基于ITS序列的东亚七筋姑多倍体复合体研究
摘要
东亚七筋姑C.udensis Trautv.et Mey.属百合科黄精族七筋姑属(ClintoniaRaf.)多年生草本植物,具有二倍体(2n=14)和四倍体(4n=28)两种细胞型。其中二倍体的分布从中国西南的云南一直延伸到俄罗斯的滨海边区。四倍体除分布在中国云南及其以西的喜马拉雅地区,日本和萨哈林岛以外,在陕西南部化龙山北坡海拔2200米以下的阔叶林或针阔混交林下和湖北木林子较狭的范围内也有分布。在东亚七筋姑种内叶、果实、花序和花粉等形态特征变异复杂多样,但反映不出倍性上的差异,显示出该种目前正处于强烈的分化阶段。本研究采用克隆测序的方法检测了13个东亚七筋姑二倍体居群和5个四倍体居群,共18个居群的rDNA ITS序列。研究表明东亚七筋姑ITS序列长度显示出较高的多态性,ITS1序列的长度在171—270bp之间,ITS2序列的长度在205—238bp之间,5.8s序列的长度在162—164bp之间,序列中GC的含量为45.83%—54.13%。当空位(gap)作缺失处理时,东亚七筋姑ITS区全序列排序后的长度为682个,其中简约性位点289个。用MEGA4.0计算Kimura双参数法遗传距离,得到18个居群之间遗传距离平均值为0.249,结果显示ITS序列在东亚七筋姑种内有着很大的变异和分歧。用SPSS进行双变量相关性分析显示,居群间遗传距离和地理距离有着较强的正相关性(r=0.372,P=0.01)。
采取最大简约法(Maximum Parsimony method,MP)、邻接法(Neighbor-joiningmethod,NJ)和最小进化法(Minimum Evolution,ME)对获得的序列进行分析,并构建分子系统树,探讨东亚七筋姑各居群间的关系和多倍体的起源方式。利用PAUP4.0b4a软件,采取最大简约法分析,获得严格一致树,其步长1056,一致性指数(CI)为0.630,保留系数(RI)为0.684。结果表明四倍体居群没有聚为单独的一支,而是和相邻地理单元的二倍体居群聚为一支,支持东亚七筋姑异地多起源的推断。
Clniotnia udensis Trautv.et Mey.,a perennial herb,belongs to the genus Clintonia Raf in the tribe Polygonateae(Liliaceae).Two ploid levels exist in this species(2n=14 and 2n=48).The diploid C.udensis is distributed from Northwest China's Yunnan Province to Russia's Primorskiy Kray,while the tetraploids are distributed in Yunnan,Himalayas,Japan,Mulinzi in Hubei and a narrow area in the northern hillside of Mt.Hualongshan below 2200 altitude in Shaanxi province.The characters of the shape in leaf,fruit,inflorescence and pollen et.al differentiate greatly in the inner species,but no differences are showed in ploid level,the results infer the species is in the stage of great differentiation.ITS sequences of 13 Clintonia undensis diploid populations and 5 tetraploid populations are reported by cloned sequencing. Sequence analysis shows that the length of ITS1 varies from 171bp to 270 bp;the length of ITS2 is from 205bp to 238 bp and 5.8S is from 162bp to 164bp.Among the 18 ITS sequences 289 sites are informative.The contents of GC of the sequences are between 45.83%～54.13%.Genetic distances are gained by MEGA4.0,the average genetic distance of 18 populations is 0.249,the results indicate that ITS sequences among the 18 populations differentiate significantly.Correlative analysis by SPSS reveals that genetic distances and geographical distances in these populations show positive correlation(r=0.372,P=0.01).
