中国梨果实挥发性物质鉴定及酯类物质生物合成相关基因表达的研究
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摘要
果实香气是重要的果实品质性状之一,本研究利用固相微萃取技术(Solid Phase Microextraction, SPME)结合气相色谱-质谱联用仪(Gas Chromatography/Mass Spetrum, GC/MS)鉴定了中国梨主要栽培系统果实挥发性物质,同时研究了酯类物质生物合成途径编码脂氧合酶、醇脱氢酶及醇酰基转移酶等基因在秋子梨品种后熟及贮藏中的表达特性,以期为改进果实品质提供理论依据。本研究主要结果如下：
1、以‘库尔勒香梨’为试验材料对固相微萃取技术进行优化。结果发现,65μm的PDMS/DVB萃取头具有较好的萃取效果；优化后的萃取条件为：样品量为10g,萃取温度40℃,萃取时间45min。
2、利用电子鼻对不同栽培系统梨品种的果实挥发性物质进行检测分析。Alpha FOX4000的18个传感器能对不同栽培系统的梨品种挥发性组分产生不同的反应值；运用主成分分析(PCA)能够将所检测的8个品种聚为两大类：一类表现为传感器反应值较大,挥发性物质浓度较高；另一类则表现为传感器反应值较低,挥发性物质浓度较低。运用判别因子分析(DFA)可以将所检测的8个品种根据种类或产地很好地进行区分。因此,利用电子鼻检测挥发性物质结合一定的数学分析能对不同栽培系统梨品种进行区分。
3、从不同栽培系统的11个梨品种果实中共检测到70种挥发性物质,包括3种醇类、6种醛类、52种酯类、3种酸类、3种萜类和3种酮类化合物；酯类物质为主要的挥发性物质；根据各组分在品种中的相对含量及其阈值浓度看,已醛、乙酸已酯、己酸乙酯、2-甲基丁酸乙酯和丁酸乙酯是主要的挥发性物质和特征香气成分。秋子梨系统品种果实的挥发性物质含量最高,尤其是酯类物质含量要显著高于其他系统各品种；基于挥发性组分对11个品种进行主成分分析可以将其分为四大类。
4、对三个秋子梨品种‘油红梨’‘苹香梨’及‘小香水’梨果实后熟过程中果实挥发性组分的变化进行了跟踪调查,共检测到数十种种挥发性物质,其中酯类物质最多。果实后熟过程中,酯类物质含量逐渐升高,而醛类物质含量逐渐下降,使得果实具有浓郁的香气。‘苹香梨’果实基因表达结果显示,乙烯合成相关基因与酯类物质合成基因具有一定的相关性,乙烯合成途径中的PuACS1、PuACO1、 PuACO6与酯类物质合成途径中的PuLOX1、PuLOX8、PuADH2及PuAAT均在果实后熟第3d达到最高表达丰度,而PuACO2、PuACO5及酯类合成途径中的PuADH3等基因的表达丰度出现在果实后熟的第6d。当酯类合成相关底物供给充足的情况下,乙烯调控PuAAT基因表达使得果实中酯类物质合成增加。
5、醛类物质和酯类物质在‘软儿梨,果实贮藏过程中的变化趋势不同,随着果实贮藏期延长,果实酯类物质含量呈现出升高的趋势,而醛类物质含量维持在一定的水平。1-MCP处理‘软儿梨’果实后,酯类物质释放明显受1-MCP抑制,而醛类物质则不明显。1-MCP处理果实的PuLOXs和PuAAT等基因的表达受到抑制,而PuADHs等基因的表达不受抑制,表明在果实酯类物质合成中,PuLOXs和PuAAT等基因与果实酯类物质合成相关,而PuADH并不是果实酯类物质合成途径中的关键限制因子。
Fruit aroma is one of the most important attributes of fruit quality. In our research, we compared different aromatic constituents of different species and cultivars native to China using SPME and GC/MS. We also studied the expression of some genes which are related to ester biosynthesis including PuLOXs, PuADHs and PuAAT in the fruits of Pyrus ussuriensis during ripening and storage. The main results are as follows:
1. We optimized the different parameters of SPME using'Kuerle fragrance'pear as material. Results showed that65μm PDMS/DVB fiber had the best effect of extraction; the optimum factors such as sample volume, extraction time and temperature were10g,45min and40℃respectively.
2. In this study, volatile compounds of pear cultivars originating from different species were detected by using an electronic nose. The results revealed that18sensors of the Alpha FOX4000electronic nose responded differently to the volatile compounds of pear cultivars. Likewise, different cultivars resulted in different responses to these sensors. The eight cultivars were clustered into two groups by principal component analysis (PCA). One group with a high concentration of some volatile compounds exhibited higher response values to sensors, and another group with a low concentration of some volatiles showed lower response values to sensors. In addition, these cultivars were clearly differentiated on the basis of their origination and producing area using discrimination factor analysis (DFA). Therefore, the detection of volatile compounds using electronic nose combining with appropriate mathematic analysis could differentiate pear cultivars.
