生物技术通报 ›› 2021, Vol. 37 ›› Issue (6): 24-35.doi: 10.13560/j.cnki.biotech.bull.1985.2020-1245
张谦1(), 薛宇2, 贺凌霄1, 吴疆3, 程玉渊3, 杨铁钊1, 丁永乐1, 徐世晓1, 薛刚1()
收稿日期:
2020-10-09
出版日期:
2021-06-26
发布日期:
2021-07-08
作者简介:
张谦,硕士研究生,研究方向:烟草品种筛选;E-mail: 基金资助:
ZHANG Qian1(), XUE Yu2, HE Ling-xiao1, WU Jiang3, CHENG Yu-yuan3, YANG Tie-zhao1, DING Yong-le1, XU Shi-xiao1, XUE Gang1()
Received:
2020-10-09
Published:
2021-06-26
Online:
2021-07-08
摘要:
烟叶中类西柏烷类及其降解产物和石油醚提取物是烟草中重要的香味物质。本研究探究种植密度、施氮量和留叶数三种栽培措施对烟叶类西柏烷类化合物和石油醚提取物的影响,为进一步彰显烟叶浓香型风格提供有效方法。以烤烟品种“渠首1号”为试材,按L9(34)正交试验设计,将上述三因素各设3个不同水平进行研究。观察不同处理的腺毛形态和密度,测定不同处理的叶面腺毛分泌物,并利用荧光定量PCR 技术比较不同处理中与西柏三烯二醇代谢相关CYC-1、CYP71D16等基因的表达量,对比了不同处理初烤烟叶茄酮含量和石油醚提取物含量,并进行了感官质量评价。种植密度是影响腺毛总密度、叶面烷烃类含量以及烤后烟石油醚提取物的关键因子;留叶数对烟叶茄酮含量的影响最大;施氮量是影响西柏三烯一醇、西柏三烯二醇、叶面分泌物总量,CYC-1和CYP71D16的相对表达量以及感官质量评价的关键因子。适宜的栽培措施能促进西柏三烯二醇代谢相关基因的上调表达,改良叶面分泌物,提高石油醚提取物和茄酮含量,从而改善感官质量,进一步彰显烟叶的香气风格特征。
张谦, 薛宇, 贺凌霄, 吴疆, 程玉渊, 杨铁钊, 丁永乐, 徐世晓, 薛刚. 密度、施氮量和留叶数对渠首1号香气特性彰显的影响[J]. 生物技术通报, 2021, 37(6): 24-35.
ZHANG Qian, XUE Yu, HE Ling-xiao, WU Jiang, CHENG Yu-yuan, YANG Tie-zhao, DING Yong-le, XU Shi-xiao, XUE Gang. Effects of Planting Density,Amount of Nitrogen Application and Left Leaf Number on the Highlighting of Flavor Style in Qushou 1[J]. Biotechnology Bulletin, 2021, 37(6): 24-35.
