椪柑和脐橙果实柠檬酸合成与降解相关基因表达及其调控研究
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摘要
柠檬酸是柑橘果实的主要有机酸,其合成途径以三羧酸(TCA)循环为主,其降解有GABA途径和GS途径。本研究以椪柑(Citrus reticulata)和脐橙(Citrus sinensis)果实为试材,分析了不同种类与品种、生境及采后热处理对柠檬酸代谢及相关基因表达的影响,探讨了柑橘果实柠檬酸代谢的分子调控机制。主要结果如下：
1、椪柑果实发育过程中柠檬酸呈先上升后下降的变化,且早熟椪柑品种果实的柠檬酸含量始终低于普通碰柑品种果实,其变化趋势与总有机酸和TA含量相一致。进一步研究柠檬酸代谢相关基因的结果显示,柠檬酸合成相关基因,CitCSs、 CitPEPCs和CitPEPCKs的表达在不同碰柑品种果实间无显著差异；而柠檬酸降解相关基因,CitAco3、CitIDH1/3、CitGAD4/5、CitGS2的转录丰度在早熟椪柑果实中均显著高于普通椪柑。推测早熟椪柑果实中的低柠檬酸含量,可能与其合成速率无明显关系,而与其降解途径中相关酶编码基因的级联上调有关。
2、以‘纽荷尔’、‘朋娜’脐橙果实为对象,研究不同生境对有机酸代谢的影响,结果发现赣州(北纬25°26')脐橙成熟果实有机酸含量显著低于松阳(北纬28°27')脐橙的,该差异也以柠檬酸含量差异为主。与椪柑果实柠檬酸代谢类似,不同生境脐橙果实柠檬酸水平差异与柠檬酸合成相关基因无显著相关性,而与柠檬酸降解基因有关。赣州脐橙果实的CitAco3、CitIDH1和CitGS2表达水平显著高于松阳脐橙果实,这些基因的高水平表达有利于果实柠檬酸的降解,而GABA途径的CitGADs基因的表达在两个生境间无显著差异。
3、研究了采后热处理对柑橘果实有机酸水平的影响,发现采后40℃热空气处理2d对椪柑果实的CCI和TSS无显著影响,但可显著促进果实有机酸尤其是柠檬酸含量的下降。基因表达结果显示,热处理显著诱导了CitAco3、CitDH2/3及CitGAD4等基因的表达增强,而对CitGS2表达无影响,表明热处理加速柠檬酸分解主要通过GABA途径,尤其是通过CitAco3-CitIDH2/3-CitGAD4的级联调节。
综述所述,本研究中不同品种、不同生境和采后热处理的柑橘果实有机酸含量差异均由柠檬酸水平决定,柠檬酸降解的速率决定了柑橘果实有机酸含量,但其降解机制存在差异。椪柑不同品种果实间的柠檬酸差异可能由GABA和GS降解途径共同参与,而不同生境和采后热处理则分别特异上调GS途径和GABA途径。表明不同品种间柠檬酸降解调控位点具有多样性,而不同生境和采后热处理降酸可有特异的途径,其中GABA途径可能与逆境胁迫相关。
Citric acid, the major organic acid in citrus fruit, is produced by the tricarboxylic acid (TCA) cycle and degraded through gamma-aminobutirate (GABA) shunt and glutamine synthesis (GS) pathway. In this study, Ponkan (Citrus reticulata) and navel orange (Citrus sinensis) fruits were used to investigate the effects of varieties and cultivars, ecological environment, and postharvest heat treatment on citric acid metabolism and related gene expression, and the molecular mechanisms of citric acid metabolism and its regulation in citrus fruit were discussed. The main results are as follows:
1. During the development of Ponkan fruit, citric acid increased in the early stage which was followed by a later decrease until the maturation stage. Earlier-ripening mutant Ponkan (ZSPG) fruit accumulated lower concentration of citric acid than the normal Ponkan (PTPG), which was consistent with their total organic acids and TA. Results of gene expression showed that there was no significant difference in the expression of citric acid biosynthesis related genes such as CitCSs, CitPEPCs and CitPEPCKs between PTPG and ZSPG fruit. However, significant higher transcript levels of CitAco3, CitIDHl/3, CitGAD4/5and CitGS2, which took part in the degradation of citric acid, were found in ZSPG than PTPG fruits. Such results indicated that lower citric acid level in ZSPG was caused by the upregulation of cascade genes encoded enzymes involved in citric acid degradation, rather than its biosynthesis.
