香料烟腺毛和结晶细胞发育及其与环境关系研究
|
摘要
2001-2002年以云南和新疆大田试验和河南农业大学校内盆栽试验香料烟为材料，就香料烟腺毛和结晶细胞的发育方面作了一些研究，并分析了部分环境条件对腺毛密度和结晶细胞的影响。结果如下：
1 香料烟腺毛的种类及分布
香料烟腺毛主要包括长柄多细胞腺头腺毛、长柄单细胞腺头腺毛和短柄多细胞腺头腺毛三种。长柄分枝腺毛基本上很难观察到。随叶位升高，长柄、短柄腺毛密度及短柄腺毛比例均增加。长柄腺毛为叶片(尤其幼叶)上优势种类，且叶背面比例高于正面。
2 香料烟叶腺毛的形态发生
长柄腺毛和短柄腺毛最早产生于茎尖第一对叶原基高约100-200μm时，腺毛原初细胞大部分发育成长柄腺毛。长柄腺毛和短柄腺毛由原表皮细胞突起分化产生，经过不同的形态形成过程，逐渐形成。长柄腺毛柄较长，由2-5个柄细胞组成，腺头棒状，由1-8个分泌细胞构成。短柄腺毛柄较短，只有一个柄细胞，腺头球状，由8-16个分泌细胞组成。
3 香料烟叶片生长成熟过程中腺毛密度的变化规律
香料烟叶片腺毛密度随叶片的生长基本上呈逐渐下降趋势。腺毛密度在叶长1cm时最高，此时长柄腺毛密度远高于短柄腺毛密度，随后短柄腺毛产生速度稍大于长柄腺毛，其比例逐渐提高，至叶长约占定长1／2时，两种腺毛数目都达最高，但长柄腺毛仍占优势，以后腺毛数目逐渐下降。腺毛产生的关键时期在叶长约占定长1／2前，以后很少产生并逐渐脱落。香料烟叶成熟过程中腺毛密度的变化与叶片成熟度、叶位和腺毛类型有关。就腺毛总密度来看，香料烟叶从欠熟变为适熟过程中，上部叶与下部叶腺毛明显脱落，中部叶腺毛没有明显脱落；从适熟再到过熟，只有下部叶腺毛明显脱落。
香料烟腺毛和结晶细胞发育及其与环境关系研究
4香料烟腺毛的分泌活动规律
香料烟两种类型腺毛（长柄腺毛和短柄腺毛）分泌活动明显不同。香
料烟长柄腺毛分泌活动与叶片发育尤其是叶片的成熟密切相关，从欠熟叶
开始腺毛腺头细胞分泌活动开始增强，至叶片适熟时大部分腺毛达到分泌
高峰，腺头细胞着生大量胶状分泌物，叶片过熟时腺毛分泌活动减弱，腺
毛分泌物逐渐干缩。但少数腺毛分泌活动滞后，当叶片适熟时只有少量分
泌，在调制初期才大量分泌。短柄腺毛在叶片生长成熟直到调制陈化过程
中腺头细胞不会分泌像长柄腺毛一样的胶状分泌物。
5香料烟叶腺毛分泌物成分分析
经分析定性出香料烟叶15种表面化合物，这些化合物中属于腺毛分泌
物及其降解物的有5种，含量比较高的是西柏三烯二醇。
6环境对腺毛密度的影响
不同产区腺毛密度存在差异，云南香料烟长柄多细胞腺头腺毛密度、
短柄腺毛密度及腺毛总密度均高于新疆，尤其是长柄多细胞腺毛密度明显
较高。
栽培措施对香料烟腺毛密度有一定影响。肥料类型（化肥和有机肥）
对长柄多细胞腺毛密度影响较大，对长柄单细胞腺毛密度影响不大，对短
柄腺毛密度影响很小。肥料类型对上部叶和中部叶腺毛密度影响较大，对
下部叶腺毛密度影响较小。施用有机肥能明显增加上部叶长柄多细胞腺毛
密度及腺毛总密度，纯施化肥增加中部叶长柄多细胞腺毛密度。灌水次数
主要影响香料烟上部叶腺毛密度，不灌水处理上部叶腺毛密度明显较高，
灌水越多，上部叶腺毛密度越低。灌水次数对中部和下部叶腺毛密度影响
不大。
7香料烟结晶细胞的超微结构和晶体形态
2
香料烟腺毛和结晶细胞发育及其与环境关系研究
结晶细胞是一类特化的含晶细胞，细胞内充满晶体，质体等被挤到细
胞边缘。由扫描电镜观察到结晶体形状似角锥状。叶片成熟过程中结晶细
胞超微结构发生变化，欠熟叶结晶细胞胞基质、质体处于细胞边缘，质体
逐渐退化，但仍保持质体形态，胞基质清晰可见；适熟叶结晶细胞己看不
到质体，胞基质收缩；过熟叶结晶细胞胞基质干缩。
8香料烟结晶细胞的发育和分布
结晶细胞最早产生于叶长 Icm左右，发生于从下表皮往上数第 3层细
胞处。叶长2叶cm左右时结晶细胞含量迅速增加，之后随着叶长增加其含
量逐渐下降，并处于栅栏组织和海绵组织之间，呈不连续的条带状。结晶
细胞随着叶片的生长逐渐增大，叶片成熟时一些结晶细胞破裂，颗粒状结
晶物散出。结晶细胞在香料烟烟株根茎叶中均有分布，以叶中较多，并随
叶位的升高含量增加。
9香料烟叶结晶细胞部分矿质元素相对含量
香料烟叶结晶细胞与普通细胞相比有较高的钙含量，因此推测细胞内
结晶体可能是一种钙盐结晶。同时不排除结晶体可能含有尚未证实的有机
物成分。
11不同产区自然条件及栽培措施对香料烟结晶细胞含量的影响
初步试验结果表明新疆碱性土壤下生长的香料烟叶中结晶细胞的含量
稍高于云南酸性土壤生长的香料烟。雨水较少年份结晶细胞含量较高。灌
水对叶片结晶细胞含量影响较大，不灌水处理各部位叶片结晶细胞含量均
较高，灌水越多，结晶细胞含量越低。
This paper studied on the development of glandular trichomes and crystal idioblasts of oriental tobacco and the effect of environment on them in Yunnan and Xinjiang in addition to pot experiment in Henan Agriculture University during 2001-2002. The results were as follows:
The glandular trichomes of oriental tobacco was principally classified by multicell-headed long stalk trichomes,monocell-headed long stalk trichomes and multicell-headed short stalk trichomes .Branched trichomes was hardly observed. Density of short trichomes and long ones and their ratio raised with the increasing of stalk position.The long trichomes had absolute superiority on the leaves,especially on the yonger leaves. The percentage on the lower surface was higher than that on the upper surface.
