藜蒿离体培养与试管苗分泌结构的研究
|
摘要
藜蒿(Artemisia selengensis)为菊科蒿属多年生草本植物,也是该属唯一的多年生水生草本植物。分布于东北、华北、华中和西南等省区,江西鄱阳湖和湖南洞庭湖区是主产区。特别是八十年代以后,形成了采食与开发的热潮。藜蒿嫩茎不仅味正香脆,还含有丰富的营养成分,被视为宴中佳品。此外,它还可以入药。它是大自然赐予人类的一种宝贵的植物资源。为了使藜蒿得到良好的保护和可持续发展的开发利用,需要为藜蒿种质保存、新种质的培育及其产业化提供一个基础性技术平台。此外,藜蒿具有特殊香味的挥发油与分泌结构密切相关。本文从云南产藜蒿离体培养入手,建立离体培养再生体系,并研究试管苗的分泌结构发生与发育,以填补藜蒿分泌结构无研究报道的空白。
主要研究结果如下:
1.建立了以云南产藜蒿为供试材料的离体培养再生体系。
云南产藜蒿的细胞全能性表达能力强,MS+BA1mg·L~(-1)+2.4-D4mg·L~(-1)培养基适宜于它的致密胚性愈伤组织的诱导;MS+BA1mg·L~(-1)+NAA0.4mg·L~(-1)培养基适宜丛芽分化和增殖;1/2MS+NAA0.5mg·L~(-1)+IAA0.5mg·L~(-1)培养基适宜试管苗生根及根状茎的形成。
2.云南产藜蒿外植体不同,所诱导出的愈伤组织在颜色、质地上普遍存在差异。
嫩茎的愈伤组织一般为浅黄绿色的胚性愈伤组织;幼叶的愈伤组织有深绿色、淡紫色、紫红色等。愈伤组织的多样性为藜蒿体细胞无性系变异的筛选提供了良好的基础。
3.细胞组织学观察,表明愈伤组织脱分化与不同外值体有差异,并发现嫩茎愈伤组织的有两种来源。
第一种是由表皮及以下多处皮层薄壁细胞脱分化而产生;第二种,是由切口处维管组织周围基本组织的薄壁细胞脱分化产生。幼叶愈伤组织主要来自部分叶肉细胞脱分化产生分生细胞,从叶缘到整个外植体都为愈伤组织所包被。与此同时,相当一部分叶肉细胞原生质体自溶,细胞壁加厚成为管状分子,这是叶肉细胞分化的标志性特征。
藜蒿幼嫩愈伤组织在结构以胚性细胞、胚性细胞团及分生组织结节的出现,但胚性细胞仍然为主,因而藜蒿愈伤组织继代能力比较强。
藜蒿愈伤组织芽分化为外起源。试管苗生根培养,根原基发生于皮层的髓射线处,由薄壁细胞脱分化产生分生细胞形成根原基,属内起源。
4.藜蒿试管苗有分泌腔和分泌腺两种分泌结构,为人工条件下系统研究植物分泌结构与发育提供了新途径。
藜蒿分泌腔分布在茎和根状茎皮层内,为裂生分泌腔。它是由排列紧密的一团细胞,彼此间的中层溶解消失,细胞互相分离并向外推移,形成一个球形囊腔,在腔内由一层具有分泌能力的上皮细胞围绕。
藜蒿试管苗茎和表面分泌着两种分泌腺。一种为腺毛状分泌腺,由10个细胞构成,2个基细胞,2个柄细胞,6个腺毛状细胞,排成2列,成熟时为扇形囊状;另一种为非腺毛状,由管状腺细胞构成。
对腺毛状分泌腺发育过程中的系统观察中发现,腺毛状分泌腺存在多样性,除二列排列外,有单列和三列排列的。二列排列的腺毛状细胞有的超过10个,单列排列的细胞如何发育还有待研究。
Artemisia selengensis.is the unique aquatic perennial herb of the artemisia which is distributed in the provinces of the northeastern,the north china,central china,and the southwestern,the PoYang lake and the Dong Ting lake is the main productive area.An upsurge of ingestion and exploitation appeared especially after the 1980s. The young stem of Artemisia selengensis is not only very delicious,but also have abundant nutrient element,which is regarded as a delicacy of the dinner.Besides,it can be used as medicine as well.And it is a precious plant resource that nature bestowed on human being.For a better protection of this plants as well as a sustainable development and exploration,it's imperative to build a new technical platform for Preservation on germplasm of Artemisia selengensis,Variety Improvement and Industrialization.Besides,Artemisia selengensis has a special flavor of Artemisia volatile oil related closely to the structure.this paper begin with the culture in vitro of Artemisia selengensis in Yunnan,build regenerated system of culture in vitro,study secretory structure and development on tube seedling of Artemisia selengensis,Which fill the blank with no reports about secretory structure of Artemisia selengensis.
