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

doi:10.3864/j.issn.0578-1752.2019.07.002

中国农业科学 ›› 2019, Vol. 52 ›› Issue (7): 1136-1149.doi: 10.3864/j.issn.0578-1752.2019.07.002

所属专题：葡萄研究专题

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

王慧玲¹,王晓玥²,闫爱玲³,孙磊¹,张国军¹,任建成¹,徐海英¹()

1 北京市林业果树科学研究院,北京 100093
2 北京市落叶果树工程技术研究中心,北京 100097
3 农业部华北地区园艺作物生物学与种质创制重点实验室,北京 100097

收稿日期:2018-09-14 接受日期:2019-01-11 出版日期:2019-04-01 发布日期:2019-04-04
通讯作者: 徐海英
作者简介:王慧玲,E-mail： wanghui198216@126.com。
基金资助:
国家自然科学基金青年基金(31601712);现代农业产业技术体系建设专项(CARS-29);北京市农林科学院青年科学基金(QNJJ201604);北京市自然科学基金(6182007)

The Accumulation of Monoterpenes and the Expression of Its Biosynthesis Related Genes in ‘Aishen Meigui’ Grape Berries Cultivated in Different Trellis Systems During Ripening Stage

WANG HuiLing¹,WANG XiaoYue²,YAN AiLing³,SUN Lei¹,ZHANG GuoJun¹,REN JianCheng¹,XU HaiYing¹()

1 Beijing Academy of Forestry and Pomology Sciences, Beijing 100093
2 Beijing Engineering Research Center for Deciduous Fruit Trees, Beijing 100097
3 Key Laboratory of Biology and Genetic Improvement of Horticultural Crops (North China), Ministry of Agriculture, Beijing 100097

Received:2018-09-14 Accepted:2019-01-11 Online:2019-04-01 Published:2019-04-04
Contact: HaiYing XU

摘要/Abstract

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108.18 μg?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403.24 μg?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。

关键词: 葡萄, 栽培架式, 玫瑰香味, 单萜

Abstract:

【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-xylulose 5-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.

Key words: grape, trellis systems, muscat flavor, monoterpenes

王慧玲,王晓玥,闫爱玲,孙磊,张国军,任建成,徐海英. 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[J]. 中国农业科学, 2019, 52(7): 1136-1149.

WANG HuiLing,WANG XiaoYue,YAN AiLing,SUN Lei,ZHANG GuoJun,REN JianCheng,XU HaiYing. The Accumulation of Monoterpenes and the Expression of Its Biosynthesis Related Genes in ‘Aishen Meigui’ Grape Berries Cultivated in Different Trellis Systems During Ripening Stage[J]. Scientia Agricultura Sinica, 2019, 52(7): 1136-1149.

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链接本文: https://www.chinaagrisci.com/CN/10.3864/j.issn.0578-1752.2019.07.002

https://www.chinaagrisci.com/CN/Y2019/V52/I7/1136

图/表 6

表1

表2

不同架式葡萄果实成熟期间单萜含量分析"