Phylogenetic analysis based on the internal transcribed spacers(ITS)of 18-26S rDNA have been conducted,using maximum parsimony method(MP), neighbor-joining method(NJ)and minimum evolution(ME),phylogenetic trees are reconstructed and origin of tetraploids Clintonia undensis and the relationship among the populations are discussed.Phylogenetic analysis of ITS sequences using PAUP4. 0b4a software obtained the MP trees(Length=1056,CI=0.630 and RI=0.684) when the gaps are treated as missing.The results indicate that the diploid populations and tetraploid populations from same region are always clustered.The origin of Clintonia undensis polyploidy is polytopic.
采取最大简约法(Maximum Parsimony method,MP)、邻接法(Neighbor-joiningmethod,NJ)和最小进化法(Minimum Evolution,ME)对获得的序列进行分析,并构建分子系统树,探讨东亚七筋姑各居群间的关系和多倍体的起源方式。利用PAUP4.0b4a软件,采取最大简约法分析,获得严格一致树,其步长1056,一致性指数(CI)为0.630,保留系数(RI)为0.684。结果表明四倍体居群没有聚为单独的一支,而是和相邻地理单元的二倍体居群聚为一支,支持东亚七筋姑异地多起源的推断。
Clniotnia udensis Trautv.et Mey.,a perennial herb,belongs to the genus Clintonia Raf in the tribe Polygonateae(Liliaceae).Two ploid levels exist in this species(2n=14 and 2n=48).The diploid C.udensis is distributed from Northwest China's Yunnan Province to Russia's Primorskiy Kray,while the tetraploids are distributed in Yunnan,Himalayas,Japan,Mulinzi in Hubei and a narrow area in the northern hillside of Mt.Hualongshan below 2200 altitude in Shaanxi province.The characters of the shape in leaf,fruit,inflorescence and pollen et.al differentiate greatly in the inner species,but no differences are showed in ploid level,the results infer the species is in the stage of great differentiation.ITS sequences of 13 Clintonia undensis diploid populations and 5 tetraploid populations are reported by cloned sequencing. Sequence analysis shows that the length of ITS1 varies from 171bp to 270 bp;the length of ITS2 is from 205bp to 238 bp and 5.8S is from 162bp to 164bp.Among the 18 ITS sequences 289 sites are informative.The contents of GC of the sequences are between 45.83%～54.13%.Genetic distances are gained by MEGA4.0,the average genetic distance of 18 populations is 0.249,the results indicate that ITS sequences among the 18 populations differentiate significantly.Correlative analysis by SPSS reveals that genetic distances and geographical distances in these populations show positive correlation(r=0.372,P=0.01).
Phylogenetic analysis based on the internal transcribed spacers(ITS)of 18-26S rDNA have been conducted,using maximum parsimony method(MP), neighbor-joining method(NJ)and minimum evolution(ME),phylogenetic trees are reconstructed and origin of tetraploids Clintonia undensis and the relationship among the populations are discussed.Phylogenetic analysis of ITS sequences using PAUP4. 0b4a software obtained the MP trees(Length=1056,CI=0.630 and RI=0.684) when the gaps are treated as missing.The results indicate that the diploid populations and tetraploid populations from same region are always clustered.The origin of Clintonia undensis polyploidy is polytopic.
引文
1.中国科学院中国植物志编辑委员会.中国植物志.1992.24-26
2.Love A,Love D.Cytotaxonomy of the alpine vascular plants of Mount Washington.Univ.Colo.Stud.Ser Biol.1966.34:27.
3.Sen S.Cytotaxonomy of Liliales.Fedd Repert.1975.86(5):255-305.
4.Sokolovskaya A P.Geograficheskoe rasprostranenie poliplodnykh vidov rasteniy.Issledovanie flory Primorskogo kraya Veste Leningard Univ Ser Biol.1966.(1):92-106.
5.Utech F H,Suda Y.Biosystematic studies on Clintonia(Liliaceae Polygonatae).ⅠSometic chromosome number and chromosomal morphology of Clintonia udensis Traurvtter et Mey.Cytologia.1975.40:169-175.
6.Utech F H.Biosystenatic studies on Clintonia(Liliaceae Polygonatae)Ⅲ.Cytogeography,chromosome number and morphology of the North Aamerican speices of Clintonia Raf Cytologia.1975.40:765-786.
