苹果果实糖分形成的分子基础

doi:10.13560/j.cnki.biotech.bull.1985.2026-0069

摘要/Abstract

摘要：

苹果作为全球广泛栽培的重要水果，果实可溶性糖含量是决定其风味品质与市场竞争力的核心指标，其积累过程涉及复杂的遗传分子调控网络。本文系统综述苹果果实糖分积累的生理基础、分子机制及调控因素。生理层面，糖分积累依赖光合产物（主要为山梨醇和蔗糖）的合成、运输、卸载、代谢与液泡储存等多环节协同作用，且受蔗糖转运蛋白（SUT）、液泡糖转运蛋白（TST）、SWEET转运蛋白等关键载体介导。分子机制上，山梨醇代谢相关基因（MdS6PDH、MdSDH）、蔗糖代谢相关基因（MdSuSy、MdSPS）、己糖代谢相关基因（MdHXK、MdFRK）分别调控不同糖组分的合成与转化；MYB、bZIP、ERF、NAC、WRKY等转录因子通过形成复杂调控网络，精准调控糖代谢与转运相关基因的表达。此外，生长素、脱落酸、乙烯等植物激素通过与糖信号交叉互作，共同参与糖分积累调控；光照、温度、水分、营养元素等环境因素则通过特异性分子通路影响相关基因表达与酶活性，进而塑造果实糖分特征。目前，苹果糖分积累的核心调控因子互作、糖组分转化机制、环境与内源基因互作等问题仍需深入探索。本综述为解析苹果果实糖分积累的分子机制提供全面参考，对推动苹果品质改良的分子育种进程具有重要意义。

关键词: 苹果, 果实品质, 糖分积累, 分子调控, 转运蛋白, 转录因子

Abstract:

Apple is an important fruit widely cultivated worldwide. The soluble sugar content of the fruit is a core index determining its flavor quality and market competitiveness, and its accumulating process involves a complex genetic and molecular regulatory network. This paper systematically reviews the physiological basis, molecular mechanisms, and regulatory factors of sugar accumulation in apple fruits. At the physiological level, sugar accumulation relies on the coordinated action of multiple processes, including the synthesis, transport, unloading, metabolism, and vacuolar storage of photosynthates (mainly sorbitol and sucrose), which are mediated by key carriers such as sucrose transporters (SUTs), tonoplast sugar transporters (TSTs), and SWEET transporters. In terms of molecular mechanisms, sorbitol metabolism-related genes (MdS6PDH and MdSDH), sucrose metabolism-related genes (MdSuSy and MdSPS), and hexose metabolism-related genes (MdHXK and MdFRK) regulate the synthesis and conversion of different sugar components respectively; transcription factors such as MYB, bZIP, ERF, NAC, and WRKY accurately regulate the expression of sugar metabolism and transport-related genes by forming a complex regulatory network. In addition, plant hormones including auxin, abscisic acid, and ethylene participate in the regulation of sugar accumulation through cross-talk with sugar signals; environmental factors such as light, temperature, water, and nutrients affect the expression of related genes and enzyme activities through specific molecular pathways, thereby shaping the sugar characteristics of fruits. At present, issues such as the interaction of core regulatory factors, the mechanism of sugar component conversion, and the interaction between environmental factors and endogenous genes in apple sugar accumulation still need further exploration. This review provides a comprehensive reference for elucidating the genetic and molecular mechanisms of sugar accumulation in apple fruits, and is of great significance for promoting the molecular breeding process of apple quality improvement.

Key words: apple, fruit quality, sugar accumulation, molecular regulation, transporter protein, transcription factor

罗龙鑫, 李智, 李彤, 冯资权, 翟欣悦, 梁成林, 张亚丽, 吴上, 李媛媛, 姜翰. 苹果果实糖分形成的分子基础[J]. 生物技术通报, 2026, 42(3): 156-171.

LUO Long-xin, LI Zhi, LI Tong, FENG Zi-quan, ZHAI Xin-yue, LIANG Cheng-lin, ZHANG Ya-li, WU Shang, LI Yuan-yuan, JIANG Han. Molecular Basis of Sugar Accumulation in Apple Fruits[J]. Biotechnology Bulletin, 2026, 42(3): 156-171.

图/表 3

图1 苹果果实糖分积累过程糖分从叶片光合产物合成、经韧皮部长距离运输与果实韧皮部装载，再到果肉细胞内转运、糖分转化与形成，最终在液泡中储存

Fig. 1 Sugar accumulation process in apple fruitSugars are synthesized as photosynthate in leaves, transported over long distances via the phloem and loaded into the fruit phloem, then undergo intracellular transport, conversion and formation within the flesh cells, and are finally stored in the vacuoles

图2 苹果果实糖分形成分子机制核心转录因子家族（MYB、bZIP、ERF、NAC、WRKY）通过调控糖转运相关蛋白家族（SUT、TST、SWEET），分别介导韧皮部运输、液泡糖富集及跨细胞/胞内糖转运过程，最终调控果糖、葡萄糖、蔗糖、山梨醇等糖分在果实中的积累

Fig. 2 Molecular mechanism of sugar formation in apple fruitCore transcription factor families (MYB, bZIP, ERF, NAC, and WRKY) orchestrate the expression of sugar transport-related protein families (SUT, TST, and SWEET). These transporters mediate key processes including phloem transport, vacuolar sugar sequestration, and intercellular/intracellular sugar partitioning, ultimately governing the accumulation of fructose, glucose, sucrose, and sorbitol in the fruit

图3 环境因素与植物激素通过糖信号通路协同调控苹果果实发育的分子网络环境因素（光照、低温、营养元素、适度干旱等）通过激活特定信号分子（如MdCBF1/2、MdCIPK22、MdDREB2A）调控糖信号通路；植物激素（CTK/GA、生长素、乙烯、ABA等）则通过调控光合面积、库强、糖代谢及干旱响应，与糖信号通路形成交叉响应与信号协同，共同影响果实发育过程

Fig. 3 Molecular network of environmental factors and plant hormones synergistically regulating apple fruit development via the sugar signaling pathwayEnvironmental factors (light, low temperature, nutrients, moderate drought, etc.) regulate the sugar signaling pathway by activating specific signaling molecules (e.g., MdCBF1/2, MdCIPK22, MdDREB2A). Plant hormones (CTK/GA, auxin, ethylene, ABA, etc.) form cross-response and signal synergy with the sugar signaling pathway by regulating photosynthetic area, sink strength, sugar metabolism, and drought response, thereby collectively influencing fruit development

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