编码值 Code value | 种植密度(M) Planting density/(plants·hm-2) | 施氮量(N) Nitrogen application rate/(kg·hm-2) | 留叶数(L) Number of leaves left/(leaves/plant) |
---|---|---|---|
1 | 14 000 | 30 | 18 |
2 | 15 500 | 45 | 20 |
3 | 17 000 | 60 | 22 |
变化区间 | 1 500 | 15 | 2 |
表1 正交试验因素水平表
Table 1 Orthogonal test factor level table
编码值 Code value | 种植密度(M) Planting density/(plants·hm-2) | 施氮量(N) Nitrogen application rate/(kg·hm-2) | 留叶数(L) Number of leaves left/(leaves/plant) |
---|---|---|---|
1 | 14 000 | 30 | 18 |
2 | 15 500 | 45 | 20 |
3 | 17 000 | 60 | 22 |
变化区间 | 1 500 | 15 | 2 |
处理组合 Treatment combination | 处理 Treatment | 种植密度(M) Planting density/(plants·hm-2) | 施氮量(N) Nitrogen application rate/(kg·hm-2) | 留叶数(L) Number of leaves left/(leaves/plant) |
---|---|---|---|---|
T1 | M1N1L1 | 14 000 | 30 | 18 |
T2 | M1N2L2 | 14 000 | 45 | 20 |
T3 | M1N3L3 | 14 000 | 60 | 22 |
T4 | M2N1L2 | 15 500 | 30 | 20 |
T5 | M2N2L3 | 15 500 | 45 | 22 |
T6 | M2N3L1 | 15 500 | 60 | 18 |
T7 | M3N1L3 | 17 000 | 30 | 22 |
T8 | M3N2L1 | 17 000 | 45 | 18 |
T9 | M3N3L2 | 17 000 | 60 | 20 |
表2 试验安排表
Table 2 Test schedule
处理组合 Treatment combination | 处理 Treatment | 种植密度(M) Planting density/(plants·hm-2) | 施氮量(N) Nitrogen application rate/(kg·hm-2) | 留叶数(L) Number of leaves left/(leaves/plant) |
---|---|---|---|---|
T1 | M1N1L1 | 14 000 | 30 | 18 |
T2 | M1N2L2 | 14 000 | 45 | 20 |
T3 | M1N3L3 | 14 000 | 60 | 22 |
T4 | M2N1L2 | 15 500 | 30 | 20 |
T5 | M2N2L3 | 15 500 | 45 | 22 |
T6 | M2N3L1 | 15 500 | 60 | 18 |
T7 | M3N1L3 | 17 000 | 30 | 22 |
T8 | M3N2L1 | 17 000 | 45 | 18 |
T9 | M3N3L2 | 17 000 | 60 | 20 |
名称 Name | GenBank accession | 引物 Primer(5'-3') |
---|---|---|
Actin | AB158612 | F:GCTGTTTTTCCTAGTATTGTTGGTC |
R:AATACCTGTTGTACGGCCACTG | ||
CYC-1 | AY049090 | F:CGAGGGTGGGATCAAGGAAATACA |
R:ATCGCATCCGTGAAAAGCATAAG | ||
CYP17D16 | AF166332 | F:ATGGACCAATTATGCACCTTCAACT |
R:GGCACTGAGCAATTCCAAGAGACA |
表3 各基因名称及引物
Table 3 Gene names and primers
名称 Name | GenBank accession | 引物 Primer(5'-3') |
---|---|---|
Actin | AB158612 | F:GCTGTTTTTCCTAGTATTGTTGGTC |
R:AATACCTGTTGTACGGCCACTG | ||
CYC-1 | AY049090 | F:CGAGGGTGGGATCAAGGAAATACA |
R:ATCGCATCCGTGAAAAGCATAAG | ||
CYP17D16 | AF166332 | F:ATGGACCAATTATGCACCTTCAACT |
R:GGCACTGAGCAATTCCAAGAGACA |
处理组合 Treatment combination | T1 | T2 | T3 | T4 | T5 | T6 | T7 | T8 | T9 |
---|---|---|---|---|---|---|---|---|---|
短柄腺毛 Glandular hairs with short stalks | 9.00de | 11.21ab | 10.06bcd | 11.67a | 10.82abc | 9.51cde | 8.22e | 8.62e | 8.15e |
长柄腺毛 Glandular hairs with long stalks | 17.38de | 19.38bc | 18.32cd | 21.63a | 18.75c | 16.56e | 20.09b | 17.44de | 16.59e |