化合物 Compound	年份 Year	架式Trellis	盛花后天数 Days after full blossom (d)
化合物 Compound	年份 Year	架式Trellis	30	39	48	57	67	76
β-月桂烯 β-Myrcene	2016	T V	0.25±0.02a 0.24±0.03a	0.86±0.06b 4.71±0.06a	0.22±0.01b 0.34±0.02a	1.28±0.09a 0.26±0.02b	1.32±0.12a 0.44±0.07b	0.45±0.04a 0.16±0.00b
	2017	T V	0.95±0.01b	1.02±0.01b	1.99±0.10b	10.30±0.31a	7.21±0.06a	-
	2017	T V	1.17±0.40a	1.24±0.34a	3.35±0.11a	1.56±0.09b	1.17±0.02b	-
柠檬烯 Limonene	2016	T V	8.30±0.04a 3.69±0.11b	7.31±0.00a 3.71±0.11b	1.86±0.01b 5.86±1.04a	9.08±0.15a 2.33±0.12b	11.03±0.19a 1.68±0.08b	7.73±0.14a 2.56±0.07b
	2017	T V	5.16±0.10a	5.08±0.12a	5.42±0.04b	17.57±0.31a	16.98±0.32a	-
	2017	T V	3.43±0.12b	5.20±0.09a	9.18±0.51a	6.50±0.20b	3.35±0.07b	-
水芹烯 Phellandrene	2016	T V	1.95±0.06a 0.87±0.02b	2.07±0.01a 0.77±0.02b	0.56±0.10b 1.11±0.07a	1.08±0.11a 0.53±0.01b	1.91±0.07a 0.47±0.08b	1.49±0.05a 0.56±0.03b
	2017	T V	1.72±0.01a	3.40±0.01a	0.74±0.02b	1.56±0.05a	1.82±0.02a	-
	2017	T V	0.79±0.01b	0.73±0.08b	1.10±0.06a	0.73±0.01b	0.56±0.02b	-
β-trans-罗勒烯 β-trans-Ocimene	2016	T V	3.08±0.00a 1.17±0.08b	2.61±0.03a 1.21±0.01b	0.71±0.00b 1.96±0.34a	3.68±0.02a 0.83±0.10b	4.41±0.03a 0.57±0.05b	3.30±0.03a 1.00±0.03b
	2017	T V	3.07±0.01a	2.96±0.03a	3.99±0.22b	9.24±0.26a	8.89±0.06a	-
	2017	T V	2.73±0.04b	2.97±0.41a	6.15±0.35a	3.07±0.06b	2.64±0.12b	-
γ-松油烯 γ-Terpinen	2016	T V	5.26±0.02a 3.10±0.16b	3.84±0.10a 1.41±0.08b	0.91±0.03a 0.87±0.15b	1.34±0.03a 0.30±0.01b	1.93±0.05a 0.39±0.18b	0.87±0.03a 0.32±0.01b
	2017	T V	5.79±0.61a	3.74±0.04a	2.17±0.07a	2.06±0.40a	1.56±0.07a	-
	2017	T V	2.59±0.03b	2.19±0.21b	1.53±0.06b	1.14±0.07b	1.08±0.12b	-
β-cis-罗勒烯 β-cis-Ocimene	2016	T V	4.22±0.00a 2.52±0.17b	3.95±0.04a 2.60±0.04b	1.50±0.01b 3.60±0.34a	6.51±0.05a 1.86±0.12b	8.44±0.16a 1.14±0.09b	5.44±0.14a 2.18±0.00b
	2017	T V	6.36±0.21a	5.07±0.11a	3.54±0.01b	18.94±0.03a	19.06±0.15a	-
	2017	T V	3.81±0.02b	4.01±0.25b	8.60±0.31a	3.92±0.20b	0.98±0.31b	-
异松油烯 Terpinolen	2016	T V	5.15±0.00a 2.71±0.13b	4.57±0.00a 2.32±0.10b	1.06±0.04b 3.47±0.96a	4.94±0.05a 0.98±0.06b	5.50±0.06a 0.73±0.18b	4.50±0.04a 1.20±0.10b
	2017	T V	7.32±0.02a	6.67±0.30a	6.12±0.03b	7.31±0.41a	9.94±0.08a	-
	2017	T V	4.32±0.40b	4.33±0.13b	7.79±0.16a	3.12±0.04b	1.83±0.17b	-
cis-氧化玫瑰 cis Rose oxide	2016	T V	0.35±0.01b 0.80±0.02a	tr 0.25±0.01	tr 0.21±0.02	0.30±0.01b 0.46±0.00a	0.23±0.00 tr	tr tr
	2017	T V	0.64±0.01a	0.68±0.01b	0.73±0.13b	0.85±0.02a	0.59±0.12a	-
	2017	T V	0.68±0.01a	0.82±0.01a	0.82±0.03a	0.70±0.03b	0.36±0.04b	-
trans-氧化玫瑰 trans-Rose oxide	2016	T V	tr tr	tr tr	tr tr	tr tr	tr tr	tr tr
	2017	T V	tr	tr	0.62±0.02a	2.93±0.07a	0.61±0.01	-
	2017	T V	tr	tr	0.64±0.01a	0.56±0.04b	tr	-
别罗勒烯 Allo-Ocimene	2016	T V	3.16±0.01a 1.12±0.06b	2.54±0.04a 1.10±0.02b	0.70±0.03b 1.85±0.33a	3.79±0.02a 0.87±0.01b	4.48±0.07a 0.55±0.03b	3.24±0.04a 0.89±0.01b