2. Effects of ecological environment on organic acid metabolism during fruit development were investigated by using two cultivars of navel orange, i.e.'Newhall' and 'SkaggsBonanza', as fruit materials. Organic acids in matured navel orange cultivated in Ganzhou (GZ) city of Jiangxi province (25°26'N) was significantly lower than those in Songyang (SY) county of Zhejiang province (28°27'N), and the difference was mainly caused by citric acid contents. Similar to the result of Ponkan, the different citric acid levels in navel orange in different habitats were closely related with the degradation of citric acid, rather than its biosynthesis, since the expression levels of CitAco3, CitIDH1and CitGS2in navel orange fruits grown in GZ were significantly higher than those in SY. Meanwhile, the expression of CitGADs genes, which involved in GABA shunt, had no significantly differences between GZ and SY.
3. Hot air (40℃,2d) treatment (HAT) showed significant effects on the degradation of organic acids, especially on citric acid, but it had no obvious effect on CCI or TSS. Gene expression results showed that HAT significantly up-regulated the expression of CitAco3, CitIDH2/3and CitGAD4, while it had no effect on CitGS2. Our results indicated that HAT might accelerate citric acid degradation via the GABA shunt pathway, particularly by modulating the CitAco3-CitIDH2/3-CitGAD4cascade.
In conclusion, the difference of organic acids content in citrus of different cultivars, habitats and postharvest heat treatment were all determined by their citric acid levels, and degradation rate, rather than the biosynthesis rate, of citric acid showed significant effect on the total organic acids concentration in citrus fruit, though the mechanisms varied for different cases. The different citric acid levels between two Ponkan cultivars might be caused by both GS and GABA degradation pathway, while ecological environment and HAT treatment up-regulated GS and GABA degradation pathway, respectively. The results indicated that the regulatory sites of citric acid degradation varied under different circumstances, and GABA pathway is also related to environment stress.
1、椪柑果实发育过程中柠檬酸呈先上升后下降的变化,且早熟椪柑品种果实的柠檬酸含量始终低于普通碰柑品种果实,其变化趋势与总有机酸和TA含量相一致。进一步研究柠檬酸代谢相关基因的结果显示,柠檬酸合成相关基因,CitCSs、 CitPEPCs和CitPEPCKs的表达在不同碰柑品种果实间无显著差异；而柠檬酸降解相关基因,CitAco3、CitIDH1/3、CitGAD4/5、CitGS2的转录丰度在早熟椪柑果实中均显著高于普通椪柑。推测早熟椪柑果实中的低柠檬酸含量,可能与其合成速率无明显关系,而与其降解途径中相关酶编码基因的级联上调有关。
2、以‘纽荷尔’、‘朋娜’脐橙果实为对象,研究不同生境对有机酸代谢的影响,结果发现赣州(北纬25°26')脐橙成熟果实有机酸含量显著低于松阳(北纬28°27')脐橙的,该差异也以柠檬酸含量差异为主。与椪柑果实柠檬酸代谢类似,不同生境脐橙果实柠檬酸水平差异与柠檬酸合成相关基因无显著相关性,而与柠檬酸降解基因有关。赣州脐橙果实的CitAco3、CitIDH1和CitGS2表达水平显著高于松阳脐橙果实,这些基因的高水平表达有利于果实柠檬酸的降解,而GABA途径的CitGADs基因的表达在两个生境间无显著差异。
3、研究了采后热处理对柑橘果实有机酸水平的影响,发现采后40℃热空气处理2d对椪柑果实的CCI和TSS无显著影响,但可显著促进果实有机酸尤其是柠檬酸含量的下降。基因表达结果显示,热处理显著诱导了CitAco3、CitDH2/3及CitGAD4等基因的表达增强,而对CitGS2表达无影响,表明热处理加速柠檬酸分解主要通过GABA途径,尤其是通过CitAco3-CitIDH2/3-CitGAD4的级联调节。
综述所述,本研究中不同品种、不同生境和采后热处理的柑橘果实有机酸含量差异均由柠檬酸水平决定,柠檬酸降解的速率决定了柑橘果实有机酸含量,但其降解机制存在差异。椪柑不同品种果实间的柠檬酸差异可能由GABA和GS降解途径共同参与,而不同生境和采后热处理则分别特异上调GS途径和GABA途径。表明不同品种间柠檬酸降解调控位点具有多样性,而不同生境和采后热处理降酸可有特异的途径,其中GABA途径可能与逆境胁迫相关。
Citric acid, the major organic acid in citrus fruit, is produced by the tricarboxylic acid (TCA) cycle and degraded through gamma-aminobutirate (GABA) shunt and glutamine synthesis (GS) pathway. In this study, Ponkan (Citrus reticulata) and navel orange (Citrus sinensis) fruits were used to investigate the effects of varieties and cultivars, ecological environment, and postharvest heat treatment on citric acid metabolism and related gene expression, and the molecular mechanisms of citric acid metabolism and its regulation in citrus fruit were discussed. The main results are as follows:
