七种野生铁线莲属植物核型分析
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摘要
铁线莲属植物种间在形态、香味、花量、绿期及生活型等方面差异较大,遗传背景也相对复杂,因此在杂交育种前了解其核型特征意义重大。采用染色体常规压片法,对东方铁线莲、大花铁线莲、大叶铁线莲、褐毛铁线莲、棉团铁线莲、齿叶铁线莲和东北铁线莲7种野生铁线莲属植物的倍性及核型进行分析,结果表明,1)7种野生铁线莲中除东北铁线莲为四倍体(2n=4x=32)外,另外6种均为二倍体(2n=2x=16),染色体基数为8。染色体类型有中部(m)、亚中部(sm)、亚端部(st)、端部(t)4种,其中大花铁线莲和大叶铁线莲的核型公式为2n=2x=16=5m+2st+t,褐毛铁线莲的核型公式为2n=2x=16=5m+3st,棉团铁线莲的核型公式为2n=2x=16=5m+2st(SAT)+t,齿叶铁线莲的核型公式为2n=2x=16=5m(SAT)+2st+t,东方铁线莲的核型公式为2n=2x=16=5m(SAT)+sm(SAT)+2st,东北铁线莲的核型公式为2n=4x=32=5m+2st+t。东方铁线莲、棉团铁线莲、齿叶铁线莲的短臂上带有随体。2)核型不对称系数为62.27%~66.77%,核型类型均为2A型,表明7种野生铁线莲属植物的进化程度均较原始。其中,褐毛铁线莲和齿叶铁线莲为首次报道。
There are many significant differences in the wild species of Clematis,including differences in morphology,aroma,flower number,green period and life form.These differences reflect the species' complex genetic background.It is advantageous to study the chromosome karyotypes of the various Clematis species prior to any attempts at cross-breeding.In this study,traditional squashing methods were used to study the karyotypes of 7 wild Clematis species:Clematis orientalis,C.patens,C.heracleifolia,C.fusca,C.hexapetala,C.serratifoliaand C.ternifloravar.mandshurica.The results showed that six of the Clematis species were diploid(2 n=2 x=16),with the exception being C.terniflora var.mandshurica,which is tetraploid(2 n=4 x=32).The basic chromosome number was 8.The karyotypes of all the tested Clematis species were mainly made of median,submedian,subterminal and terminal chromosomes.Their karyotype formulae are as follows C.patens and C.heracleifolia:2 n=2 x=16=5 m+2 st+t;C.fusca:2 n=2 x=16=5 m+3 st;C.hexapetala:2 n=2 x=16=5 m+2 st(SAT)+t;C.serratifolia:2 n=2 x=16=5 m(SAT)+2 st+t;C.orientalis:2 n=2 x=16=5 m(SAT)+sm(SAT)+2 st;C.ternifloravar.mandshurica:2 n=4 x=32=5 m+2 st+t.The karyotype formulae of C.orientalis,C.hexapetala and C.serratifolia had satellites in the long arm of their chromosomes.The asymmetry index of the chromosomes ranged from 62.27% to 66.77%.The karyotypes of these Clematis species were 2 A,showing that these seven species are relatively primitive.The karyotypes of C.fuscaand C.serratifoliaare reported here for the first time.
There are many significant differences in the wild species of Clematis,including differences in morphology,aroma,flower number,green period and life form.These differences reflect the species' complex genetic background.It is advantageous to study the chromosome karyotypes of the various Clematis species prior to any attempts at cross-breeding.In this study,traditional squashing methods were used to study the karyotypes of 7 wild Clematis species:Clematis orientalis,C.patens,C.heracleifolia,C.fusca,C.hexapetala,C.serratifoliaand C.ternifloravar.mandshurica.The results showed that six of the Clematis species were diploid(2 n=2 x=16),with the exception being C.terniflora var.mandshurica,which is tetraploid(2 n=4 x=32).The basic chromosome number was 8.The karyotypes of all the tested Clematis species were mainly made of median,submedian,subterminal and terminal chromosomes.Their karyotype formulae are as follows C.patens and C.heracleifolia:2 n=2 x=16=5 m+2 st+t;C.fusca:2 n=2 x=16=5 m+3 st;C.hexapetala:2 n=2 x=16=5 m+2 st(SAT)+t;C.serratifolia:2 n=2 x=16=5 m(SAT)+2 st+t;C.orientalis:2 n=2 x=16=5 m(SAT)+sm(SAT)+2 st;C.ternifloravar.mandshurica:2 n=4 x=32=5 m+2 st+t.The karyotype formulae of C.orientalis,C.hexapetala and C.serratifolia had satellites in the long arm of their chromosomes.The asymmetry index of the chromosomes ranged from 62.27% to 66.77%.The karyotypes of these Clematis species were 2 A,showing that these seven species are relatively primitive.The karyotypes of C.fuscaand C.serratifoliaare reported here for the first time.