The morphology development of trichomes of two types was studied. Initial cell of the trichomes generally developed into long ones. The primary long trichomes and short ones both issumed from the first and second leaf primordium about 100-200um.The differentiation cells bulging from protoderm developed into long trichomes or short ones in different mode of morphology formation. The long trichomes had longer stalk composed of 2-5 cells and one rod-liking gland head composed of l-8secretory cells. The short trichomes had shorter stalk composed of only one cell and one ball-liking gland head composed of 8-16secretory cells.
The trichome density was highest when the leaves was 1 centimetre and appeared to descend gradually during the growth of leaves. The number of trichomes came to maxium when the leaves length was half of mature leaves,then decreased. The density and number of the long trichomes prevailed over short ones during the growth of the leaves though the forming velocity of short trichomes exceeded that of long ones. The trichome density of the leaves of three maturity varied by the leaf position. The trichome density varied by maturity ,leaf position and trichome types during leaf maturing. When the immature leaves turned into proper mature, the total trichome density on the uppeer and lower leaves decreased with obvious variation while no variation on the middle leaves. When the leaves appeared over mature ,only the total trichome density on the lower leaves obviously decreased.
The secretion activities of two trichomes was observed. The long trichomes were
closely related with the stage of leaf development .especially the mature stage.The secretion of gland cells grandually strengthened in the ver-matured leaves.and came to climax when the leaves were proper mature,then weakened when the leaves appeared over mature. A minority of trichomes delayed and had a little of gummy exudate in the fine-matured leaves,consequently secreting much during curing of the leaves.The short trichomes were obviously different from the long ones.The exudate observed in the long trichomes was hardly observed in the short trichomes during the growth ,mature and aging of the leaves.
The cuticular constituents about 15 were analyzed by GC/MS,5 compounds were belonged to secretory products of the glandular trichomes and their degradations.The content of cembratriendiol was higher.
The trichome density varied in different regions.Compared with Xinjiang,there were higher densities of multicell-headed long stalk trichomes,multicell-headed short stalk trichomes and total trichomes in Yunnan,and multicell-headed long stalk trichomes were especially significant.