1. During the development of Ponkan fruit, citric acid increased in the early stage which was followed by a later decrease until the maturation stage. Earlier-ripening mutant Ponkan (ZSPG) fruit accumulated lower concentration of citric acid than the normal Ponkan (PTPG), which was consistent with their total organic acids and TA. Results of gene expression showed that there was no significant difference in the expression of citric acid biosynthesis related genes such as CitCSs, CitPEPCs and CitPEPCKs between PTPG and ZSPG fruit. However, significant higher transcript levels of CitAco3, CitIDHl/3, CitGAD4/5and CitGS2, which took part in the degradation of citric acid, were found in ZSPG than PTPG fruits. Such results indicated that lower citric acid level in ZSPG was caused by the upregulation of cascade genes encoded enzymes involved in citric acid degradation, rather than its biosynthesis.
2. Effects of ecological environment on organic acid metabolism during fruit development were investigated by using two cultivars of navel orange, i.e.'Newhall' and 'SkaggsBonanza', as fruit materials. Organic acids in matured navel orange cultivated in Ganzhou (GZ) city of Jiangxi province (25°26'N) was significantly lower than those in Songyang (SY) county of Zhejiang province (28°27'N), and the difference was mainly caused by citric acid contents. Similar to the result of Ponkan, the different citric acid levels in navel orange in different habitats were closely related with the degradation of citric acid, rather than its biosynthesis, since the expression levels of CitAco3, CitIDH1and CitGS2in navel orange fruits grown in GZ were significantly higher than those in SY. Meanwhile, the expression of CitGADs genes, which involved in GABA shunt, had no significantly differences between GZ and SY.
3. Hot air (40℃,2d) treatment (HAT) showed significant effects on the degradation of organic acids, especially on citric acid, but it had no obvious effect on CCI or TSS. Gene expression results showed that HAT significantly up-regulated the expression of CitAco3, CitIDH2/3and CitGAD4, while it had no effect on CitGS2. Our results indicated that HAT might accelerate citric acid degradation via the GABA shunt pathway, particularly by modulating the CitAco3-CitIDH2/3-CitGAD4cascade.
In conclusion, the difference of organic acids content in citrus of different cultivars, habitats and postharvest heat treatment were all determined by their citric acid levels, and degradation rate, rather than the biosynthesis rate, of citric acid showed significant effect on the total organic acids concentration in citrus fruit, though the mechanisms varied for different cases. The different citric acid levels between two Ponkan cultivars might be caused by both GS and GABA degradation pathway, while ecological environment and HAT treatment up-regulated GS and GABA degradation pathway, respectively. The results indicated that the regulatory sites of citric acid degradation varied under different circumstances, and GABA pathway is also related to environment stress.
引文
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