The main results of this study are as follows:
1.Build culture in vitro regenerating system which take Artemisia selengensis of Yunnan as the test material.
The expressive ability about cell all-roundness of Yunnan Artemisia selengensis is very strong.It is suitable for inducing of the rapid and intense embryo callus in MS+BA1mg·L~(-1)+2.4-D4mg·L~(-1)culture medium;It is suitable for differentiation and multiplication of the bud in MS+BA1mg·L~(-1)+NAA0.4mg·L~(-1)culture medium;It is suitable for the in vitro seedling striking root and forming rhizome in 1/2MS+NAA0.5mg·L~(-1)+IAA0.5mg·L~(-1)culture medium.
2.If explants of Yunnan Artemisia selengensis are different,the colour and texture of the callus that explants induced are universally different.
Generally,the callus induced by the young stems are pistachio embryo callus; the callus induced by young leaves are deep green,mauve,purplish red,etc.The various callus lay afford a good foundation for selecting mutations of Artemisia Selengensis
3.Through cytohistology observation,it was found that there are some differences with callus dedifferentiation and different values,and there were two kind of source of the young stem callus.
The first kind is dedifferentiated from the cortical parenchyma cells of the epidermis and its below;the second kind is dedifferentiated from the parenchyma cells of the basic tissue around the vascular tissue at the incision.The young leaves callus mainly come from meristematic cells which dedifferentiate from a part of mesophyll cells,they are enveloped with the callus from the leaf margin to the whole explants.In the meanwhile,a large proportion of mesophyll cells protoplasts autolyzes,cell walls thickening into the tracheary elements,which is the character sympolizing the differentiation of the mesophyll cells.
The young callus of Artemisia selengensis appeared with structure of embryogenic cells,embryogenic cell masses and meristematic nodules,but the embryogenic cells is predominant of them,so the subculture capacity of Artemisia selengensis callus is strong.
Differentiation of the bud of Artemisia selengensis callu belongs to external origination.In rooting cultivation of the tube seedling,the root primordia is originated from the pith rays of the cortex,which is formed by the meristematic cells that dedifferentiate from the parenchyma cells.This is endogenesis in origin.
4.There are two secretory cavities and glands in Tube seedling of Artemisia selengensis,Which Provides a new way to study systemly structure and development of plant secretory under the artificial conditions.
Secretory cavities of Artemisia selengensis are cleft secretion cavity,Which are distributed in cortex of Stem and rhizome.And it is a group cell with working closely, Among which the middle class dissolved and disappear,Cell mutually separated and passaged outward,formed a spherical cavity which is arounded by a layer of Epithelial cells with secretion capacity in secretory cavities
There are two kind of glands in the young stemand young leaf of tube seedling of Artemisia selengensis.One is glandularia gland which consist of 10 cells,2 basal cells, 2 stalk cells,6 glandularia cells,they arrange in 2 rows.It was sectorial cystic when matured.The other one is non- glandularia gland which consist of tubular gland cell.
After systematic observation of the glandularia gland,we found that they are diversified,besides 2-row-arranged,they also appeared paracytic and 3-row-arranged.Some glandularia cells consist of overlOcells,We still need further study about how on paracytic cells decelop.