引文
[1]Pei H M,Mao H L.Study on the germplasm resources and fancy application of wild Clematis L.in south Gansu province.Chinese Wild Plant Resources,2004,23(6):30-32.裴会明,毛浩龙.甘肃南部野生铁线莲属植物种质资源及观赏应用.中国野生植物资源,2004,23(6):30-32.
[2]Ma D.Research on Cold Resistance of Clematis heracleifoliain Harbin[D].Harbin:Northeast Forestry University,2010.马迪.哈尔滨地区大叶铁线莲抗寒性的研究[D].哈尔滨:东北林业大学,2010.
[3]Jiang L Y.Effect of Na2CO3,NaHCO3 Stress on Morphological and Physiological of Clematis heracleifolia[D].Harbin:Northeast Forestry University,2011.姜丽颖.Na2CO3、NaHCO3胁迫对大叶铁线莲形态和生理的影响[D].哈尔滨:东北林业大学,2011.
[4]Yang L,Yao G Y,Xue H Y,et al.Antitum or activities of total saponins fromClematis hexapetala Pall.Acta Chinese Medicine and Pharmacology,2011,39(2):21-24.杨林,姚广玉,薛鸿燕,等.棉团铁线莲总皂苷抗肿瘤作用研究.中医药学报,2011,39(2):21-24.
[5]Feng L F,Weng M Y,Tian F R,et al.Comparasion on total flavonoid content from aerial parts of 11 medicinal plants of Clematis L.Modern Chinese Medicine,2011,13(7):14-18.冯玲枫,翁美娅,田富饶,等.11种铁线莲属药用植物地上部分总黄酮含量比较研究.中国现代中药,2011,13(7):14-18.
[6]Liu H M,Zhi L,Zhao L H,et al.Karyotype analysis of four wild Liliumspecies.Journal of Plant Genetic Resources,2010,11(4):469-473.刘华敏,智丽,赵丽华,等.四种野生百合核型分析.植物遗传资源学报,2010,11(4):469-473.
[7]Gong W Z,Long Y Y,Li M X.Karyotype studies on Clematis from Beijing China.Journal of Wuhan Botanical Research,1985,3(4):371-379.龚维忠,龙雅宜,李懋学.北京地区铁线莲属植物的核型研究.武汉植物学研究,1985,3(4):371-379.
[8]Zhang Y L,He S Y.Chromosome studies on 6species of Clematis in China.Journal of Wuhan Botanical Research,1990,8(2):115-121.张镱锂,贺士元.国产6种铁线莲的染色体研究.武汉植物学研究,1990,8(2):115-121.
[9]Zhang Y L.Chromosome studies on 7species of Clematis in China.Journal of Wuhan Botanical Research,1991,9(2):107-111,201-202.张镱锂.7种铁线莲的染色体研究.武汉植物学研究,1991,9(2):107-111,201-202.
[10]Yang Q E,Luo Y B,Hong D Y.A karyotypic study of species in the Ranunculaceae from Hunan.Guihaia,1994,14(1):27-36,99-101.杨亲二,罗毅波,洪德元.湖南6种毛茛科植物的核型研究.广西植物,1994,14(1):27-36,99-101.
[11]Peng L C,Yu H J,Yu N,et al.Karyotype features of 10species of Clematis and cluster analysis of karyotypes resemblancenear coefficient.Journal of Hunan Agricultural University:Natural Sciences,2012,38(6):617-622.彭绿春,于恒隽,余娜,等.10种铁线莲的核型特征及核型似近系数聚类分析.湖南农业大学学报:自然科学版,2012,38(6):617-622.
[12]Sheng L.Karyotype Analysis on Sixteen Species of Genus Clematis[D].Nanjing:Nanjing Forestry University,2011.盛璐.铁钱莲属16种植物的核型分析[D].南京:南京林业大学,2011.
[13]Sheng L.Karyotype Analysis and Molecular Systematics Study of Genus Clematis[D].Nanjing:Nanjing Forestry University,2015.盛璐.铁线莲属植物的核型分析及分子系统学研究[D].南京:南京林业大学,2015.
[14]Sheidai M,Habibi M,Azizian D.Cytology and palynology of the Clematis L.species(Ranunculaceae)in Iran.Acta Botanica Croatica,2009,68(1):67-77.
[15]Duan Q,Cui G F,Wang X N,et al.Genetic diversity of Lilium sargentiae based on phenotypic traits and ISSR markers.Acta Botanica Boreali-Occidentalia Sinica,2013,33(6):1106-1113.段青,崔光芬,王祥宁,等.云南泸定百合遗传多样性的表型与ISSR分析.西北植物学报,2013,33(6):1106-1113.
