锡盟沙地榆（Ulmus pumila var.sabulosa）遗传多样性及其与家榆（U.pumila）的比较研究
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摘要
本文以分布于内蒙古浑善达克沙地的锡盟沙地榆( Ulmus pumila L.var.sabulosa)及对照家榆(Ulmus pumila)为研究对象,运用形态学、孢粉学、解剖学和RAPD分子标记等方法对其进行系统的研究,通过实验研究表明:
锡盟沙地榆与家榆在植株形态、花粉形态特征和叶片解剖结构方面具有相同的特征,但同时也存在一定的差异,研究结果表明,在漫长的演化过程中,锡盟沙地榆因长期适应其所处的浑善达克沙地生态环境而产生植株形态、花粉形态及解剖特征的变异。
建立了适于锡盟沙地榆的稳定的RAPD反应体系及反应程序,RAPD研究结果如下:14个随机引物在锡盟沙地榆和家榆中共检测到118个位点,平均每个引物扩增带数为8.4条,其中多态位点83个,占70.34%。选取了多态位点比率、Nei's多样性指数(h)和Shannon's多样性指数(I)来评价锡盟沙地榆与对照家榆居群的遗传多样性水平,结果表明锡盟沙地榆与家榆均具有较高水平的遗传多样性,且锡盟沙地榆遗传多样性水平明显高于家榆。在假设Hardy-Weinberg平衡时计算出锡盟沙地榆六个居群间的遗传分化系数Gst为0.3762,即在总的遗传变异中有37.62%存在于居群之间,居群内的遗传变异所占比例为62.38%,六个居群之间有相似的遗传多样性,但由于地理和距离的隔离,居群间存在有一定的遗传分化。根据遗传距离和聚类分析得出:锡盟沙地榆和对照家榆明显聚为两大类,表明经过漫长的进化,锡盟沙地榆产生并保留了较大的遗传变异。锡盟沙地榆及对照家榆各居群间的地理分布与其聚类结果存在着极大的相关性。
结合形态学、解剖学、孢粉学和RAPD研究结果,本文支持将锡盟沙地榆作为家榆一变种的结论。
The research objects are Ulmus pumila L.var.sabulosa which distribution on Hunshandake sandland and Ulmus pumila as comparision.The results of the morphologic,palynology,anatomy and RAPD indicate that:
There are same characteristics between Ulmus pumila L.var.sabulosa and U.s pumila in the shape characteristic, pollen shape characteristic and the structure of leaf, but also has differences. The results show that U. pumila L.var.sabulosa has occurred variation in the shape characteristic, pollen shape characteristic and the structure of leaf for acclimatize itself to entironment in the long process of evolvement.
118 reproducible loci are yielded based on 14 decamer primers from U.s pumila L.var.sabulosa and U. pumila,of which 83 are polymorphic .The average is 8.4 per decamer primers and the percentage of polymorphic loci equaled 70.34%. The percentage of polymorphic loci, Nei's (1973)gene diversity and Shannon's information index are choosed to estimate the genetic diversity of U. pumila L.var.sabulosa and U. pumila, the results show that U. pumila L.var.sabulosa and U. pumila has higher genetic diversity level, of which U. pumila L.var.sabulosa are higher than U. pumila distinctly. The coefficient of gene differentiation (Gst) based on supposing Hardy-Weinberg among populations is 0.3762, which reveal that 37.62% of the genetic variance existed among populations and 62.38% within populations. There were conform genetic diversity in the six populations, while there were definite gene differentiation for being obstructed. According to the genetic distance and cluster analysis (UPGMA), U. pumila L.var.sabulosa and U. pumila can be sorted into 2 different groups. The results demonstrate that U. pumila L.var.sabulosa has existenced and keeped biggest gene differentiation as endless evolution. There are pertinence between the geography distribution and cluster results of the populations U. pumila L.var.sabulosa and U. pumila.
The results demonstrate that U. pumila L.var. sabulosa can be regarded as a variety of U. pumila based on the results of the morphologic, palynology, anatomy and RAPD.
