生物技术通报 ›› 2026, Vol. 42 ›› Issue (3): 79-95.doi: 10.13560/j.cnki.biotech.bull.1985.2025-1025
杜连达(
), 魏萌萌, 陈泽, 郭伟, 赵婷婷(
), 胡大刚(
)
收稿日期:2025-09-26
出版日期:2026-03-26
发布日期:2026-04-23
通讯作者:
胡大刚,男,博士,教授,研究方向 :苹果果实品质生物学与营养健康;E-mail: fap_296566@163.com作者简介:杜连达,男,博士,研究方向 :苹果果实品质生物学;E-mail: liandadu@126.com
基金资助:
DU Lian-da(
), WEI Meng-meng, CHEN Ze, GUO Wei, ZHAO Ting-ting(
), HU Da-gang(
)
Received:2025-09-26
Published:2026-03-26
Online:2026-04-23
摘要:
糖酸平衡是苹果果实品质评价体系中的关键参数,既综合反映了糖类与有机酸的代谢平衡,又直接决定其风味特征与市场价值。苹果果实糖酸的形成受遗传背景与环境因子双重调控,其代谢机制及调控模式已成为果实品质研究的核心热点之一。目前,光照、温度、水分与土壤养分等环境因子不仅独立影响糖与有机酸的合成、降解及转运过程,更通过复杂的交互网络共同塑造果实代谢格局。本文综述了苹果糖酸代谢与转运的分子调控机制,系统分析了光照、温度、水分与养分供应等环境因子协同调控信号转导途径间接影响碳流分配与酸积累。这些多维度的环境交互作用构成了糖酸代谢调控的核心生态网络,同时探讨了农业管理措施在调节糖酸平衡与提升果实风味中的调控潜力与现存问题。未来研究应结合多组学、单细胞及空间转录组等前沿技术,解析基因型‒环境‒管理互作下的糖酸代谢网络,揭示环境信号整合与表观遗传记忆的分子基础。深入理解这些机制,不仅有助于开展面向环境适应性的苹果育种,还为应对气候变化背景下的果实品质稳定与优质风味形成提供理论支撑与实践方向。
杜连达, 魏萌萌, 陈泽, 郭伟, 赵婷婷, 胡大刚. 环境因子对苹果果实糖酸品质的影响:多因子交互作用与环境适应性育种[J]. 生物技术通报, 2026, 42(3): 79-95.
DU Lian-da, WEI Meng-meng, CHEN Ze, GUO Wei, ZHAO Ting-ting, HU Da-gang. Effects of Environmental Factors on Sugar-acid Quality in Apple Fruits: Multi-factor Interactions and Environmental Adaption Breeding[J]. Biotechnology Bulletin, 2026, 42(3): 79-95.
图1 苹果糖酸代谢途径网络(细胞膜-胞质-线粒体)糖经过膜转运蛋白跨膜转运到细胞质中,其中蔗糖通过蔗糖合成酶以及蔗糖转化酶分解为果糖和葡萄糖等,同时在细胞质中葡萄糖发生糖酵解过程产生丙酮酸和苹果酸,丙酮酸进一步通过线粒体的三羧酸循环产生苹果酸等有机酸,同时磷酸烯醇式丙酮酸通过糖异生作用生成葡萄糖。OAA:草酰乙酸;PEP:磷酸烯醇式丙酮酸;PEPCK:磷酸烯醇式丙酮酸羧激酶;PPDK:丙酮酸磷酸双激酶;NADP-ME:NADP-苹果酸酶;NAD-MDH:NAD-苹果酸脱氢酶;TCA:三羧酸循环
Fig. 1 Metabolic pathway network of sugar-acid in apple fruit (Cell membrane-cytoplasm-mitochondria)Sugars are transported across membranes into the cytoplasm via specific membrane transport proteins. Within the cytoplasm, sucrose is hydrolyzed to fructose and glucose by sucrose synthase and invertase. Subsequently, glucose undergoes glycolysis to produce pyruvate and malate. Pyruvate is further metabolized through the mitochondrial TCA cycle to generate malate and other organic acids. Meanwhile, phosphoenolpyruvate regenerate glucose by gluconeogenesis. OAA: Oxaloacetic acid. PEP: Phosphoenolpyruvate. PEPCK: Phosphoenolpyruvate carboxykinase. PPDK: Pyruvate phosphate dikinase. NADP-ME: NADP-malic enzyme. NAD-MDH: NAD-malate dehydrogenase. TCA: Tricarboxylic acid cycle
图2 苹果液泡内糖酸转运和“酸陷阱”模型苹果液泡作为重要的糖酸交互场所,其液泡膜承载的多种转运蛋白影响糖酸的储存与外排,包括糖分内向转运蛋白、糖分外向转运蛋白、糖分双向转运蛋白、苹果酸/柠檬酸转运蛋白、铝激活型苹果酸转运蛋白和质子泵。糖一般以葡萄糖、果糖以及蔗糖形式,而酸主要为苹果酸和柠檬酸穿梭于液泡与细胞质间。通过质子泵作用,苹果液泡pH值一般维持在5.5左右,使苹果酸和柠檬酸产生梯度解离储存于液泡中。Fru:果糖;Glc:葡萄糖;Suc:蔗糖