腺毛总密度 Total density of glandular hairs | 26.38de | 30.59b | 28.38c | 33.30a | 29.57bc | 26.07de | 27.98cd | 26.06de | 24.74e |
表4 各处理组合间烟叶腺毛密度差异分析
Table 4 Analysis of the difference in density of tobacco leaf glandular hair among the treatment combinations(root/mm2)
处理组合 Treatment combination | T1 | T2 | T3 | T4 | T5 | T6 | T7 | T8 | T9 |
---|---|---|---|---|---|---|---|---|---|
短柄腺毛 Glandular hairs with short stalks | 9.00de | 11.21ab | 10.06bcd | 11.67a | 10.82abc | 9.51cde | 8.22e | 8.62e | 8.15e |
长柄腺毛 Glandular hairs with long stalks | 17.38de | 19.38bc | 18.32cd | 21.63a | 18.75c | 16.56e | 20.09b | 17.44de | 16.59e |
腺毛总密度 Total density of glandular hairs | 26.38de | 30.59b | 28.38c | 33.30a | 29.57bc | 26.07de | 27.98cd | 26.06de | 24.74e |
试验因子 Test factor | 水平 Level | 短柄腺毛密度 Density of glandular hair with short stalk | 长柄腺毛密度 Density of glandular hair with long stalk | 总腺毛密度 Total glandular hair density |
---|---|---|---|---|
M | M1 | 10.09a | 18.36a | 28.45b |
M2 | 10.67a | 18.98ab | 29.65a | |
M3 | 8.33b | 18.04b | 26.26c | |
Rm | 2.3367 | 0.9433 | 3.3911 | |
N | N1 | 9.63ab | 19.70a | 29.22a |
N2 | 10.22a | 18.52b | 28.74a | |
N3 | 9.24b | 17.16c | 26.4b | |
Rn | 0.9778 | 2.5422 | 2.8244 | |
L | L1 | 9.04b | 17.13b | 26.17b |
L2 | 10.34a | 19.20a | 29.54a | |
L3 | 9.7ab | 19.05a | 28.64a | |
Rl | 1.2978 | 2.0733 | 3.3711 |
表5 叶面腺毛密度的边际值和极差
Table 5 The marginal value and range of leaf glandular hair density
试验因子 Test factor | 水平 Level | 短柄腺毛密度 Density of glandular hair with short stalk | 长柄腺毛密度 Density of glandular hair with long stalk | 总腺毛密度 Total glandular hair density |
---|---|---|---|---|
M | M1 | 10.09a | 18.36a | 28.45b |
M2 | 10.67a | 18.98ab | 29.65a | |
M3 | 8.33b | 18.04b | 26.26c | |
Rm | 2.3367 | 0.9433 | 3.3911 | |
N | N1 | 9.63ab | 19.70a | 29.22a |
N2 | 10.22a | 18.52b | 28.74a | |
N3 | 9.24b | 17.16c | 26.4b | |
Rn | 0.9778 | 2.5422 | 2.8244 | |
L | L1 | 9.04b | 17.13b | 26.17b |
L2 | 10.34a | 19.20a | 29.54a | |
L3 | 9.7ab | 19.05a | 28.64a | |
Rl | 1.2978 | 2.0733 | 3.3711 |
处理组合 Treatment combination | 西柏三烯一醇含量 Content of cephtriene monoalcohol | 西柏三烯二醇含量 Contentof of cephtriene diols | 烷烃类 Alkanes | 腺毛分泌物总量 Total secretion of glandular hair |
---|---|---|---|---|
T1 | 0.1833bc | 46.9267cd | 9.7933c | 65.23d |
T2 | 0.1833bc | 50.8767b | 9.66c | 70.6872b |
T3 | 0.2167b | 44.54de | 8.9933de | 64.033e |
T4 | 0.29a | 54.21a | 11.0667b | 74.382a |
T5 | 0.2133b | 49.5033bc | 11.2467a | 68.8589c |
T6 | 0.1267c | 42.06ef | 9.0633d | 61.7715f |
T7 | 0.1833bc | 49.0667bc | 9.8267c | 71.0882b |