The effect of different-typed fertilizers on trichome density varied by trichome category and leaf position. The multicell-headed long stalk trichomes were obviously influenced,The monocell-headed long stalk trichomes were affected any the less,however, the short trichomes were hardly influenced.The trichome density in the upper and middle leaves were more influenced than that in the lower leaves.The application of organic fertilizers enhanced the denity of multicell-headed long stalk trichomes and total trichomes of the upper leaves while chemical fertilizers increased the denity of multicell-heade
1 香料烟腺毛的种类及分布
香料烟腺毛主要包括长柄多细胞腺头腺毛、长柄单细胞腺头腺毛和短柄多细胞腺头腺毛三种。长柄分枝腺毛基本上很难观察到。随叶位升高，长柄、短柄腺毛密度及短柄腺毛比例均增加。长柄腺毛为叶片(尤其幼叶)上优势种类，且叶背面比例高于正面。
2 香料烟叶腺毛的形态发生
长柄腺毛和短柄腺毛最早产生于茎尖第一对叶原基高约100-200μm时，腺毛原初细胞大部分发育成长柄腺毛。长柄腺毛和短柄腺毛由原表皮细胞突起分化产生，经过不同的形态形成过程，逐渐形成。长柄腺毛柄较长，由2-5个柄细胞组成，腺头棒状，由1-8个分泌细胞构成。短柄腺毛柄较短，只有一个柄细胞，腺头球状，由8-16个分泌细胞组成。
3 香料烟叶片生长成熟过程中腺毛密度的变化规律
香料烟叶片腺毛密度随叶片的生长基本上呈逐渐下降趋势。腺毛密度在叶长1cm时最高，此时长柄腺毛密度远高于短柄腺毛密度，随后短柄腺毛产生速度稍大于长柄腺毛，其比例逐渐提高，至叶长约占定长1／2时，两种腺毛数目都达最高，但长柄腺毛仍占优势，以后腺毛数目逐渐下降。腺毛产生的关键时期在叶长约占定长1／2前，以后很少产生并逐渐脱落。香料烟叶成熟过程中腺毛密度的变化与叶片成熟度、叶位和腺毛类型有关。就腺毛总密度来看，香料烟叶从欠熟变为适熟过程中，上部叶与下部叶腺毛明显脱落，中部叶腺毛没有明显脱落；从适熟再到过熟，只有下部叶腺毛明显脱落。
香料烟腺毛和结晶细胞发育及其与环境关系研究
4香料烟腺毛的分泌活动规律
香料烟两种类型腺毛（长柄腺毛和短柄腺毛）分泌活动明显不同。香
料烟长柄腺毛分泌活动与叶片发育尤其是叶片的成熟密切相关，从欠熟叶
开始腺毛腺头细胞分泌活动开始增强，至叶片适熟时大部分腺毛达到分泌
高峰，腺头细胞着生大量胶状分泌物，叶片过熟时腺毛分泌活动减弱，腺
毛分泌物逐渐干缩。但少数腺毛分泌活动滞后，当叶片适熟时只有少量分
泌，在调制初期才大量分泌。短柄腺毛在叶片生长成熟直到调制陈化过程
中腺头细胞不会分泌像长柄腺毛一样的胶状分泌物。
5香料烟叶腺毛分泌物成分分析
经分析定性出香料烟叶15种表面化合物，这些化合物中属于腺毛分泌
物及其降解物的有5种，含量比较高的是西柏三烯二醇。
6环境对腺毛密度的影响
不同产区腺毛密度存在差异，云南香料烟长柄多细胞腺头腺毛密度、
短柄腺毛密度及腺毛总密度均高于新疆，尤其是长柄多细胞腺毛密度明显
较高。
栽培措施对香料烟腺毛密度有一定影响。肥料类型（化肥和有机肥）
对长柄多细胞腺毛密度影响较大，对长柄单细胞腺毛密度影响不大，对短
柄腺毛密度影响很小。肥料类型对上部叶和中部叶腺毛密度影响较大，对
下部叶腺毛密度影响较小。施用有机肥能明显增加上部叶长柄多细胞腺毛
密度及腺毛总密度，纯施化肥增加中部叶长柄多细胞腺毛密度。灌水次数
主要影响香料烟上部叶腺毛密度，不灌水处理上部叶腺毛密度明显较高，
灌水越多，上部叶腺毛密度越低。灌水次数对中部和下部叶腺毛密度影响
不大。
7香料烟结晶细胞的超微结构和晶体形态
2
香料烟腺毛和结晶细胞发育及其与环境关系研究
结晶细胞是一类特化的含晶细胞，细胞内充满晶体，质体等被挤到细
胞边缘。由扫描电镜观察到结晶体形状似角锥状。叶片成熟过程中结晶细
胞超微结构发生变化，欠熟叶结晶细胞胞基质、质体处于细胞边缘，质体
逐渐退化，但仍保持质体形态，胞基质清晰可见；适熟叶结晶细胞己看不
到质体，胞基质收缩；过熟叶结晶细胞胞基质干缩。
8香料烟结晶细胞的发育和分布
结晶细胞最早产生于叶长 Icm左右，发生于从下表皮往上数第 3层细
胞处。叶长2叶cm左右时结晶细胞含量迅速增加，之后随着叶长增加其含
量逐渐下降，并处于栅栏组织和海绵组织之间，呈不连续的条带状。结晶
细胞随着叶片的生长逐渐增大，叶片成熟时一些结晶细胞破裂，颗粒状结
晶物散出。结晶细胞在香料烟烟株根茎叶中均有分布，以叶中较多，并随
叶位的升高含量增加。
9香料烟叶结晶细胞部分矿质元素相对含量
香料烟叶结晶细胞与普通细胞相比有较高的钙含量，因此推测细胞内
结晶体可能是一种钙盐结晶。同时不排除结晶体可能含有尚未证实的有机
物成分。
11不同产区自然条件及栽培措施对香料烟结晶细胞含量的影响
初步试验结果表明新疆碱性土壤下生长的香料烟叶中结晶细胞的含量
稍高于云南酸性土壤生长的香料烟。雨水较少年份结晶细胞含量较高。灌
水对叶片结晶细胞含量影响较大，不灌水处理各部位叶片结晶细胞含量均
较高，灌水越多，结晶细胞含量越低。
This paper studied on the development of glandular trichomes and crystal idioblasts of oriental tobacco and the effect of environment on them in Yunnan and Xinjiang in addition to pot experiment in Henan Agriculture University during 2001-2002. The results were as follows:
The glandular trichomes of oriental tobacco was principally classified by multicell-headed long stalk trichomes,monocell-headed long stalk trichomes and multicell-headed short stalk trichomes .Branched trichomes was hardly observed. Density of short trichomes and long ones and their ratio raised with the increasing of stalk position.The long trichomes had absolute superiority on the leaves,especially on the yonger leaves. The percentage on the lower surface was higher than that on the upper surface.
The morphology development of trichomes of two types was studied. Initial cell of the trichomes generally developed into long ones. The primary long trichomes and short ones both issumed from the first and second leaf primordium about 100-200um.The differentiation cells bulging from protoderm developed into long trichomes or short ones in different mode of morphology formation. The long trichomes had longer stalk composed of 2-5 cells and one rod-liking gland head composed of l-8secretory cells. The short trichomes had shorter stalk composed of only one cell and one ball-liking gland head composed of 8-16secretory cells.
The trichome density was highest when the leaves was 1 centimetre and appeared to descend gradually during the growth of leaves. The number of trichomes came to maxium when the leaves length was half of mature leaves,then decreased. The density and number of the long trichomes prevailed over short ones during the growth of the leaves though the forming velocity of short trichomes exceeded that of long ones. The trichome density of the leaves of three maturity varied by the leaf position. The trichome density varied by maturity ,leaf position and trichome types during leaf maturing. When the immature leaves turned into proper mature, the total trichome density on the uppeer and lower leaves decreased with obvious variation while no variation on the middle leaves. When the leaves appeared over mature ,only the total trichome density on the lower leaves obviously decreased.
The secretion activities of two trichomes was observed. The long trichomes were
closely related with the stage of leaf development .especially the mature stage.The secretion of gland cells grandually strengthened in the ver-matured leaves.and came to climax when the leaves were proper mature,then weakened when the leaves appeared over mature. A minority of trichomes delayed and had a little of gummy exudate in the fine-matured leaves,consequently secreting much during curing of the leaves.The short trichomes were obviously different from the long ones.The exudate observed in the long trichomes was hardly observed in the short trichomes during the growth ,mature and aging of the leaves.
The cuticular constituents about 15 were analyzed by GC/MS,5 compounds were belonged to secretory products of the glandular trichomes and their degradations.The content of cembratriendiol was higher.
The trichome density varied in different regions.Compared with Xinjiang,there were higher densities of multicell-headed long stalk trichomes,multicell-headed short stalk trichomes and total trichomes in Yunnan,and multicell-headed long stalk trichomes were especially significant.