主要研究结果如下:
1.建立了以云南产藜蒿为供试材料的离体培养再生体系。
云南产藜蒿的细胞全能性表达能力强,MS+BA1mg·L~(-1)+2.4-D4mg·L~(-1)培养基适宜于它的致密胚性愈伤组织的诱导;MS+BA1mg·L~(-1)+NAA0.4mg·L~(-1)培养基适宜丛芽分化和增殖;1/2MS+NAA0.5mg·L~(-1)+IAA0.5mg·L~(-1)培养基适宜试管苗生根及根状茎的形成。
2.云南产藜蒿外植体不同,所诱导出的愈伤组织在颜色、质地上普遍存在差异。
嫩茎的愈伤组织一般为浅黄绿色的胚性愈伤组织;幼叶的愈伤组织有深绿色、淡紫色、紫红色等。愈伤组织的多样性为藜蒿体细胞无性系变异的筛选提供了良好的基础。
3.细胞组织学观察,表明愈伤组织脱分化与不同外值体有差异,并发现嫩茎愈伤组织的有两种来源。
第一种是由表皮及以下多处皮层薄壁细胞脱分化而产生;第二种,是由切口处维管组织周围基本组织的薄壁细胞脱分化产生。幼叶愈伤组织主要来自部分叶肉细胞脱分化产生分生细胞,从叶缘到整个外植体都为愈伤组织所包被。与此同时,相当一部分叶肉细胞原生质体自溶,细胞壁加厚成为管状分子,这是叶肉细胞分化的标志性特征。
藜蒿幼嫩愈伤组织在结构以胚性细胞、胚性细胞团及分生组织结节的出现,但胚性细胞仍然为主,因而藜蒿愈伤组织继代能力比较强。
藜蒿愈伤组织芽分化为外起源。试管苗生根培养,根原基发生于皮层的髓射线处,由薄壁细胞脱分化产生分生细胞形成根原基,属内起源。
4.藜蒿试管苗有分泌腔和分泌腺两种分泌结构,为人工条件下系统研究植物分泌结构与发育提供了新途径。
藜蒿分泌腔分布在茎和根状茎皮层内,为裂生分泌腔。它是由排列紧密的一团细胞,彼此间的中层溶解消失,细胞互相分离并向外推移,形成一个球形囊腔,在腔内由一层具有分泌能力的上皮细胞围绕。
藜蒿试管苗茎和表面分泌着两种分泌腺。一种为腺毛状分泌腺,由10个细胞构成,2个基细胞,2个柄细胞,6个腺毛状细胞,排成2列,成熟时为扇形囊状;另一种为非腺毛状,由管状腺细胞构成。
对腺毛状分泌腺发育过程中的系统观察中发现,腺毛状分泌腺存在多样性,除二列排列外,有单列和三列排列的。二列排列的腺毛状细胞有的超过10个,单列排列的细胞如何发育还有待研究。
Artemisia selengensis.is the unique aquatic perennial herb of the artemisia which is distributed in the provinces of the northeastern,the north china,central china,and the southwestern,the PoYang lake and the Dong Ting lake is the main productive area.An upsurge of ingestion and exploitation appeared especially after the 1980s. The young stem of Artemisia selengensis is not only very delicious,but also have abundant nutrient element,which is regarded as a delicacy of the dinner.Besides,it can be used as medicine as well.And it is a precious plant resource that nature bestowed on human being.For a better protection of this plants as well as a sustainable development and exploration,it's imperative to build a new technical platform for Preservation on germplasm of Artemisia selengensis,Variety Improvement and Industrialization.Besides,Artemisia selengensis has a special flavor of Artemisia volatile oil related closely to the structure.this paper begin with the culture in vitro of Artemisia selengensis in Yunnan,build regenerated system of culture in vitro,study secretory structure and development on tube seedling of Artemisia selengensis,Which fill the blank with no reports about secretory structure of Artemisia selengensis.