7.常朝阳 李思锋.七筋姑的细胞地理学研究.植物分类学报.1996.34(1):29-38.
8.朱兆云.云南天然药物图鉴(第二卷).2003.
9.王丽,顾志建,龚洵.百合科六属十五种植物的细胞学研究.植物分类学报.1993.31(6):549-559.
10.常朝阳 李思锋,黎斌..七筋姑花粉形态种内变异及其系统学意义.西北植物学报.1997.17(1):39-42.
11.李思锋 常朝阳,杨均..七筋姑属种皮形态特征的研究.武汉植物学研究.1998.16(1):1-4.
12.李思锋.常朝阳.七筋姑属叶表皮形态特征及其系统学意义.西北植物学报.1997.17(4):468-473.
13.Seiji Yoshida Kazuhiko Hayashi,Frederick H.Utech§ and Shoichi Kawano..Molecular systematics in the genus Clintonia and related taxa based on rbcL and matK gene sequence data.Plant Species Biology.2001.16:119-137.
14.王祎玲,赵桂仿.七筋姑自然居群的遗传结构分析.云南植物研究. 2007.29(3):293-299.
15.郭晶.东亚七筋姑的遗传多样性及遗传分化——基于cpSSR标记.2007.西安:西北大学硕士论文
16.邹喻萍,葛颂.系统与进化植物学中的分子标记.2001.北京:科学出版社.
17.徐宏发,王静波.分子系统学研究进展.生态学杂志.2001.20(3):41-46.
18.Palmer J D.Olmstead RG.Chloroplast DNA systematics:a review of methods and data analysis.Amer J Bot.1994.81:1205-1224.
19.Li W-H Wolfe K H,Sharp P M.Rates of nucleotide substitution vary greatly among plant mitochondrial,choroplast,and nuclear DNAs.Proc Natl Acad Sci USA.1987.84:9054-9058.
20.Bayer R Ainouche M L.On the origins of the tetraploidBromusspecies (sectionBromus,Poaceae):insights from internal transcribed spacer sequences of nuclear ribosomal DNA.Genome.1997:730-743
21.Badaeva E D,Friebe B,Gill B S.Genome differntiation in Aegilops.2.Physical mapping of 5S and 18-26S ribosomal RNA gene famlies in diploid species.Genome.1996.39:1150-1158.
22.jiang J,Gill B S.New 18S-26S ribosomal RNA gene loci:chromosomal land marks for the evlution of polyploid wheats.Chromosoma.1994.103:179-185.
23.Baldwin B G,Sanderson M J,Porter J Met al.The ITS region of nuclear ribosomal DNA:a valuable source of evidence on angiosperm phylogeny.Ann Missouri Bot Gard.1995:247-277.
24.Rogers S O,Bendich A J.Ribosomal RNA genes in plants:variability in copy number and in the intergenic spacer.Plant Mot Biol.1987.9:509-520.
25.Turner B J Elder J R.Concerted evolution of repetitive DNA sequences in eukaryotes.Quart Rev Biol.1995.70:297-319
26.王建波,张文驹,陈家宽.核rDNA的ITS序列在被子植物系统与进化研究种的应用.植物分类学报.1999.37(4):407-416.
27.Hsiao C,Chatterton N L,Asay K H.Phylogenetic relationship of 10 grass species:an assessment of phylogenetic utility of the internal transcribed spacer region in nuclear ribosomal DNA in monocots.Genome.1994.37:112-120.
28.Baldwin B G.Phylogenetic utility of the internal transcribed spacers of nuclear ribosomal DNA in plants:an example from the Compositae.Molecular Phylogenetics and Evolution.1992.1:3-16.
29.Steve Law Nina Ronsted,Hannah Thornton,Michael F.Fay,Mark W.Chase.Molecular phylogenetic evidence for the monophyly of Fritillaria and Lilium (Liliaceae;Liliales)and the infrageneric classification of Fritillaria.Molecular Phylogenetics and Evolution.2005.35:509-527.
