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

Abstract

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

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.

Figures/Tables 6

Table 1

Table 2

Analysis of monoterpenes content in grape berries cultivated in different trellis systems during maturity (μg?L-1)"

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	-

Table 2

Fig. 1

Fig. 2

Fig. 3

Fig. 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		-	-	-

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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