不同架式‘爱神玫瑰’葡萄果实成熟期间单萜积累及相关基因的表达
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摘要
【目的】探讨不同架式对葡萄果实成熟期间单萜类化合物合成的影响,进一步从基因表达水平揭示基因转录与单萜积累的关系,以期为生产中架式选择及葡萄果实香味品质的提高提供一定的理论依据。【方法】以T型和V型架式栽培的‘爱神玫瑰’葡萄果实为试材,于果实成熟初期(花后30 d)开始取样,直至果实完全成熟。连续两年常规方法测定果实样品可溶性固形物及可滴定酸含量,利用顶空固相微萃取结合气相色谱与质谱联用技术(SPEME-GC-MS)测定果实中单萜类组分和含量的变化,采用实时荧光定量PCR技术分析单萜合成途径中关键基因脱氧木酮糖磷酸合酶基因(DXS1和DXS3)、脱氧木酮糖磷酸还原异构酶基因(DXR)、异戊烯基焦磷酸还原酶基因(HDR)、里那醇合成酶基因(Liner syn)和萜品醇合成酶基因(Terp syn)的表达变化。【结果】随着果实成熟,可溶性固形物含量逐渐升高,而可滴定酸含量逐渐降低。成熟期的T型架葡萄果实可溶性固形物含量显著高于V型架,可滴定酸含量没有显著差异。2016年和2017年两种架式果实样品中分别检测到27和28种单萜类化合物。检测结果表明不同架式主要萜烯类化合物组分不尽相同,随着果实成熟,主要萜烯类成分也发生变化。成熟时,T型架果实中主要单萜类化合物有里那醇、柠檬烯、α-萜品醇、β-cis-罗勒烯和香叶醇;V型架主要有里那醇、α-萜品醇、柠檬烯、橙花醚和β-cis-罗勒烯等,其中以里那醇含量最高。2016年成熟期T型架果实单萜总量达到108.18μg·L~(-1),是V型架最高含量的1.9倍。而2017年成熟期T型架果实单萜总量达到403.24μg·L~(-1),是V型架最高含量的1.5倍;大多数单萜类化合物含量在成熟时表现为T型架显著高于V型架。在整个果实成熟期间,两种架式葡萄果实单萜类化合物积累表现为两种变化模式,包括里那醇、香叶醇、橙花醇及萜品醇等在内的大部分化合物遵循第一种模式,即在果实成熟时含量达到最高。但是不同架式表现又略微不同,(E,Z)-别罗勒烯、β-cis-罗勒烯、柠檬烯和α-萜品醇等化合物在T型架果实中表现为先下降,花后57 d急剧升高,成熟后期(花后76 d)又下降的趋势。而在V型架果实中这些化合物含量随着果实成熟逐渐上升,花后48 d达到积累高峰,之后又逐渐下降至最低含量。另外,果实成熟期间单萜合成途径基因(DXS1、DXS3、DXR、DHR、Liner syn和Terp syn)表达量随着果实成熟呈上升趋势。不同架式葡萄果实成熟期间单萜总量积累规律与DXS3、HDR、Liner syn和Terp syn表达规律相似。成熟期T型架果实中各个基因表达量明显高于V型架,与单萜类化合物积累模式相一致。【结论】T型架式栽培更有利于果实单萜类物质的积累,T型架式单萜类化合物的高效积累与其代谢途径多个关键酶基因高效表达密切相关。
【Objective】 The effects of two trellis systems on the synthesis of monoterpenes and the expression of its biosynthesis related genes during the maturity of grape berries were studied, in order to reveal the relationship between gene transcription and monoterpenoids accumulation and provide a theoretical basis for trellis selection in production and improvement of aroma quality of grape fruits. 【Method】 The grape berries of Aishen Meigui cultivated in the T and V trellis were used as materials.The berry samples were collected from the beginning of the color change period until the fruit ripened completely. Total soluble solid and titratable acid content in fruits were detected. The changes of monoterpene components and contents in fruits were determined by headspace solid-phase microextraction combined with gas chromatography-mass spectrometry(SPEME-GC-MS) in two consecutive years. Additionally, the expression of key enzyme genes in monoterpene biosynthesis pathway including 1-deoxy-D-xylulose5-phosphate synthase genes(DXS1 and DXS3), 1-deoxy-D-xylulose-5-phosphate reductoisomerase gene(DXR), isopentenyl pyrophosphate reductase gene(HDR), linalool synthase gene(Liner syn) and terpineol synthase gene(Terp syn) were analyzed by real-time fluorescence quantitative analysis(qPCR).【Result】During grape berries maturity, the total soluble solids content increased gradually, while titratable acid content decreased gradually. The total soluble solids content of the Aishen Meigui grape cultivated in T trellis was significantly higher than that of grape cultivated in V trellis at ripening stage, and there was no significant difference in titratable acid content. 27 and 28 monoterpene compounds were detected in the grape berries of Aishen Meigui in 2016 and 2017,respectively. The main monoterpene components changed during berry maturity and were different in grape berries cultivated in T and V trellis. As the results shown, linalool, limonene, α-Terpineol, β-cis-Ocimene and geraniol were the main monoterpenes in grape berries cultivated in T trellis, while abundant linalool, α-Terpineol, limonene, nerol oxide and β-cis-Ocimene were detected in the berries cultivated in V trellis, and the content of linalool was the highest among them. At the maturity stage, total monoterpene content of T trellis grape berries reached 108.18 μg·L~(-1), which was 1.9 times higher than that of V trellis berries in 2016. While in 2017, total monoterpene content of T trellis grape berries reached 403.24 μg·L~(-1), which was 1.5 times higher than that of V trellis