30.葛颂,B.A.Schaal,洪德元.用核糖体DNA的ITS序列探讨裂叶沙参的系统学位置—兼论ITS片段在沙参属系统学研究中的价值.植物分类学报.1997.35(5):385-395.
31.沈洁,丁小余,贺佳,褚必海,刘冬扬,丁鸽.泽泻rDNA ITS区序列特征及其居群鉴别研究.南京师大学报.2005.28(3):86-88.
32.张富民,葛颂,陈文俐.紫乌头复合体nrDNA的ITS序列与系统发育分析.植物分类学报.2003.41(3):220-228.
33.黎维平.三脉紫菀多倍体复合体的物种生物学研究.2002.长沙:湖南农业大学博士论文
34.潘豪杰 王祎玲,沈张波等.七筋姑基因组DNA提取方法的比较研究.西北大学学报(自然科学版).2007.37(4):617-620.
35.White T J,Bruns T,Amplification and direct sequencing of fungal ribosomal RNA gene for phylogenetics.In:Innis M Lee S,et al.,Gelfand D,Sninsky J,White T eds.PCR protocols:A Guide to methods and application[C].San Diego,California:Academic Press:315-322.
36.张文驹 王建波,陈家宽.核rDNA的ITS序列在被子植物系统与进化研究中的应用 植物分类学报.1999.37(4):407-416.
37.陈月琴,庄丽,屈良鹄,张宏达.“活化石”植物银杏形态与分子进化Ⅰ.中山大学学报.1999.38(1):16-19.
38.卢海英.中国北方8种鸢尾属植物ITS序列分析及其分子系统学意义的研究[D].2006.东北师范大学硕士论文.
39.Okazaki K Nishikawa T,Uchino T,et al.A molecular phylogeny of Lilium in the internal transcribed spacer region of nuclear ribosomal DNA.J Mol Evol. 1999.49(2):238-249.
40.曾建,张利,凡星.仲彬草属植物ITS序列PCR产物直接测序与克隆测序比较分析.四川农业大学学报.2006.24(4):369-374.
41.郑和斌,王作仁,吕作舟.美味侧耳ITS序列直接测序与克隆测序比较分析.食用菌学报.2005.12(1):1-4.
42.张文驹,应用rDNA的ITS区探讨多倍体小麦的基因组起源.1998,武汉:武汉大学博士学位论文.
43.Hamrick,JL,1990.Allozyme diversity in plant species.In:Plant Population Genetics Godt MJW,Breeding and Genetic Resources(eds Brown AHD,Clegg MT,Kahler AL,Wier BS).p:43-63.
44.Karron JD.A compareison of levels of genetic polymorphism and self-compatibility in geographically restricted and widespred plant congeners.Evolutionary Ecology.1987.1:47-58.
45.M.E.Dorken,B.C.Husband.Self-Rility in the understory herb Clintonia Borealis (Liliaceae).International Journal of Plant Sciences.1999.160(3):577.
46.J Masterson.Stomal size in fossil plants:evidence for polyploidy in majority of angiosperm.Science.1994.264:421-424.
47.Hia J.Leitch,Nichaei D.Bennett.Polyploidy in angiosperms.Trends in plant science.1997.12(2):470-476.
48.Love D Love a Origin and evolution of the arctic and alpine.1974:571-604
49.Husband BC.The role of triploid in the evolutionary dynamics of mixed-ploidy populations.Biological Journal of the Linnean Society.2004.82:537-546.
50.David JL,Benavente E,Bres-Patry C,Dusautoir J-C,Echaide M.Are neopolyploids a likely route for atransgene walk in the wild The Aegilops ovata×Triticum durum case.Biological Journal of the Linnean Society.2004.82:503-510.
51.Ramsey J,Schemske DW.Pathways,mechanisms,and rates of polyploid formation in flowering plants.Annual Review of Ecology and Systematics.1998.29:467-501.