[16]Li Z Y,Hua Y W,Ge X H,et al.Chromosomal behaviors in plant wide hybridizations and their genetic and evolutionary implications.Hereditas(Beijing),2005,(2):315-324.李再云,华玉伟,葛贤宏,等.植物远缘杂交中的染色体行为及其遗传与进化意义.遗传,2005,(2):315-324.
[17]Li M X,Zhang Z P.Chromosome and Research Techniques of Crops[M].Beijing:China Agriculture Press,1996:292-297.李懋学,张赞平.作物染色体及其研究技术[M].北京:中国农业出版社,1996:292-297.
[18]Li M X,Chen R Y.The standardization about the karyotype analysis.Journal of Wuhan Botanical Research,1985,3(4):297-302.李懋学,陈瑞阳.关于核型分析的标准化问题.武汉植物学研究,1985,3(4):297-302.
[19]Stebbins G L.Chromosomal Evolution in Higher Plants[M].London:Edward Arnold Press,1971:87-89.
[20]Levan A,Fredga K,Sandberg A A.Nomenclature for centromeric position on chromosomes.Hereditas,1964,52(2):201-220.
[21]Arano H.Cytological studies in subfamily Carduoideae(Compositae)of Japan.Botanical Magazine of Tokyo,1963,76:32-39.
[22]Zhou Q X,Yang Y P,Zhang M L.Karyotypes of fourteen species in Caragana.Bulletin of Botanical Research,2002,22(4):492-496,518-519.周其兴,杨永平,张明理.锦鸡儿属植物14个种类的核型.植物研究,2002,22(4):492-496,518-519.
[23]Shang B L,Meng K B,Wang Y J.Karyotype analysis of five Saussureaspecies.Acta Botanica Boreali-Occidentalia Sinica,2014,34(11):2220-2226.尚宝龙,蒙奎宾,王玉金.风毛菊属5种植物的核型分析.西北植物学报,2014,34(11):2220-2226.
[24]Zhang F,Xing S Y,Han C J,et al.Analysis on karyotype and its evolutional trend of Ginkgo biloba var.epiphyllagermplasms resources.Acta Horticulturae Sinica,2011,38(12):2245-2252.张芳,邢世岩,韩晨静,等.叶籽银杏种质资源染色体核型分析及进化趋势.园艺学报,2011,38(12):2245-2252.
[25]Li Z,Chen S M,Chen F D,et al.Distribution of 45SrDNA sequence on chromosomes of five species in Artemisia.Acta Horticulturae Sinica,2011,38(2):353-360.李真,陈素梅,陈发棣,等.45SrDNA在蒿属5种植物染色体中的分布.园艺学报,2011,38(2):353-360.
[2]Ma D.Research on Cold Resistance of Clematis heracleifoliain Harbin[D].Harbin:Northeast Forestry University,2010.马迪.哈尔滨地区大叶铁线莲抗寒性的研究[D].哈尔滨:东北林业大学,2010.
[3]Jiang L Y.Effect of Na2CO3,NaHCO3 Stress on Morphological and Physiological of Clematis heracleifolia[D].Harbin:Northeast Forestry University,2011.姜丽颖.Na2CO3、NaHCO3胁迫对大叶铁线莲形态和生理的影响[D].哈尔滨:东北林业大学,2011.
[4]Yang L,Yao G Y,Xue H Y,et al.Antitum or activities of total saponins fromClematis hexapetala Pall.Acta Chinese Medicine and Pharmacology,2011,39(2):21-24.杨林,姚广玉,薛鸿燕,等.棉团铁线莲总皂苷抗肿瘤作用研究.中医药学报,2011,39(2):21-24.
[5]Feng L F,Weng M Y,Tian F R,et al.Comparasion on total flavonoid content from aerial parts of 11 medicinal plants of Clematis L.Modern Chinese Medicine,2011,13(7):14-18.冯玲枫,翁美娅,田富饶,等.11种铁线莲属药用植物地上部分总黄酮含量比较研究.中国现代中药,2011,13(7):14-18.
[6]Liu H M,Zhi L,Zhao L H,et al.Karyotype analysis of four wild Liliumspecies.Journal of Plant Genetic Resources,2010,11(4):469-473.刘华敏,智丽,赵丽华,等.四种野生百合核型分析.植物遗传资源学报,2010,11(4):469-473.
[7]Gong W Z,Long Y Y,Li M X.Karyotype studies on Clematis from Beijing China.Journal of Wuhan Botanical Research,1985,3(4):371-379.龚维忠,龙雅宜,李懋学.北京地区铁线莲属植物的核型研究.武汉植物学研究,1985,3(4):371-379.
[8]Zhang Y L,He S Y.Chromosome studies on 6species of Clematis in China.Journal of Wuhan Botanical Research,1990,8(2):115-121.张镱锂,贺士元.国产6种铁线莲的染色体研究.武汉植物学研究,1990,8(2):115-121.
