Biotechnology Bulletin ›› 2026, Vol. 42 ›› Issue (3): 79-95.doi: 10.13560/j.cnki.biotech.bull.1985.2025-1025
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DU Lian-da(
), WEI Meng-meng, CHEN Ze, GUO Wei, ZHAO Ting-ting(
), HU Da-gang(
)
Received:2025-09-26
Online:2026-03-26
Published:2026-04-23
Contact:
ZHAO Ting-ting, HU Da-gang
E-mail:liandadu@126.com;datingsdau@126.com;fap_296566@163.com
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.
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
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 | [ |
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 | [ |
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 | [ |
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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