52.王超,王建波,施苏华.山羊草属异源多倍体物种核rDNA ITS区的进化.植物分类学报.2000.38(3):211-217.
53.刘艳玲,徐立铭,倪学明等.睡莲科的系统发育:核糖体DNA ITS区序列证据.植物分类学报.2005.43(1):22-30.
54.TAO SANG,JIN PAN,DAMING ZHANG.Origins ofpolyploids:an example from peonies(Paeonia)and a model for angiosperms.Biological Journal of the Linnean Society.2004.82:561-571.
55.Levin DA.Minority cytotype exclusion in local plant populations.Taxon.1975.24:35-43.
56.张宏达,黄云晖,缪汝槐,叶创兴,廖文波,金建华.种子植物系统学.2004:598-601
57.中国科学院《中国自然地理》编辑委员会.中国植物地理—植物地理.1983:1-6
58.Lewis W.H,Biologica Relevance[M](ed.Lewis W.H).Plenum Press Polyploidy in species populations.In:Polyploidy,New York,1980:104-143.
59.Levin D.A.Polyploidy and novely in flowering plants.The American Naturalist.1983.122:1-25.
60.Lumaret R.Adaptive strategies and ploidy levels.Aacta Oecological/Oecologia Plantarum.1988.9:83-93.
61.Li SF,Chang ZY,Yuan YM.The origin and dispersal of the genus Clintonia Raf (Liliaceae):Evidence from its cytogeography and morphology.Caryologia.1996.49(2):125-135.
2.Love A,Love D.Cytotaxonomy of the alpine vascular plants of Mount Washington.Univ.Colo.Stud.Ser Biol.1966.34:27.
3.Sen S.Cytotaxonomy of Liliales.Fedd Repert.1975.86(5):255-305.
4.Sokolovskaya A P.Geograficheskoe rasprostranenie poliplodnykh vidov rasteniy.Issledovanie flory Primorskogo kraya Veste Leningard Univ Ser Biol.1966.(1):92-106.
5.Utech F H,Suda Y.Biosystematic studies on Clintonia(Liliaceae Polygonatae).ⅠSometic chromosome number and chromosomal morphology of Clintonia udensis Traurvtter et Mey.Cytologia.1975.40:169-175.
6.Utech F H.Biosystenatic studies on Clintonia(Liliaceae Polygonatae)Ⅲ.Cytogeography,chromosome number and morphology of the North Aamerican speices of Clintonia Raf Cytologia.1975.40:765-786.
7.常朝阳 李思锋.七筋姑的细胞地理学研究.植物分类学报.1996.34(1):29-38.
8.朱兆云.云南天然药物图鉴(第二卷).2003.
9.王丽,顾志建,龚洵.百合科六属十五种植物的细胞学研究.植物分类学报.1993.31(6):549-559.
10.常朝阳 李思锋,黎斌..七筋姑花粉形态种内变异及其系统学意义.西北植物学报.1997.17(1):39-42.
11.李思锋 常朝阳,杨均..七筋姑属种皮形态特征的研究.武汉植物学研究.1998.16(1):1-4.
12.李思锋.常朝阳.七筋姑属叶表皮形态特征及其系统学意义.西北植物学报.1997.17(4):468-473.
13.Seiji Yoshida Kazuhiko Hayashi,Frederick H.Utech§ and Shoichi Kawano..Molecular systematics in the genus Clintonia and related taxa based on rbcL and matK gene sequence data.Plant Species Biology.2001.16:119-137.
14.王祎玲,赵桂仿.七筋姑自然居群的遗传结构分析.云南植物研究. 2007.29(3):293-299.
15.郭晶.东亚七筋姑的遗传多样性及遗传分化——基于cpSSR标记.2007.西安:西北大学硕士论文
16.邹喻萍,葛颂.系统与进化植物学中的分子标记.2001.北京:科学出版社.
17.徐宏发,王静波.分子系统学研究进展.生态学杂志.2001.20(3):41-46.