[9]Zhang Y L.Chromosome studies on 7species of Clematis in China.Journal of Wuhan Botanical Research,1991,9(2):107-111,201-202.张镱锂.7种铁线莲的染色体研究.武汉植物学研究,1991,9(2):107-111,201-202.
[10]Yang Q E,Luo Y B,Hong D Y.A karyotypic study of species in the Ranunculaceae from Hunan.Guihaia,1994,14(1):27-36,99-101.杨亲二,罗毅波,洪德元.湖南6种毛茛科植物的核型研究.广西植物,1994,14(1):27-36,99-101.
[11]Peng L C,Yu H J,Yu N,et al.Karyotype features of 10species of Clematis and cluster analysis of karyotypes resemblancenear coefficient.Journal of Hunan Agricultural University:Natural Sciences,2012,38(6):617-622.彭绿春,于恒隽,余娜,等.10种铁线莲的核型特征及核型似近系数聚类分析.湖南农业大学学报:自然科学版,2012,38(6):617-622.
[12]Sheng L.Karyotype Analysis on Sixteen Species of Genus Clematis[D].Nanjing:Nanjing Forestry University,2011.盛璐.铁钱莲属16种植物的核型分析[D].南京:南京林业大学,2011.
[13]Sheng L.Karyotype Analysis and Molecular Systematics Study of Genus Clematis[D].Nanjing:Nanjing Forestry University,2015.盛璐.铁线莲属植物的核型分析及分子系统学研究[D].南京:南京林业大学,2015.
[14]Sheidai M,Habibi M,Azizian D.Cytology and palynology of the Clematis L.species(Ranunculaceae)in Iran.Acta Botanica Croatica,2009,68(1):67-77.
[15]Duan Q,Cui G F,Wang X N,et al.Genetic diversity of Lilium sargentiae based on phenotypic traits and ISSR markers.Acta Botanica Boreali-Occidentalia Sinica,2013,33(6):1106-1113.段青,崔光芬,王祥宁,等.云南泸定百合遗传多样性的表型与ISSR分析.西北植物学报,2013,33(6):1106-1113.
[16]Li Z Y,Hua Y W,Ge X H,et al.Chromosomal behaviors in plant wide hybridizations and their genetic and evolutionary implications.Hereditas(Beijing),2005,(2):315-324.李再云,华玉伟,葛贤宏,等.植物远缘杂交中的染色体行为及其遗传与进化意义.遗传,2005,(2):315-324.
[17]Li M X,Zhang Z P.Chromosome and Research Techniques of Crops[M].Beijing:China Agriculture Press,1996:292-297.李懋学,张赞平.作物染色体及其研究技术[M].北京:中国农业出版社,1996:292-297.
[18]Li M X,Chen R Y.The standardization about the karyotype analysis.Journal of Wuhan Botanical Research,1985,3(4):297-302.李懋学,陈瑞阳.关于核型分析的标准化问题.武汉植物学研究,1985,3(4):297-302.
[19]Stebbins G L.Chromosomal Evolution in Higher Plants[M].London:Edward Arnold Press,1971:87-89.
[20]Levan A,Fredga K,Sandberg A A.Nomenclature for centromeric position on chromosomes.Hereditas,1964,52(2):201-220.
[21]Arano H.Cytological studies in subfamily Carduoideae(Compositae)of Japan.Botanical Magazine of Tokyo,1963,76:32-39.
[22]Zhou Q X,Yang Y P,Zhang M L.Karyotypes of fourteen species in Caragana.Bulletin of Botanical Research,2002,22(4):492-496,518-519.周其兴,杨永平,张明理.锦鸡儿属植物14个种类的核型.植物研究,2002,22(4):492-496,518-519.
[23]Shang B L,Meng K B,Wang Y J.Karyotype analysis of five Saussureaspecies.Acta Botanica Boreali-Occidentalia Sinica,2014,34(11):2220-2226.尚宝龙,蒙奎宾,王玉金.风毛菊属5种植物的核型分析.西北植物学报,2014,34(11):2220-2226.
[24]Zhang F,Xing S Y,Han C J,et al.Analysis on karyotype and its evolutional trend of Ginkgo biloba var.epiphyllagermplasms resources.Acta Horticulturae Sinica,2011,38(12):2245-2252.张芳,邢世岩,韩晨静,等.叶籽银杏种质资源染色体核型分析及进化趋势.园艺学报,2011,38(12):2245-2252.
[25]Li Z,Chen S M,Chen F D,et al.Distribution of 45SrDNA sequence on chromosomes of five species in Artemisia.Acta Horticulturae Sinica,2011,38(2):353-360.李真,陈素梅,陈发棣,等.45SrDNA在蒿属5种植物染色体中的分布.园艺学报,2011,38(2):353-360.
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