Research Progress on the Synergistic Regulation of Sugar and Acid in Fresh Fruits

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

Acta Horticulturae Sinica ›› 2026, Vol. 53 ›› Issue (7): 1891-1909.doi: 10.16420/j.issn.0513-353x.2026-0447

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Research Progress on the Synergistic Regulation of Sugar and Acid in Fresh Fruits

JIAO Tiantian, DANG Suqing, ZHU Lingcheng, LI Mingjun^*()

State Key Laboratory of Crop Stress Resistance and High-Efficiency Production， Shaanxi Key Laboratory of Apple，College of Horticulture，Northwest A & F University，Yangling， Shaanxi 712100， China

Received:2026-06-04 Revised:2026-07-01 Online:2026-07-25 Published:2026-07-23
Contact: LI Mingjun

Abstract

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

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.

Figures/Tables 3

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

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

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）

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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Research Progress on the Synergistic Regulation of Sugar and Acid in Fresh Fruits

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