	2017	T V	2.98±0.01a	2.86±0.30a	2.86±0.06b	5.36±0.06a	6.12±0.09a	-
	2017	T V	2.62±0.04b	2.85±0.03a	3.66±0.16a	1.37±0.02b	0.73±0.02b	-
化合物 Compound	年份 Year	架式Trellis	盛花后天数 Days after full blossom (d)
化合物 Compound	年份 Year	架式Trellis	30	39	48	57	67	76
(E,Z)-别罗勒烯 (E,Z)-Allo- Ocimene	2016	T V	0.55±0.02a 0.37±0.01b	0.57±0.02a 0.36±0.01b	0.35±0.03b 0.58±0.08a	1.27±0.03a 0.48±0.12b	1.47±0.07a 0.26±0.03b	0.82±0.01a 0.32±0.01b
	2017	T V	0.62±0.10b	0.63±0.03a	0.74±0.04b	1.42±0.02a	1.82±0.08a	-
	2017	T V	0.69±0.01a	0.64±0.01a	0.81±0.09a	0.64±0.04b	0.41±0.02b	-
cis-呋喃型氧化里那醇 cis-furan linalool oxide	2016	T V	9.31±0.01a 2.50±0.04b	5.03±0.13a 2.15±0.14b	1.46±0.05b 2.66±0.00a	2.66±0.10a 1.32±0.05b	3.37±0.16a 1.09±0.03b	1.81±0.07a 1.18±0.00a
	2017	T V	4.92±0.07a	2.65±0.01a	1.23±0.31b	3.19±0.02a	4.00±0.22a	-
	2017	T V	1.43±0.28b	1.80±0.17b	2.71±0.50a	1.32±0.15b	0.30±0.07b	-
trans-呋喃型氧化里那醇 trans-furan linalool oxide	2016	T V	3.49±0.11a 1.73±0.03b	2.32±0.05a 1.52±0.03b	1.14±0.01b 1.47±0.03a	1.68±0.04a 1.19±0.02b	1.84±0.04a 1.05±0.01b	1.23±0.02a 1.01±0.00b
	2017	T V	1.76±0.04a	0.79±0.09b	0.35±0.08b	1.09±0.49a	1.83±0.07a	-
	2017	T V	1.41±0.05b	1.14±0.31a	0.58±0.12a	0.53±0.01b	0.23±0.04b	-
橙花醚 Nerol oxide	2016	T V	10.02±0.17a 5.92±0.05b	10.18±0.30a 8.98±0.55b	8.86±0.14a 3.09±2.40b	7.05±0.94b 11.79±0.26a	3.53±1.13a 3.38±0.60a	3.65±0.02a 2.31±0.07b
	2017	T V	25.16±0.09a	23.70±1.74a	17.78±0.62b	16.86±0.73a	10.41±0.16a	-
	2017	T V	14.15±0.53b	17.16±0.73b	21.77±0.22a	15.12±0.12b	8.97±0.21b	-
香茅醛 Citronellal	2016	T V	0.81±0.07a 0.42±0.05b	0.59±0.06a 0.50±0.01a	0.34±0.01b 0.65±0.04a	0.77±0.04a 0.24±0.10b	0.99±0.04a 0.10±0.01b	0.60±0.06a 0.28±0.04b
	2017	T V	-	-	-	-	-	-
	2017	T V	-	-	-	-	-	-
里那醇 Linalool	2016	T V	3.63±0.04b 5.58±0.04a	8.33±0.12a 6.79±0.35b	4.44±0.13b 14.62±0.77a	41.67±0.94a 10.48±0.32b	19.29±1.00a 4.23±0.55b	33.60±0.70a 11.12±0.26b
	2017	T V	8.38±1.12a	15.22±0.15b	82.89±3.07a	96.64±2.25a	43.56±0.75a	-
	2017	T V	7.01±0.05b	15.26±0.37a	71.08±1.03b	67.56±0.54b	8.90±0.43b	-
4-松油烯醇 4-Terpineol	2016	T V	2.95±0.01a 0.88±0.03b	2.72±0.00a 0.52±0.05b	0.39±0.03b 0.47±0.01a	0.81±0.06a 0.22±0.01b	1.24±0.02b 3.08±0.23a	0.51±0.01a 0.16±0.01b
	2017	T V	2.91±0.09a	3.23±0.04a	2.89±0.07a	2.56±0.01a	1.98±0.02a	-
	2017	T V	0.90±0.02b	0.87±0.01b	0.83±0.02b	0.72±0.02b	0.70±0.02b	-
橙花醛 Neral	2016	T V	0.67±0.01a 0.27±0.01b	0.34±0.02a 0.33±0.05b	0.13±0.01b 0.60±0.05a	0.49±0.06b 0.59±0.04a	0.67±0.08a 0.14±0.03b	0.40±0.03a 0.15±0.08b
	2017	T V	0.87±0.03a	0.75±0.01a	0.63±0.01b	0.65±0.05b	0.84±0.01a	-
	2017	T V	0.53±0.01b	0.61±0.02b	1.01±0.10a	0.83±0.01a	0.62±0.03b	-
α-衣兰油烯 α-Muurolene	2016	T V	0.22±0.00a 0.10±0.00b	0.09±0.00a 0.07±0.00b	0.07±0.00a 0.07±0.02a	0.08±0.01b 0.09±0.01a	0.09±0.00a 0.08±0.00b	0.07±0.02b 0.08±0.01a
	2017	T	0.60±0.01a	0.54±0.01b	0.55±0.01b	0.58±0.02a	0.52±0.05b	-