The main results of this study are as follows:
1.Build culture in vitro regenerating system which take Artemisia selengensis of Yunnan as the test material.
The expressive ability about cell all-roundness of Yunnan Artemisia selengensis is very strong.It is suitable for inducing of the rapid and intense embryo callus in MS+BA1mg·L~(-1)+2.4-D4mg·L~(-1)culture medium;It is suitable for differentiation and multiplication of the bud in MS+BA1mg·L~(-1)+NAA0.4mg·L~(-1)culture medium;It is suitable for the in vitro seedling striking root and forming rhizome in 1/2MS+NAA0.5mg·L~(-1)+IAA0.5mg·L~(-1)culture medium.
2.If explants of Yunnan Artemisia selengensis are different,the colour and texture of the callus that explants induced are universally different.
Generally,the callus induced by the young stems are pistachio embryo callus; the callus induced by young leaves are deep green,mauve,purplish red,etc.The various callus lay afford a good foundation for selecting mutations of Artemisia Selengensis
3.Through cytohistology observation,it was found that there are some differences with callus dedifferentiation and different values,and there were two kind of source of the young stem callus.
The first kind is dedifferentiated from the cortical parenchyma cells of the epidermis and its below;the second kind is dedifferentiated from the parenchyma cells of the basic tissue around the vascular tissue at the incision.The young leaves callus mainly come from meristematic cells which dedifferentiate from a part of mesophyll cells,they are enveloped with the callus from the leaf margin to the whole explants.In the meanwhile,a large proportion of mesophyll cells protoplasts autolyzes,cell walls thickening into the tracheary elements,which is the character sympolizing the differentiation of the mesophyll cells.
The young callus of Artemisia selengensis appeared with structure of embryogenic cells,embryogenic cell masses and meristematic nodules,but the embryogenic cells is predominant of them,so the subculture capacity of Artemisia selengensis callus is strong.
Differentiation of the bud of Artemisia selengensis callu belongs to external origination.In rooting cultivation of the tube seedling,the root primordia is originated from the pith rays of the cortex,which is formed by the meristematic cells that dedifferentiate from the parenchyma cells.This is endogenesis in origin.
4.There are two secretory cavities and glands in Tube seedling of Artemisia selengensis,Which Provides a new way to study systemly structure and development of plant secretory under the artificial conditions.
Secretory cavities of Artemisia selengensis are cleft secretion cavity,Which are distributed in cortex of Stem and rhizome.And it is a group cell with working closely, Among which the middle class dissolved and disappear,Cell mutually separated and passaged outward,formed a spherical cavity which is arounded by a layer of Epithelial cells with secretion capacity in secretory cavities
There are two kind of glands in the young stemand young leaf of tube seedling of Artemisia selengensis.One is glandularia gland which consist of 10 cells,2 basal cells, 2 stalk cells,6 glandularia cells,they arrange in 2 rows.It was sectorial cystic when matured.The other one is non- glandularia gland which consist of tubular gland cell.
After systematic observation of the glandularia gland,we found that they are diversified,besides 2-row-arranged,they also appeared paracytic and 3-row-arranged.Some glandularia cells consist of overlOcells,We still need further study about how on paracytic cells decelop.
引文
1.陶桂权.中国野菜图谱[M].北京:解放军出版社,1989.
2.中国科学院植物研究所.中国高等植物图鉴[M].北京:科学出版.
3.赖芳兰,鄱阳湖藜蒿的栽培技术[J].特种蔬菜.2005:34-35.
4.杨红.西昌地区野生藜蒿资源的开发利用[J].生物资源,2003,19(2):88-89.
5.李曙轩.中国百科全书[M].蔬菜卷,北京:农业出版社,1990.
6.谢怡孙,袁梦仙,李保同.蒌蒿精油的杀虫活性与化学成分研究[J].江西植保,2001,24(4)229-230.
7.郑功源,陈红兵,邓丹雯,白琼,佘世望.藜蒿提取物抑菌作用的初步研究[J].天然产物研究与开发,1999,11(3):72-76.
