摘要
本文以‘左山一’、‘左山二’、‘双红’、‘双优’、‘双丰’5个山葡萄品种和2个山-欧杂种葡萄品种‘左红一’、‘左优红’果皮和酒为试验材料,应用HPLC-MS/MS技术,对果实发育过程中果皮及酒中花色苷和非花色苷酚种类及含量变化规律进行了研究。同时,以‘双红’山葡萄果实为试材,应用HPLC-MS/MS和实时荧光定量PCR技术对果实发育过程中酚类物质的含量变化及生物合成相关的14个结构基因mRNA转录水平进行了定量分析。
研究结果表明,不同品种果皮中花色苷组成不同,在7个品种果实发育过程中果皮中共检测到花色苷15种。其中,双糖苷6种,单糖苷9种,单糖苷中检测出乙酰化葡萄糖苷1种,香豆酰化葡萄糖苷1种。7个品种转色前均未检测到花色苷,随着果实发育,花色苷含量不断增加,到达成熟时含量最高。主成分分析表明,花青素类、二甲花翠素类和甲基花青素类花色苷对果皮中花色苷的组成起决定作用。7个品种酒中共检测到花色苷15种。其中,双糖苷6种,单糖苷9种,在果皮及酒中首次检测到花葵素-3,5-双葡萄糖苷,在酒中未检测出乙酰化和香豆酰化花色苷。主成分分析表明,花青素类、甲基花翠素类和花翠素类花色苷对酒中花色苷组成起决定作用。
7个品种果实发育过程中果皮中共检测到18种非花色苷酚。其中,苯甲酸类3种,肉桂酸类4种,黄烷-3-醇类3种,黄酮醇类5种,白藜芦醇类3种。果皮中非花色苷酚的变化为转色前含量呈上升趋势,转色期至100%着色呈下降趋势,而后缓慢上升至成熟。主成分分析表明,肉桂酸类、白藜芦醇类和黄酮醇类对果实发育过程中果皮中非花色苷酚的组成起决定作用。在7个品种酒中,共检测到30种非花色苷酚。其中,苯甲酸类5种,肉桂酸类7种,黄烷-3-醇类5种,黄酮醇类9种,白藜芦醇类4种。主成分分析表明,肉桂酸类、黄烷-3-醇类和苯甲酸类对酒中非花色苷酚组成起决定作用。
‘双红’山葡萄果实的生长曲线表现为双“S”型。转色前未检测到花色苷,转色后花色苷不断积累,至成熟含量最高,花翠素类、甲氧基类、单糖苷类花色苷与总花色苷的变化规律相一致。苯甲酸类花后2-6周未检测到,花后8周至100%着色含量呈上升趋势,而后下降至成熟。肉桂酸类、黄烷-3-醇类在果实整个发育过程中含量呈下降趋势。转色前未检测到白藜芦醇类物质,转色期至成熟含量呈上升趋势。
‘双红’果实发育过程中,酚类物质生物合成相关结构基因的表达因果实发育时期而不同。PAL、CHI1、F3H2、DFR、LDOX花后2-4周为上调表达,花后6周至转色期为下调表达,转色期至成熟为上调表达,转色期至成熟花色苷的含量变化与这5个酶基因的转录水平呈正相关；CHS3、RS、UFGT、GST、OMT花后2周至转色期为下调表达,转色期至成熟为上调表达,转色期至成熟花色苷的含量变化与CHS3、UFGT、GST的转录水平呈正相关,白藜芦醇类的含量变化与RS的转录水平呈正相关,甲氧基类花色苷的含量变化与OMT的转录水平呈正相关；F’3’H花后2-4周为上调表达,花后6周为下调表达,花后8周至成熟为上调表达,转色期至成熟花青素类花色苷的含量变化与F3'H的转录水平呈正相关；F3'5'H花后2周至转色期为下调表达,转色后为上调表达,转色期至成熟花翠素类花色苷的含量变化与F3'5'H的转录水平呈正相关ANR花后2周至转色期为上调表达,转色后为下调表达,花后2周至成熟表儿茶素的含量变化与ANR的转录水平呈正相关；LAR2花后2周至转色期为上调表达,转色后为下调表达,花后2周至成熟儿茶素的含量变化与LAR2的转录水平呈正相关。
The anthocyanin and non-anthocyanin phenolic compounds profiles in grapevine skins and wines from five varieties('Zuoshanyi','Zuoshaner','Shuanghong','Shuangyou' and 'Shuangfeng')of Vitis amurensis and two hybrid varieties('Zuohongyi' and 'Zuoyouhong') were investigated by HPLC-MS/MS for revealing the changes of their varieties and contents during berry development. In combination with real-time fluorescence quantitative PCR, the phenolic compounds and mRNA transcriptional level of 14 structural genes related to biosynthesis of phenolic compounds were analyzed in variety of'Shuanghong' during berry development.