		V	0.59±0.01a	0.59±0.01a	0.60±0.01a	0.58±0.02a	0.60±0.01a	-
α-萜品醇 α-Terpineol	2016	T V	5.55±0.11a 4.49±0.02b	5.28±0.01a 4.59±0.05b	4.34±0.01b 4.99±0.11a	6.53±0.19a 4.36±0.09b	7.96±0.15a 2.80±0.97b	5.78±0.13a 4.42±0.02b
	2017	T	5.40±0.04a	9.23±1.07a	6.65±0.57b	22.29±0.11a	19.89±0.62a	-
		V	3.24±0.23b	6.41±0.78b	14.57±0.57a	3.03±0.03b	1.81±0.14b	-
化合物 Compound	年份 Year	架式Trellis	盛花后天数 Days after full blossom (d)
化合物 Compound	年份 Year	架式Trellis	30	39	48	57	67	76
香叶醛 Geranial	2016	T V	0.22±0.06b 0.31±0.09a	0.44±0.02b 0.53±0.03a	0.92±0.04b 1.08±0.11a	2.03±0.85a 1.20±0.21b	2.68±0.05a 0.53±0.09b	0.73±0.02a 0.35±0.02b
	2017	T	0.63±0.01a	0.74±0.01a	0.87±0.02b	1.68±0.01a	2.13±0.13a	-
		V	0.62±0.02a	0.65±0.01b	0.89±0.01a	1.20±0.10b	0.63±0.02b	-
β-香茅醇 β-Citronellol	2016	T	1.00±0.05b 3.58±0.03a	3.61±0.02a 3.65±0.04a	3.59±0.03a 2.59±0.78b	3.60±0.05a 0.53±0.09b	2.54±0.76a 0.05±0.01b	0.61±0.03a 0.23±0.01b
		V	1.00±0.05b 3.58±0.03a	3.61±0.02a 3.65±0.04a	3.59±0.03a 2.59±0.78b	3.60±0.05a 0.53±0.09b	2.54±0.76a 0.05±0.01b	0.61±0.03a 0.23±0.01b
	2017	T	2.59±0.15a	4.54±0.06a	5.56±0.06a	7.58±0.01a	3.70±0.11a	-
		V	2.49±0.04b	3.51±0.02b	4.61±0.04b	2.40±0.01b	0.64±0.01b	-
γ-香叶醇 γ-Geraniol	2016	T V	0.15±0.00a 0.06±0.01b	0.14±0.03a 0.08±0.01b	0.10±0.02a 0.08±0.02b	tr tr	tr tr	tr tr
	2017	T	0.54±0.01a	0.52±0.03a	0.53±0.04	tr	tr	-
		V	0.20±0.03b	0.21±0.01b	tr	tr	tr	-
橙花醇 Nerol	2016	T V	0.15±0.01b 0.24±0.00a	0.32±0.02a 0.29±0.04b	0.27±0.01b 0.77±0.07a	1.77±0.13a 0.45±0.07b	3.58±0.01a 0.25±0.05b	1.06±0.08a 0.25±0.04b
	2017	T	2.09±0.14b	3.43±0.07b	21.47±2.78b	82.82±8.42a	65.23±0.75a	-
		V	2.89±1.17a	4.62±0.35a	38.38±3.60a	19.56±0.54b	8.99±2.07b	-
cis-异香叶醇 cis-Isogeraniol	2016	T V	0.17±0.16 tr	tr tr	tr tr	tr tr	tr tr	tr nd
	2017	T	tr	0.25±0.08	tr	tr	tr	-
		V	tr	tr	tr	tr	tr	-
trans-异香叶醇 trans-Isogeraniol	2016	T V	tr tr	tr tr	tr tr	tr tr	tr tr	nd nd
	2017	T	0.13±0.03	tr	tr	tr	tr	tr
		V	tr	tr	tr	tr	tr	tr
香叶醇 Geraniol	2016	T V	0.62±0.05b 1.06±0.01a	3.25±1.13b 4.76±0.01a	4.91±0.02a 4.92±0.02a	5.22±0.03a 1.33±0.20b	5.62±0.02a 0.67±0.10b	5.01±0.03a 1.07±0.04b
	2017	T	3.89±0.19b	12.56±0.04b	58.97±3.42a	81.60±4.67a	82.14±0.86a	-
		V	7.31±0.10a	20.40±0.55a	54.27±6.01b	21.46±3.07b	17.59±2.26b	-
香叶酸 Geranic acid	2016	T V	0.29±0.02a 0.17±0.02b	0.51±0.20a 0.33±0.04b	0.81±0.30a 0.51±0.16b	5.22±0.03a 1.33±0.20b	0.39±0.14 tr	0.24±0.01a 0.18±0.03b
	2017	T	0.12±0.01b	2.98±0.12b	6.12±0.22a	8.12±0.56a	1.99±0.12a	-
		V	0.23±0.02a	3.53±0.10a	5.96±0.16b	6.32±0.08b	1.49±0.04b	-
总量 Total	2016	T V	71.50±0.09a 43.87±0.89b	71.40±2.69a 53.52±1.32b	39.66±0.23b 58.38±8.38a	108.18±3.15a 43.12±1.11b	94.52±4.29a 23.61±3.55b	83.08±0.46a 31.97±0.63b
	2017	T	94.63±1.23a	113.21±3.42a	235.43±0.41b	403.24±2.54a	312.82±4.12a	-
		V	65.86±3.11b	100.46±4.10b	260.91±0.91a	163.97±0.87b	64.61±5.16b	-