8.江西省农牧渔业厅编.江西蔬菜品种志.南昌:江西科技出版社,1986,500.
9.赵伯涛,钱桦,薛辉,张卫明.芦蒿花总黄酮提取工艺的研究.中国野生植物资源,1999,20(6):55-56.
10.叶文峰.藜蓠中微量元素的测定分析[J].微量元素与健康研究,2001,18(3):39-40.
11.杨凤岩.芦蒿不同部位营养成分比较[J].氨基酸和生物资源,2003,25(1):8-9.
12.杨红,余和明.藜蒿的生长习性、繁殖方法和高产栽培技术[J].西昌农业科技,2003(3):25-26.
13.卓根.藜蒿保鲜技术条件研究[J].贮运保鲜,2003,6:32-33.
14.李冬生,康旭.藜蒿的保鲜贮藏[J].食品研究与开发,2004,8(25):138-141.
15.钱晔,赵伯涛,张卫明,黄晓得.芦蒿采后处理及环境因素与生理生化变化的关系[J].中国野生资源,2003,4(22):50-53.
16.赵伯涛.芦蒿茶的加工工艺及质量研究[J].现代工贸商业,2003,8:44-45.
17.汪细桥,雷红卫.藜蒿生产中长出现的问题及解决措施[J].长江蔬菜,2004,9:14-15.
18.杨望明.对藜蒿无公害栽培及病虫草害防治技术[J].湖北植保,2005,4:13-14.
19.田迪英.不同热处理方法对芦蒿抗氧化活性的影响[J].研究与探讨,2005,12(26):45-47.
20.涂艺声,余兰平,王永强,卢伟芳.藜蒿的组织培养和快速繁殖研究[J].中国野生植物资源,2005,(2):59-61.
21.向国远,陈豪.不同施肥水平对藜蒿产量影响的研究[J].现代农业科技,2006,7:12-13.
22.欧阳崇学,郑为完,张桂珍,黄芝丰.藜蓠中营养成分分析[J].中国畜产与食品,1998,5(1):13-14.
23.范年友.蒌蒿的生物学特性和栽培技术[J].江汉大学学报,1999,6(12):26-28.
24.杨振国,陈彬,杨艺青.江西藜蒿的开发利用[J].中国野生植物资源,1995,1:61-62.
25.赖芳兰,邹桂花.鄱阳湖藜蒿的特性及繁殖方法的研究[J].江西农业大学学报,1998,(2):247-251.
26.张道宽.藜蒿及其栽培[J].中国蔬菜,1996(3):45-46.
27.管建国,宋佩扬.芦蒿的生物学特性及栽培中的几个问题[J].南京农专学报,1996,12(3):27-30.
28.邓丹雯,郑功源.藜蓠的营养保健功能及其产品开发[J].江西食品工业,2001,(3):18-19.
29.汤丽梅,李保国.江西藜蓠的生物学特性及栽培技术[J].中国野生植物资源,1999,19(4):53-54.
30.刘唐兴.蒌蒿的利用价值及栽培要点[J].湖南农业科学,2002,(3):23-25
31.李时珍.本草纲目[M].上海:商务印书馆,1982.
32.李芳霞,刘海军.蒌蒿的保健价值[J].营养与保健,2001.
33.黄树稳,黄绍华,温辉梁.藜蒿膳食纤维的制备、组成及性能研究[J].中国食品添加剂,1996,4:7-11.
34.肖玫.芦蒿的矿物元素新法测定及其开发利用[J].食品与发酵工业,2003,29(8):106-107.
35.顾昌华,郑利峰.世界名菊--墨菊花的组织培养与快速繁殖技术[J].种子,2006,(4):93-94.
36.路梅,张发成,王俊平.药用植物紫锥菊的组织培养与快速繁殖[J].浙江农业科学,2006(3):287-288.
37.胡海涛,刘永利,兰大伟.亚菊的组织培养与快速繁殖[J].植物生理学通讯,2006(4):93-94.