锡盟沙地榆与家榆在植株形态、花粉形态特征和叶片解剖结构方面具有相同的特征,但同时也存在一定的差异,研究结果表明,在漫长的演化过程中,锡盟沙地榆因长期适应其所处的浑善达克沙地生态环境而产生植株形态、花粉形态及解剖特征的变异。
建立了适于锡盟沙地榆的稳定的RAPD反应体系及反应程序,RAPD研究结果如下:14个随机引物在锡盟沙地榆和家榆中共检测到118个位点,平均每个引物扩增带数为8.4条,其中多态位点83个,占70.34%。选取了多态位点比率、Nei's多样性指数(h)和Shannon's多样性指数(I)来评价锡盟沙地榆与对照家榆居群的遗传多样性水平,结果表明锡盟沙地榆与家榆均具有较高水平的遗传多样性,且锡盟沙地榆遗传多样性水平明显高于家榆。在假设Hardy-Weinberg平衡时计算出锡盟沙地榆六个居群间的遗传分化系数Gst为0.3762,即在总的遗传变异中有37.62%存在于居群之间,居群内的遗传变异所占比例为62.38%,六个居群之间有相似的遗传多样性,但由于地理和距离的隔离,居群间存在有一定的遗传分化。根据遗传距离和聚类分析得出:锡盟沙地榆和对照家榆明显聚为两大类,表明经过漫长的进化,锡盟沙地榆产生并保留了较大的遗传变异。锡盟沙地榆及对照家榆各居群间的地理分布与其聚类结果存在着极大的相关性。
结合形态学、解剖学、孢粉学和RAPD研究结果,本文支持将锡盟沙地榆作为家榆一变种的结论。
The research objects are Ulmus pumila L.var.sabulosa which distribution on Hunshandake sandland and Ulmus pumila as comparision.The results of the morphologic,palynology,anatomy and RAPD indicate that:
There are same characteristics between Ulmus pumila L.var.sabulosa and U.s pumila in the shape characteristic, pollen shape characteristic and the structure of leaf, but also has differences. The results show that U. pumila L.var.sabulosa has occurred variation in the shape characteristic, pollen shape characteristic and the structure of leaf for acclimatize itself to entironment in the long process of evolvement.
118 reproducible loci are yielded based on 14 decamer primers from U.s pumila L.var.sabulosa and U. pumila,of which 83 are polymorphic .The average is 8.4 per decamer primers and the percentage of polymorphic loci equaled 70.34%. The percentage of polymorphic loci, Nei's (1973)gene diversity and Shannon's information index are choosed to estimate the genetic diversity of U. pumila L.var.sabulosa and U. pumila, the results show that U. pumila L.var.sabulosa and U. pumila has higher genetic diversity level, of which U. pumila L.var.sabulosa are higher than U. pumila distinctly. The coefficient of gene differentiation (Gst) based on supposing Hardy-Weinberg among populations is 0.3762, which reveal that 37.62% of the genetic variance existed among populations and 62.38% within populations. There were conform genetic diversity in the six populations, while there were definite gene differentiation for being obstructed. According to the genetic distance and cluster analysis (UPGMA), U. pumila L.var.sabulosa and U. pumila can be sorted into 2 different groups. The results demonstrate that U. pumila L.var.sabulosa has existenced and keeped biggest gene differentiation as endless evolution. There are pertinence between the geography distribution and cluster results of the populations U. pumila L.var.sabulosa and U. pumila.
The results demonstrate that U. pumila L.var. sabulosa can be regarded as a variety of U. pumila based on the results of the morphologic, palynology, anatomy and RAPD.