18.Palmer J D.Olmstead RG.Chloroplast DNA systematics:a review of methods and data analysis.Amer J Bot.1994.81:1205-1224.
19.Li W-H Wolfe K H,Sharp P M.Rates of nucleotide substitution vary greatly among plant mitochondrial,choroplast,and nuclear DNAs.Proc Natl Acad Sci USA.1987.84:9054-9058.
20.Bayer R Ainouche M L.On the origins of the tetraploidBromusspecies (sectionBromus,Poaceae):insights from internal transcribed spacer sequences of nuclear ribosomal DNA.Genome.1997:730-743
21.Badaeva E D,Friebe B,Gill B S.Genome differntiation in Aegilops.2.Physical mapping of 5S and 18-26S ribosomal RNA gene famlies in diploid species.Genome.1996.39:1150-1158.
22.jiang J,Gill B S.New 18S-26S ribosomal RNA gene loci:chromosomal land marks for the evlution of polyploid wheats.Chromosoma.1994.103:179-185.
23.Baldwin B G,Sanderson M J,Porter J Met al.The ITS region of nuclear ribosomal DNA:a valuable source of evidence on angiosperm phylogeny.Ann Missouri Bot Gard.1995:247-277.
24.Rogers S O,Bendich A J.Ribosomal RNA genes in plants:variability in copy number and in the intergenic spacer.Plant Mot Biol.1987.9:509-520.
25.Turner B J Elder J R.Concerted evolution of repetitive DNA sequences in eukaryotes.Quart Rev Biol.1995.70:297-319
26.王建波,张文驹,陈家宽.核rDNA的ITS序列在被子植物系统与进化研究种的应用.植物分类学报.1999.37(4):407-416.
27.Hsiao C,Chatterton N L,Asay K H.Phylogenetic relationship of 10 grass species:an assessment of phylogenetic utility of the internal transcribed spacer region in nuclear ribosomal DNA in monocots.Genome.1994.37:112-120.
28.Baldwin B G.Phylogenetic utility of the internal transcribed spacers of nuclear ribosomal DNA in plants:an example from the Compositae.Molecular Phylogenetics and Evolution.1992.1:3-16.
29.Steve Law Nina Ronsted,Hannah Thornton,Michael F.Fay,Mark W.Chase.Molecular phylogenetic evidence for the monophyly of Fritillaria and Lilium (Liliaceae;Liliales)and the infrageneric classification of Fritillaria.Molecular Phylogenetics and Evolution.2005.35:509-527.
30.葛颂,B.A.Schaal,洪德元.用核糖体DNA的ITS序列探讨裂叶沙参的系统学位置—兼论ITS片段在沙参属系统学研究中的价值.植物分类学报.1997.35(5):385-395.
31.沈洁,丁小余,贺佳,褚必海,刘冬扬,丁鸽.泽泻rDNA ITS区序列特征及其居群鉴别研究.南京师大学报.2005.28(3):86-88.
32.张富民,葛颂,陈文俐.紫乌头复合体nrDNA的ITS序列与系统发育分析.植物分类学报.2003.41(3):220-228.
33.黎维平.三脉紫菀多倍体复合体的物种生物学研究.2002.长沙:湖南农业大学博士论文
34.潘豪杰 王祎玲,沈张波等.七筋姑基因组DNA提取方法的比较研究.西北大学学报(自然科学版).2007.37(4):617-620.
35.White T J,Bruns T,Amplification and direct sequencing of fungal ribosomal RNA gene for phylogenetics.In:Innis M Lee S,et al.,Gelfand D,Sninsky J,White T eds.PCR protocols:A Guide to methods and application[C].San Diego,California:Academic Press:315-322.
36.张文驹 王建波,陈家宽.核rDNA的ITS序列在被子植物系统与进化研究中的应用 植物分类学报.1999.37(4):407-416.