38.马宗兴,赵红,柳申飞.芦蒿的组织培养技术[J].安徽农业学报,2003,9(5):44.
39.王新华,刘伟,赵恒国.藜蒿组织培养[J].北方园艺,2002,(4):62.
40.郑成木,刘进平.热带亚热带植物繁殖[M].长沙:湖南农业科学出版社,2001,48-106.
41.刘进平.胡椒组织培养中污染问题研究[J].热带农业科学,2003,6:6-10.
42.李群,陈丽萍,石铁松.马蹄莲组织培养过程中真菌和细菌污染的消除方法研究[J].四川师范大学学报:自然科学版,2001,24(6):607-609.
43.许智宏,刘春明.植物发育的分子机理[M].北京:北京科技出版社,1999.
44.张良波,周朴华,彭晓英.屋顶长生花的离体培养[J].湖南农业大学学报,2004,10(5):433-436.
45.吴晓霞,陈刚,张彪.植物组织培养中褐变的研究进展[J].河北林果研究,2002,17(3):284-288.
46.陈菲,李黎,宫伟.植物组织培养的防褐化探讨[J].北方园艺,2005,(2):69.
47.何康.复旦南方红豆杉愈伤组织培养及褐化抑制[J].中国林业产业,2005,(12):53-55.
48.刘庆昌,吴国良主编.植物细胞住址培养[M].北京:中国农业大学出社,2005:223-258.
49.赵国林,李师翁.黄花菜离体花梗愈伤组织发生与器官再生的细胞组织学观察[J].植物学报,1989,31(6):484-486.
50.王凯基,张丕方,兜德祥.几种木本植物组织培养的愈伤组织形成和器官再生[J].植物学报,1981,23(2):97-101.
51.蒋祖丰,周朴华,王凤翱.茶树离体子叶柄愈伤组织发生与器官再生的组织学观察[J].湖南农学院学报,1993,Vol.19,Nol:107-109.
52.李扬汉主编.植物学[M].上海科技出版社,2005,第30次印刷:105-106.
53.余迪球.甜菊叶肉细胞脱分化过程中超微结构的研究[J].植物学报,1993,35(7):499-505.
54.Fukuda H.Redifferentiation of single mesophyll cells into tracheary elements[J].Plant Sei,1994,155:262-271.
55.Fukuda H.tracheary element differentiation[J].The Plant Cell,1997,9:1147-1156.
56.Denucra T,Tashiro G.Hoyoguchi G et al.Visualization by comprehensive microarry analysis of fene expression programs during transdifferentiation of mesophyll cells into xylem cells[J].PNAS,2002,99:15794-15799.
57.李正理,张新英编著.植物解剖学[M].高等教育出版社,1983:161-162.
58.刘穆著.种子植物形态解剖学导论(第二版)[M].科学出版社,2004”100-101.
59.DuKe So,Paul RN.Development and Structure of the glandular trickomes of Artemisin annua L.Int J Plant Sci,1993,156(1):107-118.
60.Ferreeira JFS,Fanick J.Flaral morphology of Artemisin annua with special reference to trichomes Int J plant Sci,1995,156(6):807-815.
61.朱卫平.野生黄花蒿的引种驯化和高青蒿素含量栽培品种选育目标形状的研究[D].湖南农业大学.
62.Weathers PJ,Cheetham RD,Follansbee E,et al,Artemisinin Production by transformed roots of Artemisia annua.Biotechnol Lett.1994,16:1281-1286.
63.Liu BY,Yw HC,Li GF,et al.Studies on synamics of growth and biosysnthesis of artemisinin in hairy roots of Artmisia annual L[J].Chin J Biotech,1999,14(4):249-254.
2.中国科学院植物研究所.中国高等植物图鉴[M].北京:科学出版.
3.赖芳兰,鄱阳湖藜蒿的栽培技术[J].特种蔬菜.2005:34-35.
4.杨红.西昌地区野生藜蒿资源的开发利用[J].生物资源,2003,19(2):88-89.
5.李曙轩.中国百科全书[M].蔬菜卷,北京:农业出版社,1990.