The results indicated that the component of anthocyanins greatly varied with grapevine varieties. A total of 15 kinds of anthocyanins including 6 diglucosides and 9 monoglucosides of anthocyanins were identified in skins, in which one acetylglucoside and one coumarylglucoside were detected. No anthocyanins were detected until veraison, and then the anthocyanins contents increased with berry development and reached the maximum at the stage of maturity. Principal component analysis showed that cyanidin, malvidin and peonidin derivatives played key roles in the composition of anthocyanins in berry skins. In addition,15 kinds of anthocyanins were detected in wines, including 6 diglucosides and 9 monoglucosides of anthocyanidins. Pelargonidin-3,5-diglucosides was first found in the skins and wines, however, no acetyl and coumaryl anthocyanin was detected in wines. Accordingly, the cyanidin, petunidin and delphinidin derivatives played crucial roles in the composition of anthocyanins in wines, as indicated by principal component analysis.
A total of 18 kinds of non-anthocyanin phenolic compounds were identified in the skins during berry development, which included 3 benzoic acids,4 cinnamic acids,3 flavan-3-ols,5 flavonols and 3 resveratrols. The contents of non-anthocyanin phenolic compounds kept increasing before veraison, and decreased afterwards until 100% colored, thereafter the contents of non-anthocyanin phenolic compounds tended to increase again until maturity. The cinnamic acids, resveratrols and flavonols played crucial roles in the composition of non-anthocyanin phenolic compounds in berry skins, according to principal component analysis. There were also 30 non-anthocyanin phenolic compounds in their wines, including 5 benzoic acids,7 cinnamic acids,5 flavan-3-ols,9 flavonols and 4 resveratrols. Principal component analysis showed that the composition of non-anthocyanin phenolic compounds depended on cinnamic acids,flavan-3-ols and benzoic acids.
The growth dynamics of'Shuanghong'can be fitted to a double-sigmoid curve. No anthocyanins were detected before veraison, but the anthocyanins content tended to increase in the wake of veraison and reached a peak at stage of maturity. The content of delphinidin, methoxy and monoglucoside of anthocyanins varied similar to that of anthocyanin. Benzoic acids were not detected during the 2nd and 6th week after flowering, but the contents increased from the 8th week until 100% colored, and then decreased until maturity. The contents of cinnamic acids and flavan-3-ols showed a tendency to decreasing during berry development. Resveratrols were detected only after veraison, and the contents tended to increase until maturity.
The expression of structural gene in relation to biosynthesis of phenolic matters in 'Shuanghong' varied with the berry development. The genes of PAL, CHI1,F3H2, DFR,and LDOX were up-regulated from the 2nd to 4th week after flowering, down-regulated from the 6th week to veraison, and then up-regulated from veraison to maturity. The genetic transcriptional level had a positive correlation with the content of anthocyanins from veraison to maturity stage.The CHS3,RS, UFGT, GST, and OMT were down-regulated from the 2nd week during flowering and veraison stage, and then up-regulated from veraison to maturity. The genetic transcriptional level also showed a positive correlation with the content of anthocyanins from veraison to maturity stage. This positive correlation was also abserved between the RS transcriptional level and the content of resveratrols, and between OMT transcriptional level and the content of methoxyl-anthocyanins as well. F3'H expression was up-regulated from the 2nd to 4th week after flowering, down-regulated during the 6th to 8th week, and then up-regulated again from the 8th week to maturity. F3'H transcriptional level was correlated positively with the content of cyanidin anthocyanins from veraison to maturity. F3'5'H expression was down-regulated from the 2nd week after flowering to veraison stage, and then up-regulated after veraison. F3'5'H transcriptional level was correlated positively with the content of delphinidin anthocyanins from veraison to maturity. ANR expression was up-regulated from the 2nd week after flowering until the veraison stage, and then down-regulated after veraison, its transcriptional level was positively related to the content of epicatechin from the 2nd week after flowering until maturity. LAR2 expression was up-regulated from the 2nd week after flowering until the veraison stage, and then down-regulated thereafter, its transcriptional level had a positive relationship with the content of catechin since the 2nd week after flowering until maturity.
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