37.陈月琴,庄丽,屈良鹄,张宏达.“活化石”植物银杏形态与分子进化Ⅰ.中山大学学报.1999.38(1):16-19.
38.卢海英.中国北方8种鸢尾属植物ITS序列分析及其分子系统学意义的研究[D].2006.东北师范大学硕士论文.
39.Okazaki K Nishikawa T,Uchino T,et al.A molecular phylogeny of Lilium in the internal transcribed spacer region of nuclear ribosomal DNA.J Mol Evol. 1999.49(2):238-249.
40.曾建,张利,凡星.仲彬草属植物ITS序列PCR产物直接测序与克隆测序比较分析.四川农业大学学报.2006.24(4):369-374.
41.郑和斌,王作仁,吕作舟.美味侧耳ITS序列直接测序与克隆测序比较分析.食用菌学报.2005.12(1):1-4.
42.张文驹,应用rDNA的ITS区探讨多倍体小麦的基因组起源.1998,武汉:武汉大学博士学位论文.
43.Hamrick,JL,1990.Allozyme diversity in plant species.In:Plant Population Genetics Godt MJW,Breeding and Genetic Resources(eds Brown AHD,Clegg MT,Kahler AL,Wier BS).p:43-63.
44.Karron JD.A compareison of levels of genetic polymorphism and self-compatibility in geographically restricted and widespred plant congeners.Evolutionary Ecology.1987.1:47-58.
45.M.E.Dorken,B.C.Husband.Self-Rility in the understory herb Clintonia Borealis (Liliaceae).International Journal of Plant Sciences.1999.160(3):577.
46.J Masterson.Stomal size in fossil plants:evidence for polyploidy in majority of angiosperm.Science.1994.264:421-424.
47.Hia J.Leitch,Nichaei D.Bennett.Polyploidy in angiosperms.Trends in plant science.1997.12(2):470-476.
48.Love D Love a Origin and evolution of the arctic and alpine.1974:571-604
49.Husband BC.The role of triploid in the evolutionary dynamics of mixed-ploidy populations.Biological Journal of the Linnean Society.2004.82:537-546.
50.David JL,Benavente E,Bres-Patry C,Dusautoir J-C,Echaide M.Are neopolyploids a likely route for atransgene walk in the wild The Aegilops ovata×Triticum durum case.Biological Journal of the Linnean Society.2004.82:503-510.
51.Ramsey J,Schemske DW.Pathways,mechanisms,and rates of polyploid formation in flowering plants.Annual Review of Ecology and Systematics.1998.29:467-501.
52.王超,王建波,施苏华.山羊草属异源多倍体物种核rDNA ITS区的进化.植物分类学报.2000.38(3):211-217.
53.刘艳玲,徐立铭,倪学明等.睡莲科的系统发育:核糖体DNA ITS区序列证据.植物分类学报.2005.43(1):22-30.
54.TAO SANG,JIN PAN,DAMING ZHANG.Origins ofpolyploids:an example from peonies(Paeonia)and a model for angiosperms.Biological Journal of the Linnean Society.2004.82:561-571.
55.Levin DA.Minority cytotype exclusion in local plant populations.Taxon.1975.24:35-43.
56.张宏达,黄云晖,缪汝槐,叶创兴,廖文波,金建华.种子植物系统学.2004:598-601
57.中国科学院《中国自然地理》编辑委员会.中国植物地理—植物地理.1983:1-6
58.Lewis W.H,Biologica Relevance[M](ed.Lewis W.H).Plenum Press Polyploidy in species populations.In:Polyploidy,New York,1980:104-143.
59.Levin D.A.Polyploidy and novely in flowering plants.The American Naturalist.1983.122:1-25.
60.Lumaret R.Adaptive strategies and ploidy levels.Aacta Oecological/Oecologia Plantarum.1988.9:83-93.
61.Li SF,Chang ZY,Yuan YM.The origin and dispersal of the genus Clintonia Raf (Liliaceae):Evidence from its cytogeography and morphology.Caryologia.1996.49(2):125-135.