引文
1 马克平.试论生物多样性的概念,生物多样性[M].1993, 1(1) :20~22
2 陈灵芝.中国的生物多样性现状及其保护对策[M].科学出版社.1993, 1～3
3 McNeely J. A. et al. Conserving the World's Biological Diversity 1990,(生物多样性译从(一),中国科学技术出版社,1992).
4 马克平,钱迎倩.生物多样性保护及其研究进展,应用与环境生物学报[J]1998,4(1):95~99
5 董玉深,生物多样性及作物遗传多样性检测.作物品种资源[J],1995, 3:1~5
6 钱迎倩,马克平.生物多样性研究的几个国际热点.广西植物[J],1996,16 (4):295~299
7 钱迎倩,马克平.生物技术与生物安全.自然资源学报[J].1995, 10 (4) :322~331
8 王英杰.中国的生物多样性及其发展.科学对社会的影响[J].1995, 4:2~3
9 World Resources Institute(WRI)等.马克平等译.全球生物多样性策略.北京:中国标准出版社.1993
10 Solos P S,Soltis D E. Genetic variation in endemic and widespread plant species:exarnples from: Saxifragceae and polystichum, Aliso, 1991, 13:215~223
11 蒋志刚等.保护生物学(生物多样性研究丛书).杭州:浙江科学技术出版社.1997
12 沈浩,刘登义.遗传多样性的概述.生物学杂志,2001,16(3): 5～7
13 Templeton A R. Biodiversity at the genetic level: experiences form disparate microorganisms. In:Hawkks W D L. Biodiversity measurement and estimation. London: Chapman&Hall. 1996:140~141.
14 Weising K, Nybom H, Wolff K, Meyer W. DNA fingerprinting in plants and fungi. CRC Press, Inc., Bota Raton, 1995
15 Wright, S. The genetical structure of populations. Ann Eugen.1951,15:323～354
16 Brown A H. Isozymes,plant population genetic structure,genetic conservation. Theor Appl Genet.1978,52:145~157
17 Levins R. Theory of fitness in a heterogeneous environment(II).Developmental flexibility, niche selection. Amer Natur.1963, 97:75~90
18 Loveless M P, Hamrick J L. Ecological determinant of genetic structure in plant populations. Ann Rev Ecol Syst.1984, 15:65~95
19 Grant V. The evolutionary Process: A Critical Study of Evolutionary Theory(2nd ed). New York: Columbia University Press,1991
20 Hamrick J L. Isozymes ananysis of genetic structure of plant population:In Soltis P.(eds) Isozymes in Plant Biology. Washington: Dioscorides Press,1989, 87~105
21 Hamrick J L, Allard R W. Microgeographical variation in allozyme frequencies in Avena barbata. Proc Nat Sci., 1972,69:2100~2104
22 McNeilly T, Bradshaw A D. Evolutionary processes in population of copper tolerant Agrostis tenuis. Evolution,1968,22:108~118
23 Antonovics J,Bradshaw A D et al. Heavy metal tolerance in plants. Adv Ecol Res.,1971,7:1~58
24 Bradshaw A D.Evolution.Plant evolution in extreme environments.1n:Creed R ed. Ecological Genetics and Qxford: Blackwell
25 Barrett S C, Eckert C G. Variation, eolution of mating systems in seed plants.In:Kawano Sed.Bi-ological Approaches, Evolutionary Trends in Plants. London:Academic Press,1990:229~254
26 Mayr E. Animal Species, Evolution. Cambridge: Harvard University Press, 1963
27 Ehrlich P R,Raven P H. Differentiation of populations.Science,1969,165:1228~1232