The effect of different-typed fertilizers on trichome density varied by trichome category and leaf position. The multicell-headed long stalk trichomes were obviously influenced,The monocell-headed long stalk trichomes were affected any the less,however, the short trichomes were hardly influenced.The trichome density in the upper and middle leaves were more influenced than that in the lower leaves.The application of organic fertilizers enhanced the denity of multicell-headed long stalk trichomes and total trichomes of the upper leaves while chemical fertilizers increased the denity of multicell-heade
引文
1．左天觉．烟草的生产、生理和生物化学．上海：上海远东出版社，1993．
2．史宏志，刘国顺．烟草香味学．北京：中国农业出版社，1998．
3．陈瑞泰．中国烟草栽培学．上海：上海科技出版社，1987．
4．陈淑珍，高致明等．烟草叶片腺毛发育及其分泌活动对烟叶品质的影响．烟草科技，1993，(4)：33．
5．王宝华，吴帼英等．烟叶的植物学特性的观察．Ⅰ烤烟叶片腺毛密度及其与烟叶品质和化学成分的关系．中国烟草科学，1983，(2)：1-6．
6．高致明，刘国顺等．香料烟叶片腺毛及分泌细胞的研究．河南农大学报，1996，(4)：329-332．
7．高致明，刘国顺等．香料烟叶片发育和结构与品质关系的研究．中国烟草学报，1993，(4)：33-39．
8．时向东，刘国顺等不同类型肥料对烤烟叶片腺毛密度、种类及分布规律的影响．中国烟草学报，1999，2：19-22．
9．时向东．不同类型肥料对烤烟生长发育和烟叶品质影响机理的研究．南京农业大学博士论文．1998．
10．韩锦峰，王广山等．烤烟叶面分泌物初步研究．中国烟草科学，1995，10-12．
11．李扬汉．植物学．上海：上海科技出版社，1993．
12．李正理．植物组织制片学．北京师范大学出版社．1996．
13．黄建成，董忠民等．腺毛的形态结构和发育．植物学报．1986，28(4)：437-440．
14．郭凤根，王仕玉．紫苏腺毛的形态结构和发育的研究．云南农业大学学报．1997，12(1)．
15．于丽杰，崔继杰．野薄荷叶腺毛的发育形态学研究．植物研究。1997，17(1)．
16．杜廷发 现代仪器分析．国防科技大学出版社．1994．
17．陆静梅．一种新型的植物细胞结晶．武汉植物研究．1991，9(3)：206-208．
18．慕小倩，安德荣等．红豆草中含晶细胞的形态学研究．西北植物学报．1994，14(5)：4-6．
19．张友玉，刘志玲等．烟草叶片细胞分离方法与扫描电镜观察．
20．林金星．裸子植物次生韧皮部中晶体的分布、大小和形状及其分类意义．Cathaya,1994，6：163-173．
21．洪健，高其康等．植物细胞结晶体的SEM观察和EDX分析．电子显微学报．2001，20(4)：517-518．
22．王学东，崔琳，小麦叶肉细胞的电镜观察．电子显微学报．1995(6)：403-405．
23．滕崇德．植物学(上册)．东北师范大学出版社．1990，34．
24．刘捷平．植物形态解剖学．北京：北京师范学院出版社．1991，66．
25．牟其云，张华云等．酶解法测定梨果肉中石细胞的含量．中国果品研究．1997，1：24-25．
26．刘鼎新等．细胞生物学研究方法与技术．北京医科大学中国协和医科大学联合出版
社．1997．
27．汤章城．现代植物生理学实验指南．北京：科学出版社．1999．
28. Barnes,J.D.The influence of UV-B radiation on the physicochemical nature of tobacco (Nicotiana tabacum L.) leaf surfaces. Joumal-of-Experimental-Botany. 1996, 47: 294, 99-109.
29. Bogdanieski,M.Trichome density in the variety Prihp7.depending on tobacco irrigations and leaf insertions. Tutun 1993,43(1-2):3-16.
30. Burk L.G. Inheritance of the glandless leaf trichome trait in Nicotiana tobaccum. Tob. Sci. 1982,26:51-53.
31. Carolyn P. Akers. Ultrastructure of glandular trichomes of leaves of Nicotiana tabacum L.,cv Xanthi. Amer. J.Bot. 1985,65(3):282-292.
32. Chakraborty M.K.The chemistry of tobacco trichomes. Tob. Sci. 1963,7:122-127.
33. Chang K.W. et.al. Changs in the surface chemisty of tobacco leaf during curing with particular emphasis ontrichomes. Tob. Int. 1985,27.
34. Chang,S.Y. et.al. Duvatrienediols,alkane,and fatty acids in cuticularwaxes of tobacco leaves of various physiological maturity. Phyto. Chem. 1976,15:961-963.
35. Court, W.A.Factors affecting the concentration of the duvatrienediols of flue-cured tobacco. Tobacco-International. 1982, 184: 7, 94-97.
36. Court, W.A.Influence of applied nitrogen fertilization on certain lipids,terpenes,andothercharacteristicsofflue-cured tobacco.Tob.Sci. 1984,28:69-72.