Fig. 2 Sugar and acid transport within the vacuole and the ‘acid trap’ model in appleAs a crucial site for sugar-acid interactions, the vacuole in apple cells hosts multiple transport proteins on its tonoplast that regulate the storage and efflux of these compounds. Transporters include sugar influx transporters, sugar efflux transporters, bidirectional movement of sugars, as well as malate/citrate transporters, aluminum-activated malate transporters, and proton pumps. Sugars are mainly transported in the forms of glucose, fructose, and sucrose, while organic acids such as malate and citrate shuttle between the vacuole and the cytosol. Driven by proton pump activity, the vacuolar pH in apples is typically maintained at around 5.5, which facilitates the gradient-dependent dissociation and accumulation of malate and citrate within the vacuole. Fru: Fructose. Glc: Glucose. Suc: Sucrose
发育阶段 Developmental stages | 温度参数 Temperature parameter | 主要品质影响 Primary effects on quality | 潜在机制 Potential mechanism | 参考文献 Reference |
|---|---|---|---|---|
6月下旬‒7月上旬 Late June to early July | 平均最低温 Average minimum temperature | 可溶性固形物 Soluble solids | 影响糖合成代谢 Influence sugar biosynthesis and metabolism | [ |
日均温 Daily mean temperature | ||||
7月中下旬 Mid-to-late July | 日均温 Daily mean temperature | 调节糖酸比 Regulate the sugar-acid ratio | 协调糖转运与有机酸代谢 Coordination of sugar transport and organic acid metabolism | |
整个生长期 Whole growth period | 昼夜温差 Diurnal temperature variation | 提升总糖含量 Increase the total sugar content | 提高净光合效率,减少夜间呼吸消耗 Enhance net photosynthesis efficiency and minimize nocturnal respiratory consumption | [ |
表1 温度对苹果糖酸品质影响的关键阶段及潜在机制
Table 1 Temperature-sensitive stages and potential mechanisms shaping sugar-acid quality in apple
发育阶段 Developmental stages | 温度参数 Temperature parameter | 主要品质影响 Primary effects on quality | 潜在机制 Potential mechanism | 参考文献 Reference |
|---|---|---|---|---|
6月下旬‒7月上旬 Late June to early July | 平均最低温 Average minimum temperature | 可溶性固形物 Soluble solids | 影响糖合成代谢 Influence sugar biosynthesis and metabolism | [ |
日均温 Daily mean temperature | ||||
7月中下旬 Mid-to-late July | 日均温 Daily mean temperature | 调节糖酸比 Regulate the sugar-acid ratio | 协调糖转运与有机酸代谢 Coordination of sugar transport and organic acid metabolism | |
整个生长期 Whole growth period | 昼夜温差 Diurnal temperature variation | 提升总糖含量 Increase the total sugar content | 提高净光合效率,减少夜间呼吸消耗 Enhance net photosynthesis efficiency and minimize nocturnal respiratory consumption | [ |