6.谢怡孙,袁梦仙,李保同.蒌蒿精油的杀虫活性与化学成分研究[J].江西植保,2001,24(4)229-230.
7.郑功源,陈红兵,邓丹雯,白琼,佘世望.藜蒿提取物抑菌作用的初步研究[J].天然产物研究与开发,1999,11(3):72-76.
8.江西省农牧渔业厅编.江西蔬菜品种志.南昌:江西科技出版社,1986,500.
9.赵伯涛,钱桦,薛辉,张卫明.芦蒿花总黄酮提取工艺的研究.中国野生植物资源,1999,20(6):55-56.
10.叶文峰.藜蓠中微量元素的测定分析[J].微量元素与健康研究,2001,18(3):39-40.
11.杨凤岩.芦蒿不同部位营养成分比较[J].氨基酸和生物资源,2003,25(1):8-9.
12.杨红,余和明.藜蒿的生长习性、繁殖方法和高产栽培技术[J].西昌农业科技,2003(3):25-26.
13.卓根.藜蒿保鲜技术条件研究[J].贮运保鲜,2003,6:32-33.
14.李冬生,康旭.藜蒿的保鲜贮藏[J].食品研究与开发,2004,8(25):138-141.
15.钱晔,赵伯涛,张卫明,黄晓得.芦蒿采后处理及环境因素与生理生化变化的关系[J].中国野生资源,2003,4(22):50-53.
16.赵伯涛.芦蒿茶的加工工艺及质量研究[J].现代工贸商业,2003,8:44-45.
17.汪细桥,雷红卫.藜蒿生产中长出现的问题及解决措施[J].长江蔬菜,2004,9:14-15.
18.杨望明.对藜蒿无公害栽培及病虫草害防治技术[J].湖北植保,2005,4:13-14.
19.田迪英.不同热处理方法对芦蒿抗氧化活性的影响[J].研究与探讨,2005,12(26):45-47.
20.涂艺声,余兰平,王永强,卢伟芳.藜蒿的组织培养和快速繁殖研究[J].中国野生植物资源,2005,(2):59-61.
21.向国远,陈豪.不同施肥水平对藜蒿产量影响的研究[J].现代农业科技,2006,7:12-13.
22.欧阳崇学,郑为完,张桂珍,黄芝丰.藜蓠中营养成分分析[J].中国畜产与食品,1998,5(1):13-14.
23.范年友.蒌蒿的生物学特性和栽培技术[J].江汉大学学报,1999,6(12):26-28.
24.杨振国,陈彬,杨艺青.江西藜蒿的开发利用[J].中国野生植物资源,1995,1:61-62.
25.赖芳兰,邹桂花.鄱阳湖藜蒿的特性及繁殖方法的研究[J].江西农业大学学报,1998,(2):247-251.
26.张道宽.藜蒿及其栽培[J].中国蔬菜,1996(3):45-46.
27.管建国,宋佩扬.芦蒿的生物学特性及栽培中的几个问题[J].南京农专学报,1996,12(3):27-30.
28.邓丹雯,郑功源.藜蓠的营养保健功能及其产品开发[J].江西食品工业,2001,(3):18-19.
29.汤丽梅,李保国.江西藜蓠的生物学特性及栽培技术[J].中国野生植物资源,1999,19(4):53-54.
30.刘唐兴.蒌蒿的利用价值及栽培要点[J].湖南农业科学,2002,(3):23-25
31.李时珍.本草纲目[M].上海:商务印书馆,1982.
32.李芳霞,刘海军.蒌蒿的保健价值[J].营养与保健,2001.
33.黄树稳,黄绍华,温辉梁.藜蒿膳食纤维的制备、组成及性能研究[J].中国食品添加剂,1996,4:7-11.
34.肖玫.芦蒿的矿物元素新法测定及其开发利用[J].食品与发酵工业,2003,29(8):106-107.
35.顾昌华,郑利峰.世界名菊--墨菊花的组织培养与快速繁殖技术[J].种子,2006,(4):93-94.