37. Franceschi V.R.and Harry T. Horner, JR.Calcium oxalate crystals in plants.The botanical Review. 1980,46(4).
38. Gamou, K. Studies on leaf surface lipids of tobacco. I. Changes in leaf surface lipid and duvatrienediol during growth,senescence and curing of tobacco leaves. Agric. Biol.Chem. 1979,43:2173-2168.
39. Greer E.and NielsonM.T.Leaftrichomesintobaccoandinsectrelationship:Iresistancetotobaccohornwor ms.,Manduca sextal. Tob. Int. 1985,190(13): 57-61.
40. Guo,Z.H. Biosynthesis of cembratrienols in cell-free extracts from trichomes of Nicotiana tabacum. Plant-Science-Limerick. 1995, 110: 1, 1-10.
41. Guo,Z.H. Biosynthesis of the diterpene cis-abienoi in cell-free extracts of tobacco trichomes.Archives-of-Biochemistry-and-Biophysics. 1994,308:1,103-108.
42. Heeman,V. Influence of genotype and environment on the diterpenes in the wax layer of some
flued-cured tobaccos. Beitr. Tabkforsch, 1981, 11:107-113.
43. Jackson,D.M.Effects of cuticular duvane diterpenes from green tobacco leaves on tobacco budworm(lepidoptera:Noctuidae)ovipositon.J.Chem. Ecol. 1985.
44. Johnson A.W. Evaluation of several Nicationa tobacum entries resistance to two tobacco insect pests. Tob. Sci. 1978,22:41-43.
45. Johnson J C.Inheritance of glandular trichomes in tobacco. Crop Science. 1988,22:1051-1053.
46. Johnson,A.W. Tobacco budworm infestation on early, late-planted,and sucker regrowth resistant and susceptible tobacco.29th Tobacco Workers Conference,Lexington,KY, 1981,January:19-22.
47. Johnson A.W. et.al.Tobacco leaf trichomes and their exudes.Tob. Sci. 1985,29:67-72.
48. Kandra L.et.ai. Studies of the site and mode of biosynesis of tobaccotrichomeexudatecomponents.ArchivesofBiochemistry. 1988,265(2):425-432.
49. Layten Dauis. D.and Mark T. Nielsen. Tobacco Production,Chemistry and Technology. Biackwell Science,2002, 292-303.
50. Lin. Y. Surface deposition and stability of pest-interactive,trichome-exuded diterpenes and sucrose esters of tobacco.J.Agric. Food Chem. 1994,20: 1907-20.
51. Lin. Y. Rapid and simple method for estimation of sugar esters..J.Agric. Food Chem. 1994,42:1709-12.
52. Merberg, M. Ultrastructure and secretion of glandular trichomes of tobacco leaves. Tob.aba. 1994.vol(38) No4:595.
53. Nielson M.T. Variation in tobacco genetype with different density of trichomes. Jagric.Food. Chem. 1990,38:467-470.
54. Reid,W.W. The cuticular cytoplasmic lipids of Nicotina tabacum.Ann. Du. Tabac,SEITA.Sect. 197411; 151-159.
55. Roberts D. L. and Rowland R. L.Macrocyclic diterpenes, αand β-4,8,13-duvatriene-1,3-diols. From tobacco.J.Org. Chem. 1962,27:398-392.
56. Roberto Barrera and E.A.Wrensman Trichome type, density and distribution on the leaves of certain tobacco varieties and hybrids. Tob.sci., 1966,10:157-161.
57. Roberts K L.Density of types of trichomes on leaf lamina of various tobacco cultivars. Tob. Int. 1981,183(13):80-81.
58. Severson R. F. et.al. Cuticular constituents of tobacco:factors affecting their production and their
role in insect and disease resistance and smoke quality .Rev. Adv. Tob. Sci. 1985, 11:105-174.
59.Severson ,R. F. Isolation and characterization of the sucrose esters of the cuticular waxes of green tobacco leaf. J.Agric. Food. Chem. 1985,33:870-875.
60.Simpson,R.D. Leaf surface chemistry of tobacco budworm resistant and susceptible tobacco grown in the field and greenhouse at different fertilization retes. Tob. Sci. 1985.
61.Severson R F. et.al.Quantination of major cuticular components from greenleaf of different tobacco type. J.Agric. Food.Chem. 1984,32:566-570.
62.Severson,R.F. The cuticular chemistry of N.TABACUM. CORESTA Symposium 1990,34-54.
63.wahlberg,I.Noteonthestereostructureofthunbergol(isocembrol)and4-Epiisocembrol.Acta.Chem.Scand.1981,35B:65-68.