品种类型 Varietal type | 光照响应特征 Light response characteristics | 糖代谢影响 Influence on sugar metabolism | 酸代谢影响 Influence on acid metabolism | 外观表现 Phenotypic appearance | 参考文献 Reference |
|---|---|---|---|---|---|
非红色品种(Golden delicious) Non-red cultivar (Golden delicious) | 对光强敏感,光质反应弱 High sensitivity to light intensity but a weak response to light quality | 遮荫显著降低糖含量 Shading treatment leads to a significant decrease in sugar content | 遮荫轻微增加酸含量 Shading treatment leads to a slight increase in organic acid content | 微量花色苷积累,着色变化小 The fruit contains very low anthocyanins, with little coloration development | [ |
浅红色品种(Cripps pink) Light-red cultivar (Cripps pink) | 中等光质敏感性 Moderate sensitivity to light quality | 遮荫降低糖含量 Shading treatment leads to a decrease in sugar content | 遮荫增加酸含量 Shading treatment leads to an increase in organic acid content | 花色苷积累受抑制,着色变浅 The inhibition of anthocyanin accumulation leads to a lighter pigmentation | |
深红色品种(Qinguan) Deep-red cultivar (Qinguan) | 对UV和蓝光高度敏感 High sensitivity to UV and blue light | 遮荫降低糖含量 Shading treatment leads to a decrease in sugar content | 遮荫增加酸含量 Shading treatment leads to an increase in organic acid content | 完全抑制花色苷合成,失去红色 Anthocyanin biosynthesis is completely suppressed, leading to the loss of red pigmentation | |
野生苹果种质 Wild apple germplasm | 普遍适应弱光能力较强 Generally strong ability to adapt to low-light conditions | 糖组成以果糖和葡萄糖为主 The sugar composition is predominantly composed of fructose and glucose | 苹果酸含量显著高于栽培种 The malate content is significantly higher compared with that of cultivated apple varieties | 通常着色较深,果实较小 The fruit generally shows intense pigmentation and a smaller size | [ |
表2 比较光照条件对不同苹果品种糖酸品质的影响
Table 2 Effects of varying light conditions on sugar-acid composition across different apple cultivars
品种类型 Varietal type | 光照响应特征 Light response characteristics | 糖代谢影响 Influence on sugar metabolism | 酸代谢影响 Influence on acid metabolism | 外观表现 Phenotypic appearance | 参考文献 Reference |
|---|---|---|---|---|---|
非红色品种(Golden delicious) Non-red cultivar (Golden delicious) | 对光强敏感,光质反应弱 High sensitivity to light intensity but a weak response to light quality | 遮荫显著降低糖含量 Shading treatment leads to a significant decrease in sugar content | 遮荫轻微增加酸含量 Shading treatment leads to a slight increase in organic acid content | 微量花色苷积累,着色变化小 The fruit contains very low anthocyanins, with little coloration development | [ |