36.路梅,张发成,王俊平.药用植物紫锥菊的组织培养与快速繁殖[J].浙江农业科学,2006(3):287-288.
37.胡海涛,刘永利,兰大伟.亚菊的组织培养与快速繁殖[J].植物生理学通讯,2006(4):93-94.
38.马宗兴,赵红,柳申飞.芦蒿的组织培养技术[J].安徽农业学报,2003,9(5):44.
39.王新华,刘伟,赵恒国.藜蒿组织培养[J].北方园艺,2002,(4):62.
40.郑成木,刘进平.热带亚热带植物繁殖[M].长沙:湖南农业科学出版社,2001,48-106.
41.刘进平.胡椒组织培养中污染问题研究[J].热带农业科学,2003,6:6-10.
42.李群,陈丽萍,石铁松.马蹄莲组织培养过程中真菌和细菌污染的消除方法研究[J].四川师范大学学报:自然科学版,2001,24(6):607-609.
43.许智宏,刘春明.植物发育的分子机理[M].北京:北京科技出版社,1999.
44.张良波,周朴华,彭晓英.屋顶长生花的离体培养[J].湖南农业大学学报,2004,10(5):433-436.
45.吴晓霞,陈刚,张彪.植物组织培养中褐变的研究进展[J].河北林果研究,2002,17(3):284-288.
46.陈菲,李黎,宫伟.植物组织培养的防褐化探讨[J].北方园艺,2005,(2):69.
47.何康.复旦南方红豆杉愈伤组织培养及褐化抑制[J].中国林业产业,2005,(12):53-55.
48.刘庆昌,吴国良主编.植物细胞住址培养[M].北京:中国农业大学出社,2005:223-258.
49.赵国林,李师翁.黄花菜离体花梗愈伤组织发生与器官再生的细胞组织学观察[J].植物学报,1989,31(6):484-486.
50.王凯基,张丕方,兜德祥.几种木本植物组织培养的愈伤组织形成和器官再生[J].植物学报,1981,23(2):97-101.
51.蒋祖丰,周朴华,王凤翱.茶树离体子叶柄愈伤组织发生与器官再生的组织学观察[J].湖南农学院学报,1993,Vol.19,Nol:107-109.
52.李扬汉主编.植物学[M].上海科技出版社,2005,第30次印刷:105-106.
53.余迪球.甜菊叶肉细胞脱分化过程中超微结构的研究[J].植物学报,1993,35(7):499-505.
54.Fukuda H.Redifferentiation of single mesophyll cells into tracheary elements[J].Plant Sei,1994,155:262-271.
55.Fukuda H.tracheary element differentiation[J].The Plant Cell,1997,9:1147-1156.
56.Denucra T,Tashiro G.Hoyoguchi G et al.Visualization by comprehensive microarry analysis of fene expression programs during transdifferentiation of mesophyll cells into xylem cells[J].PNAS,2002,99:15794-15799.
57.李正理,张新英编著.植物解剖学[M].高等教育出版社,1983:161-162.
58.刘穆著.种子植物形态解剖学导论(第二版)[M].科学出版社,2004”100-101.
59.DuKe So,Paul RN.Development and Structure of the glandular trickomes of Artemisin annua L.Int J Plant Sci,1993,156(1):107-118.
60.Ferreeira JFS,Fanick J.Flaral morphology of Artemisin annua with special reference to trichomes Int J plant Sci,1995,156(6):807-815.
61.朱卫平.野生黄花蒿的引种驯化和高青蒿素含量栽培品种选育目标形状的研究[D].湖南农业大学.
62.Weathers PJ,Cheetham RD,Follansbee E,et al,Artemisinin Production by transformed roots of Artemisia annua.Biotechnol Lett.1994,16:1281-1286.
63.Liu BY,Yw HC,Li GF,et al.Studies on synamics of growth and biosysnthesis of artemisinin in hairy roots of Artmisia annual L[J].Chin J Biotech,1999,14(4):249-254.