3. We compared the different volatile compounds of11pear cultivars belonging to different species. A total of70volatile compounds including three alcohols, six aldehydes, fifty-two esters, three acids, three terpenes and three ketones were identified and quantified. Ester was the dominant chemical class of which a total of52compounds were detected. Hexanal, hexyl acetate, ethyl hexanoate, ethyl2-methylbutanoate and ethyl butanoate were major volatile compounds or character impact odorants. Pyrus ussuriensis Maxim showed a higher concentration of volatiles, especially ester content which was dramatically higher as compared with the other Asian pear species. Cultivars were clearly grouped into four clusters based on the quantity and occurrence of members of different classes. Clusters of different cultivars could be discriminated from each others by the first two PCs (Principal Components). It might be possible to discriminate different Pyrus cultivars by principal component analysis. Volatile profiles of pear aroma were influenced by cultivars qualitatively and quantitatively.
4. The volatile compounds of three culitvars of Pyrus ussuriensis Maxim.including'Pingxiangli','Xiaoxiangshui'and 'Youhongli' pear fruit from different ripening stage were extracted by headspace solidphase microextraction and analyzed by gas chromatography-mass spectrometry. More than twenty volatile compounds were detected at different ripening stage. Ester was predominant among the volatile profile and increased dramatically during ripening, while the content of aldehyde decreased. The gene expression of'Pingxiangli'pear fruit showed that it was highly correlated to ethylene and ester biosynthesis. The transcription level of genes including PuACSl, PuACO1, PuACO6, PuLOXl, PuLOX8, PuADH2and PuAAT were on the peak at the third day of ripening in'Pingxiangli'pear fruit. Transcription levels of PuACO2, PuACO5and PuADH3reached to peak on the6th day of ripening. Taking the production of ester during ripening of 'Pingxiangli'pear, we speculated on the condition of enough substrate, ethylene regulated the expression of PuAAT, and then enhanced the production of ester.
5. The content of volatile compounds in1-MCP treated fruits was inhibited obviously during storage of'Ruanerli' pear. As important constituents of fruit aroma, the changes of ester and aldehyde were different during all stroage time. The content of ester increased with the course of ripening, while aldehyde remained a stable content. As compared with untreated fruit, the concentration of ester in1-MCP treated fruit was lower. Nevertheless there wasn't any difference in the concentration of aldehyde between treated and untreated fruits. The expression of PuLOXs and PuAAT were inhibited in1-MCP treated fruits, while PuADHs weren't suppressed in treated fruits. These results interpreted that PuLOXs and PuAAT might to be the rate limiting steps for ester biosynthesis, while puADH wasn't.
1、以‘库尔勒香梨’为试验材料对固相微萃取技术进行优化。结果发现,65μm的PDMS/DVB萃取头具有较好的萃取效果；优化后的萃取条件为：样品量为10g,萃取温度40℃,萃取时间45min。
2、利用电子鼻对不同栽培系统梨品种的果实挥发性物质进行检测分析。Alpha FOX4000的18个传感器能对不同栽培系统的梨品种挥发性组分产生不同的反应值；运用主成分分析(PCA)能够将所检测的8个品种聚为两大类：一类表现为传感器反应值较大,挥发性物质浓度较高；另一类则表现为传感器反应值较低,挥发性物质浓度较低。运用判别因子分析(DFA)可以将所检测的8个品种根据种类或产地很好地进行区分。因此,利用电子鼻检测挥发性物质结合一定的数学分析能对不同栽培系统梨品种进行区分。
3、从不同栽培系统的11个梨品种果实中共检测到70种挥发性物质,包括3种醇类、6种醛类、52种酯类、3种酸类、3种萜类和3种酮类化合物；酯类物质为主要的挥发性物质；根据各组分在品种中的相对含量及其阈值浓度看,已醛、乙酸已酯、己酸乙酯、2-甲基丁酸乙酯和丁酸乙酯是主要的挥发性物质和特征香气成分。秋子梨系统品种果实的挥发性物质含量最高,尤其是酯类物质含量要显著高于其他系统各品种；基于挥发性组分对11个品种进行主成分分析可以将其分为四大类。
4、对三个秋子梨品种‘油红梨’‘苹香梨’及‘小香水’梨果实后熟过程中果实挥发性组分的变化进行了跟踪调查,共检测到数十种种挥发性物质,其中酯类物质最多。果实后熟过程中,酯类物质含量逐渐升高,而醛类物质含量逐渐下降,使得果实具有浓郁的香气。‘苹香梨’果实基因表达结果显示,乙烯合成相关基因与酯类物质合成基因具有一定的相关性,乙烯合成途径中的PuACS1、PuACO1、 PuACO6与酯类物质合成途径中的PuLOX1、PuLOX8、PuADH2及PuAAT均在果实后熟第3d达到最高表达丰度,而PuACO2、PuACO5及酯类合成途径中的PuADH3等基因的表达丰度出现在果实后熟的第6d。当酯类合成相关底物供给充足的情况下,乙烯调控PuAAT基因表达使得果实中酯类物质合成增加。
5、醛类物质和酯类物质在‘软儿梨,果实贮藏过程中的变化趋势不同,随着果实贮藏期延长,果实酯类物质含量呈现出升高的趋势,而醛类物质含量维持在一定的水平。1-MCP处理‘软儿梨’果实后,酯类物质释放明显受1-MCP抑制,而醛类物质则不明显。1-MCP处理果实的PuLOXs和PuAAT等基因的表达受到抑制,而PuADHs等基因的表达不受抑制,表明在果实酯类物质合成中,PuLOXs和PuAAT等基因与果实酯类物质合成相关,而PuADH并不是果实酯类物质合成途径中的关键限制因子。
Fruit aroma is one of the most important attributes of fruit quality. In our research, we compared different aromatic constituents of different species and cultivars native to China using SPME and GC/MS. We also studied the expression of some genes which are related to ester biosynthesis including PuLOXs, PuADHs and PuAAT in the fruits of Pyrus ussuriensis during ripening and storage. The main results are as follows:
1. We optimized the different parameters of SPME using'Kuerle fragrance'pear as material. Results showed that65μm PDMS/DVB fiber had the best effect of extraction; the optimum factors such as sample volume, extraction time and temperature were10g,45min and40℃respectively.
2. In this study, volatile compounds of pear cultivars originating from different species were detected by using an electronic nose. The results revealed that18sensors of the Alpha FOX4000electronic nose responded differently to the volatile compounds of pear cultivars. Likewise, different cultivars resulted in different responses to these sensors. The eight cultivars were clustered into two groups by principal component analysis (PCA). One group with a high concentration of some volatile compounds exhibited higher response values to sensors, and another group with a low concentration of some volatiles showed lower response values to sensors. In addition, these cultivars were clearly differentiated on the basis of their origination and producing area using discrimination factor analysis (DFA). Therefore, the detection of volatile compounds using electronic nose combining with appropriate mathematic analysis could differentiate pear cultivars.
3. We compared the different volatile compounds of11pear cultivars belonging to different species. A total of70volatile compounds including three alcohols, six aldehydes, fifty-two esters, three acids, three terpenes and three ketones were identified and quantified. Ester was the dominant chemical class of which a total of52compounds were detected. Hexanal, hexyl acetate, ethyl hexanoate, ethyl2-methylbutanoate and ethyl butanoate were major volatile compounds or character impact odorants. Pyrus ussuriensis Maxim showed a higher concentration of volatiles, especially ester content which was dramatically higher as compared with the other Asian pear species. Cultivars were clearly grouped into four clusters based on the quantity and occurrence of members of different classes. Clusters of different cultivars could be discriminated from each others by the first two PCs (Principal Components). It might be possible to discriminate different Pyrus cultivars by principal component analysis. Volatile profiles of pear aroma were influenced by cultivars qualitatively and quantitatively.
4. The volatile compounds of three culitvars of Pyrus ussuriensis Maxim.including'Pingxiangli','Xiaoxiangshui'and 'Youhongli' pear fruit from different ripening stage were extracted by headspace solidphase microextraction and analyzed by gas chromatography-mass spectrometry. More than twenty volatile compounds were detected at different ripening stage. Ester was predominant among the volatile profile and increased dramatically during ripening, while the content of aldehyde decreased. The gene expression of'Pingxiangli'pear fruit showed that it was highly correlated to ethylene and ester biosynthesis. The transcription level of genes including PuACSl, PuACO1, PuACO6, PuLOXl, PuLOX8, PuADH2and PuAAT were on the peak at the third day of ripening in'Pingxiangli'pear fruit. Transcription levels of PuACO2, PuACO5and PuADH3reached to peak on the6th day of ripening. Taking the production of ester during ripening of 'Pingxiangli'pear, we speculated on the condition of enough substrate, ethylene regulated the expression of PuAAT, and then enhanced the production of ester.
5. The content of volatile compounds in1-MCP treated fruits was inhibited obviously during storage of'Ruanerli' pear. As important constituents of fruit aroma, the changes of ester and aldehyde were different during all stroage time. The content of ester increased with the course of ripening, while aldehyde remained a stable content. As compared with untreated fruit, the concentration of ester in1-MCP treated fruit was lower. Nevertheless there wasn't any difference in the concentration of aldehyde between treated and untreated fruits. The expression of PuLOXs and PuAAT were inhibited in1-MCP treated fruits, while PuADHs weren't suppressed in treated fruits. These results interpreted that PuLOXs and PuAAT might to be the rate limiting steps for ester biosynthesis, while puADH wasn't.
引文
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