表2

图1

图2

图3

图4

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盛花后天数 Days after full blossom (d)	2016年				2017年
	可溶性固形物 Soluble solids (%)		可滴定酸 Titratable acid (g·L^-1)		可溶性固形物 Soluble solids (%)		可滴定酸 Titratable acid (g·L^-1)
	T	V	T	V	T	V	T	V
30	10.70±1.20a	10.54±0.87a	6.90±0.23a	6.35±0.32b	10.31±0.61a	10.42±0.53a	6.79±0.12a	6.45±0.32a
39	14.01±0.21b	14.69±0.86a	6.18±0.10a	6.20±0.15a	13.71±0.12b	14.03±0.21a	6.05±0.15a	6.13±0.04a
48	14.78±0.34b	15.72±0.53a	5.31±0.41a	4.06±0.02b	15.22±0.48b	15.81±0.57a	4.34±0.09a	4.16±0.14b
57	15.38±0.38b	16.01±0.65a	3.39±0.13a	3.26±0.17a	16.03±0.86b	16.14±0.58a	3.34±0.21a	3.48±0.04a
67	16.12±1.02a	16.03±0.21b	3.20±0.20a	3.23±0.45a	16.24±0.49a	16.03±0.12b	3.09±0.13a	3.15±0.11a
76	16.67±0.36a	16.05±0.12b	3.24±0.04a	3.20±0.25a		-	-	-

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

The Accumulation of Monoterpenes and the Expression of Its Biosynthesis Related Genes in ‘Aishen Meigui’ Grape Berries Cultivated in Different Trellis Systems During Ripening Stage

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