berries. The content of most monoterpene compounds in T trellis fruits were significantly higher than that in V trellis fruits at maturity. During grape berry development, the changes of monoterpenes accumulation in grape berries cultivated in both two trellis systems exhibited two patterns. Most of the monoterpenes including linalool, geraniol, nerol and α-Terpineol followed the first pattern: the content of monoterpenes reached the peak at maturity. But differences were observed between trellises. In the T trellis fruits, the content of compounds, such as(E,Z)-Allo-Ocimene, β-cis-Ocimene, limonene and α-terpineol, firstly decreased, and then increased sharply at 57 days after flowering, followed by a reduction during the later part of ripening stage(76 days after flowering).However, in V trellis berries, the content of these compounds increased gradually during fruit ripening, peaked at 48 days after flowering, and then decreased to the lowest level. The expression of monoterpene synthesis pathway genes(DXS1, DXS3, DXR, DHR,Liner syn and Terp syn) increased with berry ripening, of them, the changes of DXS3, HDR, Liner syn and Terp syn expression were corresponding well to the total monoterpenes accumulation patterns in grape berries cultivated in distinct trellis systems during ripening. The expression of each gene in T-trellis grape berries was significantly higher than that in V-trellis berries, which was consistent with the accumulation pattern of different monoterpenes. 【Conclusion】More accumulation of monoterpenes were detected in Aishen Meigui grape berries cultivated in T trellis. The synthesis of monoterpenes was highly correlated with the expression level of several key genes in the synthesis pathway.
【Objective】 The effects of two trellis systems on the synthesis of monoterpenes and the expression of its biosynthesis related genes during the maturity of grape berries were studied, in order to reveal the relationship between gene transcription and monoterpenoids accumulation and provide a theoretical basis for trellis selection in production and improvement of aroma quality of grape fruits. 【Method】 The grape berries of Aishen Meigui cultivated in the T and V trellis were used as materials.The berry samples were collected from the beginning of the color change period until the fruit ripened completely. Total soluble solid and titratable acid content in fruits were detected. The changes of monoterpene components and contents in fruits were determined by headspace solid-phase microextraction combined with gas chromatography-mass spectrometry(SPEME-GC-MS) in two consecutive years. Additionally, the expression of key enzyme genes in monoterpene biosynthesis pathway including 1-deoxy-D-xylulose5-phosphate synthase genes(DXS1 and DXS3), 1-deoxy-D-xylulose-5-phosphate reductoisomerase gene(DXR), isopentenyl pyrophosphate reductase gene(HDR), linalool synthase gene(Liner syn) and terpineol synthase gene(Terp syn) were analyzed by real-time fluorescence quantitative analysis(qPCR).【Result】During grape berries maturity, the total soluble solids content increased gradually, while titratable acid content decreased gradually. The total soluble solids content of the Aishen Meigui grape cultivated in T trellis was significantly higher than that of grape cultivated in V trellis at ripening stage, and there was no significant difference in titratable acid content. 