T8 | 0.1933bc | 45.3367d | 8.8567e | 64.255e |
T9 | 0.1267c | 41.2567f | 8.4367f | 61.1233f |
表6 各处理组合的烟叶腺毛分泌物含量差异分析
Table 6 Analysis of the difference in the secretion content of the tobacco leaf glandular hairs of each treatment combination(μg/cm2)
处理组合 Treatment combination | 西柏三烯一醇含量 Content of cephtriene monoalcohol | 西柏三烯二醇含量 Contentof of cephtriene diols | 烷烃类 Alkanes | 腺毛分泌物总量 Total secretion of glandular hair |
---|---|---|---|---|
T1 | 0.1833bc | 46.9267cd | 9.7933c | 65.23d |
T2 | 0.1833bc | 50.8767b | 9.66c | 70.6872b |
T3 | 0.2167b | 44.54de | 8.9933de | 64.033e |
T4 | 0.29a | 54.21a | 11.0667b | 74.382a |
T5 | 0.2133b | 49.5033bc | 11.2467a | 68.8589c |
T6 | 0.1267c | 42.06ef | 9.0633d | 61.7715f |
T7 | 0.1833bc | 49.0667bc | 9.8267c | 71.0882b |
T8 | 0.1933bc | 45.3367d | 8.8567e | 64.255e |
T9 | 0.1267c | 41.2567f | 8.4367f | 61.1233f |
试验因子 Test factor | 水平 Level | 西柏三烯一醇总和 Sum of cephtriene monoalcohol | 西柏三烯二醇总和 Sum of cephtriene diols | 烷烃类 Alkanes | 腺毛分泌物总和 Total secretion of glandular hair |
---|---|---|---|---|---|
M | M1 | 0.1944a | 47.4478a | 9.4822b | 66.6501b |
M2 | 0.21a | 48.5911a | 10.4589a | 68.3375a | |
M3 | 0.1678a | 45.22b | 9.04c | 65.4888c | |
Rm | 0.0422 | 3.3711 | 1.4189 | 2.8486 | |
N | N1 | 0.2189a | 50.0678a | 10.2289a | 70.2334a |
N2 | 0.1967ab | 48.5722a | 9.9211b | 67.9337b | |
N3 | 0.1567b | 42.6189b | 8.8311c | 62.3092c | |
Rn | 0.0622 | 7.4489 | 1.3978 | 7.9242 | |
L | L1 | 0.1678a | 44.7744b | 9.2378c | 63.7522c |
L2 | 0.2a | 48.7811a | 9.7211b | 68.7308a | |
L3 | 0.2044a | 47.7033a | 10.0222a | 67.9934b | |
Rl | 0.0367 | 4.0067 | 0.7844 | 4.9787 |
表7 腺毛分泌物含量的边际值和极差
Table 7 Marginal value and range of glandular hair secretion content(μg/cm2)
试验因子 Test factor | 水平 Level | 西柏三烯一醇总和 Sum of cephtriene monoalcohol | 西柏三烯二醇总和 Sum of cephtriene diols | 烷烃类 Alkanes | 腺毛分泌物总和 Total secretion of glandular hair |
---|---|---|---|---|---|
M | M1 | 0.1944a | 47.4478a | 9.4822b | 66.6501b |
M2 | 0.21a | 48.5911a | 10.4589a | 68.3375a | |
M3 | 0.1678a | 45.22b | 9.04c | 65.4888c | |
Rm | 0.0422 | 3.3711 | 1.4189 | 2.8486 | |
N | N1 | 0.2189a | 50.0678a | 10.2289a | 70.2334a |
N2 | 0.1967ab | 48.5722a | 9.9211b | 67.9337b | |
N3 | 0.1567b | 42.6189b | 8.8311c | 62.3092c | |
Rn | 0.0622 | 7.4489 | 1.3978 | 7.9242 | |
L | L1 | 0.1678a | 44.7744b | 9.2378c | 63.7522c |
L2 | 0.2a | 48.7811a | 9.7211b | 68.7308a | |
L3 | 0.2044a | 47.7033a | 10.0222a | 67.9934b | |
Rl | 0.0367 | 4.0067 | 0.7844 | 4.9787 |
试验因子 Test factor | M | N | L | ||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
