/（H2Cit-）：Dianion citrate/（Monoanion citrate）

图2 果实糖酸协同调控示意图植物激素（ABA、乙烯、油菜素内酯）和表观遗传修饰（m6A修饰、DNA甲基化、组蛋白甲基化）作为上游调控信号，协同影响果实糖酸代谢。激素信号直接作用于转录因子层（NAC、MYB、bZIP、bHLH、CCA1、SAR），进而调控下游糖代谢相关基因（NI、SPS、SUS、SUT、TST）和有机酸代谢相关基因（ALMT、ACO、GAD、PH8）的表达（实线箭头）；表观遗传修饰目前主要通过调控功能基因的转录活性参与糖酸调控，其对转录因子的直接调控关系尚待进一步明确（虚线箭头）。相同颜色显示转录因子与激素的诱导关系：ABA可诱导bZIP和CCA1的表达，乙烯（ETH）可诱导MYB的表达；灰色转录因子（NAC、bHLH、SAR）在本综述范围内尚未发现受特定激素直接诱导的明确证据。MDH、MPC和NADP-ME（椭圆区域）为糖酸协同调控的关键酶基因，同时参与糖与有机酸的代谢调控。多层次调控因子的协同作用最终决定果实糖酸平衡与风味品质

Fig. 2 Schematic diagram of synergistic regulation of sugar and acid in fruit Phytohormones（ABA，ETH，and BR）and epigenetic modifications（m6A modification，DNA methylation，and histone methylation）serve as upstream regulatory signals that cooperatively influence sugar and acid metabolism in fruit. Phytohormones directly act on transcription factors（NAC，MYB，bZIP，bHLH，CCA1，SAR），which subsequently modulate the expression of downstream sugar-related genes（NI，SPS，SUS，SUT，TST）and acid-related genes（ALMT，ACO，GAD，PH8）（solid arrows）；Epigenetic modifications currently appear to regulate fruit sugar-acid metabolism primarily by modulating the transcriptional activity of functional metabolic genes，while their direct regulatory effects on transcription factors remain to be further elucidated（dashed arrow）. The color correspondence between transcription factors and hormones indicates hormone-induction relationships：ABA induces the expression of bZIP and CCA1，while ethylene induces MYB expression；gray-colored transcription factors（NAC，bHLH，SAR）have not yet been found to be directly induced by specific hormones within the scope of this review. MDH，MPC，and NADP-ME （highlighted ellipse）represent key enzyme-encoding genes involved in the synergistic regulation of both sugar and organic acid metabolism. The multilevel cooperative action of these regulatory factors ultimately determines the sugar-acid balance and flavor quality of fruit

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