27 and 28 monoterpene compounds were detected in the grape berries of Aishen Meigui in 2016 and 2017,respectively. The main monoterpene components changed during berry maturity and were different in grape berries cultivated in T and V trellis. As the results shown, linalool, limonene, α-Terpineol, β-cis-Ocimene and geraniol were the main monoterpenes in grape berries cultivated in T trellis, while abundant linalool, α-Terpineol, limonene, nerol oxide and β-cis-Ocimene were detected in the berries cultivated in V trellis, and the content of linalool was the highest among them. At the maturity stage, total monoterpene content of T trellis grape berries reached 108.18 μg·L~(-1), which was 1.9 times higher than that of V trellis berries in 2016. While in 2017, total monoterpene content of T trellis grape berries reached 403.24 μg·L~(-1), which was 1.5 times higher than that of V trellis berries. The content of most monoterpene compounds in T trellis fruits were significantly higher than that in V trellis fruits at maturity. During grape berry development, the changes of monoterpenes accumulation in grape berries cultivated in both two trellis systems exhibited two patterns. Most of the monoterpenes including linalool, geraniol, nerol and α-Terpineol followed the first pattern: the content of monoterpenes reached the peak at maturity. But differences were observed between trellises. In the T trellis fruits, the content of compounds, such as(E,Z)-Allo-Ocimene, β-cis-Ocimene, limonene and α-terpineol, firstly decreased, and then increased sharply at 57 days after flowering, followed by a reduction during the later part of ripening stage(76 days after flowering).However, in V trellis berries, the content of these compounds increased gradually during fruit ripening, peaked at 48 days after flowering, and then decreased to the lowest level. The expression of monoterpene synthesis pathway genes(DXS1, DXS3, DXR, DHR,Liner syn and Terp syn) increased with berry ripening, of them, the changes of DXS3, HDR, Liner syn and Terp syn expression were corresponding well to the total monoterpenes accumulation patterns in grape berries cultivated in distinct trellis systems during ripening. The expression of each gene in T-trellis grape berries was significantly higher than that in V-trellis berries, which was consistent with the accumulation pattern of different monoterpenes. 【Conclusion】More accumulation of monoterpenes were detected in Aishen Meigui grape berries cultivated in T trellis. The synthesis of monoterpenes was highly correlated with the expression level of several key genes in the synthesis pathway.
引文
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[2]XU X Q,CHENG G,DUAN L L,JIANG R,PAN Q H,DUAN C Q,WANG J.Effect of training systems on fatty acids and their derived volatiles in cabernet sauvignon grapes and wines of the north foot of Mt.Tianshan.Food Chemistry,2015,181:198-206.
[3]XU H Y,SUN L,ZHANG G J,YAN A L.‘Aishenmeigui’:An extremely early maturing seedless grape with muscat flavor.Vitis:Journal of Grapevine Research,2014,53(1):53.
[4]MARTIN D M,CHIANG A,LUND S T,BOHLMANN J.Biosynthesis of wine aroma:Transcript profiles of hydroxyl methyl butenyl diphosphate reductase,geranyl diphosphate synthase,and linalool/nerolidol synthase parallel monoterpenol glycoside accumulation in Gewürztraminer grapes.Planta,2012,236:919-929.
[5]EI HADI M A,ZHANG F J,WU F F,ZHOU C H,TAO J.Advances in fruit aroma volatile research.Molecules,2013,18(7):8200-8229.
[6]ZHANG E P,CHAI F M,ZHANG H H,LI S H,LIANG Z C,FAN PG.Effects of sunlight exclusion on the profiles of monoterpene biosynthesis and accumulation in grape exocarp and mesocarp.Food Chemistry,2017,237:379-389.
[7]FRIEDEL M,FROTSCHER J,NITSCH M,HOFMANN M,BOGS J,STOLL M,DIETRICH H.Light promotes expression of monoterpene and flavonol metabolic genes and enhances flavour of winegrape berries(Vitis vinifera l.cv.Riesling).Australian Journal of Grape&Wine Research,2016,22(3):409-421.
[8]SCHULTZ H R.Grape canopy structure,light microclimate and photosynthesis.1.A two-dimensional model of the spatial distribution of surface area densities and leaf ages in two canopy systems.VITIS-Journal of Grapevine Research,1995,34(4):211-215.
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