M1 | M2 | M3 | Rm | N1 | N2 | N3 | Rn | L1 | L2 | L3 | Rl | ||||
CYC-1 | 1.0767b | 1.1811a | 0.9289c | 0.2522 | 1.1956a | 1.2444a | 0.7467b | 0.4978 | 0.9089b | 1.1289a | 1.1489a | 0.24 | |||
CYP71D16 | 1.0067ab | 1.0356a | 0.9522b | 0.0833 | 1.1789a | 1.0644b | 0.7511c | 0.4278 | 0.8167b | 1.1056a | 1.0722a | 0.2889 |
表8 CYC-1和CYP71D16基因相对表达量的边际值和极差
Table 8 The marginal value and range of the relative expression of CYC-1 and CYP71D16 genes
试验因子 Test factor | M | N | L | ||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
M1 | M2 | M3 | Rm | N1 | N2 | N3 | Rn | L1 | L2 | L3 | Rl | ||||
CYC-1 | 1.0767b | 1.1811a | 0.9289c | 0.2522 | 1.1956a | 1.2444a | 0.7467b | 0.4978 | 0.9089b | 1.1289a | 1.1489a | 0.24 | |||
CYP71D16 | 1.0067ab | 1.0356a | 0.9522b | 0.0833 | 1.1789a | 1.0644b | 0.7511c | 0.4278 | 0.8167b | 1.1056a | 1.0722a | 0.2889 |
试验因子Test factors | M | N | L | ||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
M1 | M2 | M3 | Rm | N1 | N2 | N3 | Rn | L1 | L2 | L3 | Rl | ||||
茄酮含量Solanone content | 58.83b | 65.92a | 56.63c | 9.2878 | 63.63b | 67.91a | 49.84c | 18.0678 | 51.93c | 61.72b | 67.73a | 15.8067 |
表9 不同因素水平下烟叶茄酮的边际值和极差
Table 9 Marginal value and range of solanone in tobacco leaves under different factor levels(μg/g)
试验因子Test factors | M | N | L | ||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
M1 | M2 | M3 | Rm | N1 | N2 | N3 | Rn | L1 | L2 | L3 | Rl | ||||
茄酮含量Solanone content | 58.83b | 65.92a | 56.63c | 9.2878 | 63.63b | 67.91a | 49.84c | 18.0678 | 51.93c | 61.72b | 67.73a | 15.8067 |
图3 各处理组合间烤后烟石油醚提取物质量分数差异分析
Fig.3 Analysis of the difference in mass fraction of petrol-eum ether extract of flue-cured tobacco among vari-ous treatment combinations
试验因子Test factors | M | N | L | ||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
M1 | M2 | M3 | Rm | N1 | N2 | N3 | Rn | L1 | L2 | L3 | Rl | ||||
石油醚提取物 | 5.60c | 7.44a | 6.09b | 1.8444 | 5.84c | 7.03a | 6.26b | 1.1889 | 6.33b | 6.03c | 6.77a | 0.7333 |
表10 不同因素烤后烟石油醚的边际值和极差
Table 10 Marginal value and range of petroleum ether of flue-cured tobacco after different factors
试验因子Test factors | M | N | L | ||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
M1 | M2 | M3 | Rm | N1 | N2 | N3 | Rn | L1 | L2 | L3 | Rl | ||||
石油醚提取物 | 5.60c | 7.44a | 6.09b | 1.8444 | 5.84c | 7.03a | 6.26b | 1.1889 | 6.33b | 6.03c | 6.77a | 0.7333 |
处理组合Treatment combination | 香气特征Aroma character | 烟气特征Smoke characteristics | 吃味特征Taste characteristics | 总计 Total | ||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