浅红色品种(Cripps pink) Light-red cultivar (Cripps pink) | 中等光质敏感性 Moderate sensitivity to light quality | 遮荫降低糖含量 Shading treatment leads to a decrease in sugar content | 遮荫增加酸含量 Shading treatment leads to an increase in organic acid content | 花色苷积累受抑制,着色变浅 The inhibition of anthocyanin accumulation leads to a lighter pigmentation | |
深红色品种(Qinguan) Deep-red cultivar (Qinguan) | 对UV和蓝光高度敏感 High sensitivity to UV and blue light | 遮荫降低糖含量 Shading treatment leads to a decrease in sugar content | 遮荫增加酸含量 Shading treatment leads to an increase in organic acid content | 完全抑制花色苷合成,失去红色 Anthocyanin biosynthesis is completely suppressed, leading to the loss of red pigmentation | |
野生苹果种质 Wild apple germplasm | 普遍适应弱光能力较强 Generally strong ability to adapt to low-light conditions | 糖组成以果糖和葡萄糖为主 The sugar composition is predominantly composed of fructose and glucose | 苹果酸含量显著高于栽培种 The malate content is significantly higher compared with that of cultivated apple varieties | 通常着色较深,果实较小 The fruit generally shows intense pigmentation and a smaller size | [ |
调控途径 Regulatory pathway | 核心措施 Core practices | 对糖酸品质的主要影响 Primary impacts on sugar-acid quality | 参考文献 Reference |
|---|---|---|---|
主动微环境调控 Active microenvironment regulation | 行间生草/树盘覆盖 | 降低土温,减少蒸发,缓解高温胁迫,促进糖分平稳积累 | [ |
| 架设遮阳网(10%‒20%遮光率) | 避免日灼,防止光抑制,保障光合作用与糖分来源 | [ | |
| 合理树形(高纺锤形等)与夏季修剪(摘叶、转果) | 改善冠层光照,提高光合效能,显著提升果实总糖含量 | [ | |
| 精准水肥一体化(基于传感器监测),成熟期控水(亏缺灌溉)并与增施钾肥协同 | 诱导碳水化合物向果实分配,显著积累糖分;有机酸稳定或略升,增大糖酸比,风味更甜。严重干旱则抑制糖酸积累 | [ | |
品种与砧木选育 Cultivar and rootstock breeding | 选育抗逆(高温、干旱)品种 | 在胁迫条件下仍能维持较高的糖分与特定酸组分(如苹果酸)的积累能力 | [ |
| 选用耐旱、耐瘠薄砧木 | 增强树体抗逆性,稳定水分和养分供应,为糖酸品质形成提供基础 | [ | |
果实负载量调控 Regulation of fruit load | 通过疏花疏果保持适宜负载量 | 避免光合产物被“稀释”,保证单果营养供给,促进糖分和酸度充分积累,形成最佳糖酸比。过高负载量导致糖度、糖酸比下降,风味变差 | [ |
表3 影响苹果果实糖酸品质的栽培措施
Table 3 Cultivation practice influencing sugar-acid quality in apple fruit
调控途径 Regulatory pathway | 核心措施 Core practices | 对糖酸品质的主要影响 Primary impacts on sugar-acid quality | 参考文献 Reference |
|---|---|---|---|
主动微环境调控 Active microenvironment regulation | 行间生草/树盘覆盖 | 降低土温,减少蒸发,缓解高温胁迫,促进糖分平稳积累 | [ |
| 架设遮阳网(10%‒20%遮光率) | 避免日灼,防止光抑制,保障光合作用与糖分来源 | [ | |
| 合理树形(高纺锤形等)与夏季修剪(摘叶、转果) | 改善冠层光照,提高光合效能,显著提升果实总糖含量 | [ | |
| 精准水肥一体化(基于传感器监测),成熟期控水(亏缺灌溉)并与增施钾肥协同 | 诱导碳水化合物向果实分配,显著积累糖分;有机酸稳定或略升,增大糖酸比,风味更甜。严重干旱则抑制糖酸积累 | [ | |
品种与砧木选育 Cultivar and rootstock breeding | 选育抗逆(高温、干旱)品种 | 在胁迫条件下仍能维持较高的糖分与特定酸组分(如苹果酸)的积累能力 | [ |
| 选用耐旱、耐瘠薄砧木 | 增强树体抗逆性,稳定水分和养分供应,为糖酸品质形成提供基础 | [ | |
果实负载量调控 Regulation of fruit load | 通过疏花疏果保持适宜负载量 | 避免光合产物被“稀释”,保证单果营养供给,促进糖分和酸度充分积累,形成最佳糖酸比。过高负载量导致糖度、糖酸比下降,风味变差 | [ |
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