2．史宏志，刘国顺．烟草香味学．北京：中国农业出版社，1998．
3．陈瑞泰．中国烟草栽培学．上海：上海科技出版社，1987．
4．陈淑珍，高致明等．烟草叶片腺毛发育及其分泌活动对烟叶品质的影响．烟草科技，1993，(4)：33．
5．王宝华，吴帼英等．烟叶的植物学特性的观察．Ⅰ烤烟叶片腺毛密度及其与烟叶品质和化学成分的关系．中国烟草科学，1983，(2)：1-6．
6．高致明，刘国顺等．香料烟叶片腺毛及分泌细胞的研究．河南农大学报，1996，(4)：329-332．
7．高致明，刘国顺等．香料烟叶片发育和结构与品质关系的研究．中国烟草学报，1993，(4)：33-39．
8．时向东，刘国顺等不同类型肥料对烤烟叶片腺毛密度、种类及分布规律的影响．中国烟草学报，1999，2：19-22．
9．时向东．不同类型肥料对烤烟生长发育和烟叶品质影响机理的研究．南京农业大学博士论文．1998．
10．韩锦峰，王广山等．烤烟叶面分泌物初步研究．中国烟草科学，1995，10-12．
11．李扬汉．植物学．上海：上海科技出版社，1993．
12．李正理．植物组织制片学．北京师范大学出版社．1996．
13．黄建成，董忠民等．腺毛的形态结构和发育．植物学报．1986，28(4)：437-440．
14．郭凤根，王仕玉．紫苏腺毛的形态结构和发育的研究．云南农业大学学报．1997，12(1)．
15．于丽杰，崔继杰．野薄荷叶腺毛的发育形态学研究．植物研究。1997，17(1)．
16．杜廷发 现代仪器分析．国防科技大学出版社．1994．
17．陆静梅．一种新型的植物细胞结晶．武汉植物研究．1991，9(3)：206-208．
18．慕小倩，安德荣等．红豆草中含晶细胞的形态学研究．西北植物学报．1994，14(5)：4-6．
19．张友玉，刘志玲等．烟草叶片细胞分离方法与扫描电镜观察．
20．林金星．裸子植物次生韧皮部中晶体的分布、大小和形状及其分类意义．Cathaya,1994，6：163-173．
21．洪健，高其康等．植物细胞结晶体的SEM观察和EDX分析．电子显微学报．2001，20(4)：517-518．
22．王学东，崔琳，小麦叶肉细胞的电镜观察．电子显微学报．1995(6)：403-405．
23．滕崇德．植物学(上册)．东北师范大学出版社．1990，34．
24．刘捷平．植物形态解剖学．北京：北京师范学院出版社．1991，66．
25．牟其云，张华云等．酶解法测定梨果肉中石细胞的含量．中国果品研究．1997，1：24-25．
26．刘鼎新等．细胞生物学研究方法与技术．北京医科大学中国协和医科大学联合出版
社．1997．
27．汤章城．现代植物生理学实验指南．北京：科学出版社．1999．
28. Barnes,J.D.The influence of UV-B radiation on the physicochemical nature of tobacco (Nicotiana tabacum L.) leaf surfaces. Joumal-of-Experimental-Botany. 1996, 47: 294, 99-109.
29. Bogdanieski,M.Trichome density in the variety Prihp7.depending on tobacco irrigations and leaf insertions. Tutun 1993,43(1-2):3-16.
30. Burk L.G. Inheritance of the glandless leaf trichome trait in Nicotiana tobaccum. Tob. Sci. 1982,26:51-53.
31. Carolyn P. Akers. Ultrastructure of glandular trichomes of leaves of Nicotiana tabacum L.,cv Xanthi. Amer. J.Bot. 1985,65(3):282-292.
32. Chakraborty M.K.The chemistry of tobacco trichomes. Tob. Sci. 1963,7:122-127.
33. Chang K.W. et.al. Changs in the surface chemisty of tobacco leaf during curing with particular emphasis ontrichomes. Tob. Int. 1985,27.
34. Chang,S.Y. et.al. Duvatrienediols,alkane,and fatty acids in cuticularwaxes of tobacco leaves of various physiological maturity. Phyto. Chem. 1976,15:961-963.
35. Court, W.A.Factors affecting the concentration of the duvatrienediols of flue-cured tobacco. Tobacco-International. 1982, 184: 7, 94-97.
36. Court, W.A.Influence of applied nitrogen fertilization on certain lipids,terpenes,andothercharacteristicsofflue-cured tobacco.Tob.Sci. 1984,28:69-72.