香气质 Aroma quality | 香气量 Aroma quantity | 透发性 Permeabilities | 浓度 Concentration | 细腻程度 Fineness | 柔细度 Flexibility | 圆润感 Mellow feeling | 劲头 Energy | 刺激性 Thrill | 杂气 Miscellaneous gas | 余味 Aftertaste | ||||
T1 | 6 | 6.5 | 5.5 | 6 | 5 | 4.5 | 5 | 5 | 5 | 6 | 5 | 64.1 | ||
T2 | 4.5 | 5.5 | 5 | 5.5 | 4.5 | 4 | 4.5 | 6 | 5 | 6.5 | 6.5 | 59.7 | ||
T3 | 5.5 | 5.5 | 5 | 5.5 | 4.5 | 5 | 4 | 6 | 6.5 | 5 | 6 | 60.4 | ||
T4 | 6.5 | 6.5 | 5.5 | 6 | 4.5 | 4.5 | 4.5 | 5.5 | 6 | 6.5 | 5 | 65.5 | ||
T5 | 6.5 | 6.5 | 6.5 | 7 | 5 | 5.5 | 5.5 | 6 | 6 | 6.5 | 6 | 69.4 | ||
T6 | 5 | 5 | 5.5 | 6 | 4.5 | 4.5 | 4.5 | 5 | 5 | 5 | 5.5 | 57.2 | ||
T7 | 5.5 | 5 | 6 | 6.5 | 5 | 4.5 | 5 | 5 | 6 | 6.5 | 6 | 61.7 | ||
T8 | 6.5 | 6.5 | 6 | 6.5 | 5 | 5 | 5 | 5 | 5 | 6 | 5.5 | 67.0 | ||
T9 | 5.5 | 5.5 | 5 | 5.5 | 4.5 | 4.5 | 4.5 | 5 | 5 | 6 | 5.5 | 60.2 |
表11 各处理组合间烤后烟感官质量评价
Table 11 Sensory quality evaluation of flue-cured tobacco in each treatment combination
处理组合Treatment combination | 香气特征Aroma character | 烟气特征Smoke characteristics | 吃味特征Taste characteristics | 总计 Total | ||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
香气质 Aroma quality | 香气量 Aroma quantity | 透发性 Permeabilities | 浓度 Concentration | 细腻程度 Fineness | 柔细度 Flexibility | 圆润感 Mellow feeling | 劲头 Energy | 刺激性 Thrill | 杂气 Miscellaneous gas | 余味 Aftertaste | ||||
T1 | 6 | 6.5 | 5.5 | 6 | 5 | 4.5 | 5 | 5 | 5 | 6 | 5 | 64.1 | ||
T2 | 4.5 | 5.5 | 5 | 5.5 | 4.5 | 4 | 4.5 | 6 | 5 | 6.5 | 6.5 | 59.7 | ||
T3 | 5.5 | 5.5 | 5 | 5.5 | 4.5 | 5 | 4 | 6 | 6.5 | 5 | 6 | 60.4 | ||
T4 | 6.5 | 6.5 | 5.5 | 6 | 4.5 | 4.5 | 4.5 | 5.5 | 6 | 6.5 | 5 | 65.5 | ||
T5 | 6.5 | 6.5 | 6.5 | 7 | 5 | 5.5 | 5.5 | 6 | 6 | 6.5 | 6 | 69.4 | ||
T6 | 5 | 5 | 5.5 | 6 | 4.5 | 4.5 | 4.5 | 5 | 5 | 5 | 5.5 | 57.2 | ||
T7 | 5.5 | 5 | 6 | 6.5 | 5 | 4.5 | 5 | 5 | 6 | 6.5 | 6 | 61.7 | ||
T8 | 6.5 | 6.5 | 6 | 6.5 | 5 | 5 | 5 | 5 | 5 | 6 | 5.5 | 67.0 | ||
T9 | 5.5 | 5.5 | 5 | 5.5 | 4.5 | 4.5 | 4.5 | 5 | 5 | 6 | 5.5 | 60.2 |
试验因子Test factors | M | N | L | ||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
M1 | M2 | M3 | Rm | N1 | N2 | N3 | Rn | L1 | L2 | L3 | Rl | ||||
评吸总分Total score | 61.4 | 64.03 | 62.97 | 2.63 | 63.77 | 65.37 | 59.27 | 6.1 | 62.77 | 61.8 | 63.83 | 2.03 |
表12 不同因素烤后烟感官质量评价边际值和极差
Table 12 Marginal value and range of sensory quality evaluation of flue-cured tobacco after different factors
试验因子Test factors | M | N | L | ||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
M1 | M2 | M3 | Rm | N1 | N2 | N3 | Rn | L1 | L2 | L3 | Rl | ||||
评吸总分Total score | 61.4 | 64.03 | 62.97 | 2.63 | 63.77 | 65.37 | 59.27 | 6.1 | 62.77 | 61.8 | 63.83 | 2.03 |
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