37. Franceschi V.R.and Harry T. Horner, JR.Calcium oxalate crystals in plants.The botanical Review. 1980,46(4).
38. Gamou, K. Studies on leaf surface lipids of tobacco. I. Changes in leaf surface lipid and duvatrienediol during growth,senescence and curing of tobacco leaves. Agric. Biol.Chem. 1979,43:2173-2168.
39. Greer E.and NielsonM.T.Leaftrichomesintobaccoandinsectrelationship:Iresistancetotobaccohornwor ms.,Manduca sextal. Tob. Int. 1985,190(13): 57-61.
40. Guo,Z.H. Biosynthesis of cembratrienols in cell-free extracts from trichomes of Nicotiana tabacum. Plant-Science-Limerick. 1995, 110: 1, 1-10.
41. Guo,Z.H. Biosynthesis of the diterpene cis-abienoi in cell-free extracts of tobacco trichomes.Archives-of-Biochemistry-and-Biophysics. 1994,308:1,103-108.
42. Heeman,V. Influence of genotype and environment on the diterpenes in the wax layer of some
flued-cured tobaccos. Beitr. Tabkforsch, 1981, 11:107-113.
43. Jackson,D.M.Effects of cuticular duvane diterpenes from green tobacco leaves on tobacco budworm(lepidoptera:Noctuidae)ovipositon.J.Chem. Ecol. 1985.
44. Johnson A.W. Evaluation of several Nicationa tobacum entries resistance to two tobacco insect pests. Tob. Sci. 1978,22:41-43.
45. Johnson J C.Inheritance of glandular trichomes in tobacco. Crop Science. 1988,22:1051-1053.
46. Johnson,A.W. Tobacco budworm infestation on early, late-planted,and sucker regrowth resistant and susceptible tobacco.29th Tobacco Workers Conference,Lexington,KY, 1981,January:19-22.
47. Johnson A.W. et.al.Tobacco leaf trichomes and their exudes.Tob. Sci. 1985,29:67-72.
48. Kandra L.et.ai. Studies of the site and mode of biosynesis of tobaccotrichomeexudatecomponents.ArchivesofBiochemistry. 1988,265(2):425-432.
49. Layten Dauis. D.and Mark T. Nielsen. Tobacco Production,Chemistry and Technology. Biackwell Science,2002, 292-303.
50. Lin. Y. Surface deposition and stability of pest-interactive,trichome-exuded diterpenes and sucrose esters of tobacco.J.Agric. Food Chem. 1994,20: 1907-20.
51. Lin. Y. Rapid and simple method for estimation of sugar esters..J.Agric. Food Chem. 1994,42:1709-12.
52. Merberg, M. Ultrastructure and secretion of glandular trichomes of tobacco leaves. Tob.aba. 1994.vol(38) No4:595.
53. Nielson M.T. Variation in tobacco genetype with different density of trichomes. Jagric.Food. Chem. 1990,38:467-470.
54. Reid,W.W. The cuticular cytoplasmic lipids of Nicotina tabacum.Ann. Du. Tabac,SEITA.Sect. 197411; 151-159.
55. Roberts D. L. and Rowland R. L.Macrocyclic diterpenes, αand β-4,8,13-duvatriene-1,3-diols. From tobacco.J.Org. Chem. 1962,27:398-392.
56. Roberto Barrera and E.A.Wrensman Trichome type, density and distribution on the leaves of certain tobacco varieties and hybrids. Tob.sci., 1966,10:157-161.
57. Roberts K L.Density of types of trichomes on leaf lamina of various tobacco cultivars. Tob. Int. 1981,183(13):80-81.
58. Severson R. F. et.al. Cuticular constituents of tobacco:factors affecting their production and their
role in insect and disease resistance and smoke quality .Rev. Adv. Tob. Sci. 1985, 11:105-174.
59.Severson ,R. F. Isolation and characterization of the sucrose esters of the cuticular waxes of green tobacco leaf. J.Agric. Food. Chem. 1985,33:870-875.
60.Simpson,R.D. Leaf surface chemistry of tobacco budworm resistant and susceptible tobacco grown in the field and greenhouse at different fertilization retes. Tob. Sci. 1985.
61.Severson R F. et.al.Quantination of major cuticular components from greenleaf of different tobacco type. J.Agric. Food.Chem. 1984,32:566-570.
62.Severson,R.F. The cuticular chemistry of N.TABACUM. CORESTA Symposium 1990,34-54.
63.wahlberg,I.Noteonthestereostructureofthunbergol(isocembrol)and4-Epiisocembrol.Acta.Chem.Scand.1981,35B:65-68.