28 Levin D A, Kerster H W. Gene flow in seed plants. Evol Biol., 1974,7:139~220
29 Merrell D J.黄瑞复等译.生态遗传学.北京:科学出版社,1991
30 Hamrick J L. Gene flow, distribution of genetic variation in plant populations. In: K. Urbanska(ed) Differentiation Patterns in Higher Plants. New York:Academic Press, 1987,53~67
31 Slatkin M. Rare alleles as indicators of gene flow. Evolution,1985,39:53~65
32 Wright S. Evolution mendelian population. Genetics, 1931,16:97
33 李难.进化论教程.北京:高等教育出版社.1999: 271
34 Wright, S. The genetical structure of populations. Ann Eugen., 1951,15:323~354
35 Welsh J ,Peterson C,Mcclelland M,et al. Genomic fingerprinting using arbitary 州 med PCRvand a matrix of pairwise combinations of primere[J].Nucleic Acids Research, 1990, 19:5275~5279
36 Williams J K,Kubelik A R,Livak K L, et al. DNA polymorphisms amplified by arbitrary primers are useful as genetic markers[J]. Nucleic Acids Research, 1990,18(22):6531~6535
37 Wu L,Lin H. IdenLifving huffalograss( Buchloe dactyloides (Nutt)Ebgekm) cultivar breeding lines using ramdom amplifide polymorphic DNARAPD markers [J].Journal of the American Society for Horticultural Science,1994,119(1):126~130
38 罗正荣,米森敬三,杨浦明.应用 PARD 技术进行柿种类和品种鉴定[J].日本园艺学会杂志,1995(64):535~541
39 傅荣昭,邵鹏柱,高文远.DNA 分子标记技术及其在药用植物研究上的应用前景[J].生物工程进展,1998, 18( 4) : 14~ 17
40 贾继增.分子标记种质资源鉴定和分子标记育种[J].中国农业科学,1996, 29( 4) : 1~10
41 Takimori N. Shinoda K. Kadotam N. et a .RAPD markers for confirmation of somatic hybrids in the dihaploed breeding of potato[J].Plant Cell Rep,1994,13:367~371
42 史水忠郭文武邓秀新.柑桔 RAPD 技术体系建立与体细胞杂种鉴定[J].园艺学报 1995,25(2): 15~110.
43 Xu Y-S. Clark M S. Pehu E. et al .Use of RAPD marker to screen somatic hybrids between Solanum cuberosum and S.brevidens[J].Nature,1993(322):275~279
44 史水忠郭文武邓秀新.柑桔 RAPD 技术体系建立与体细胞杂种鉴定[J].园艺学报 1995,25(2): 15~110
45 邹喻苹.RAPD 技术在检测遗传多样性中的应用[Z].北京: 中国科学院植物研究所系统与进化植物开放研究实验室,1994, 1~32
46 汪小荃,邹喻苹,张大明等.银杉遗传多样性的 RAPD 分析[J].中国科学(C 辑 1996.) , 26 (5): 436~441
47 李春香,杨群等.水杉自然居群遗传多样性的 RAPD 研究[J].中山大学学报(自然科学版),1999,38(1):59～63
48 李宽锐,黄敏仁等.1997.青杨的遗传分化.植物学报,39 (8 ): 753~758
49 梁红健.刘敏. 应用 RAPD 技术对 19 种中国兰花品种的分类学研究[J].园艺学报.1996, 23( 4) : 365~ 370.
50 汪小全,邹喻苹,张大明等. 1996. RAPD 应用于遗传多样性和系统学研究中的问题.杭物学报,38(12):954~962
51 乔爱民,刘佩瑛,雷建军.1998.芥菜 16 个变种的 RAPD 研究.植物学报,40 (10 ): 915~921
52 惠东威,庄炳吕,陈受宜.1996.RAPD 重建的大显属植物的亲缘关系.遗传学报,23 ( 6 ): 460~468
53 徐克学著.数量分类学[M].科学出版社,1994 .1~14
54 马骥,李俊祯,孔红.我国沙区 6 种篙属植物的种子微形态特征[J].中国沙漠,2002, 22~28
55 于海秋,徐克章等.土壤干旱对玉米叶片膜透性及叶绿体超微结构的影响[J].吉林农业大学学报,2002,(24) 3:17~23
56 敖成齐,陈功锡,张宏达.山茶属的叶表皮形态及其分类学意义[J].云南植物 2002, 24 (1) :68~74
57 蔡淑华.植物解剖学[M].台湾:国立编译馆
58 刘捷平,植物形态解剖学[M].北京:北京师范学院出版社,1991
59 郑湘如.植物解剖结构显微图谱[M].北京:农业出版社,1983
60 胡正海.结构植物学的现状和发展趋向的探[J].西北植物学报 2000,20 (6) : 899~906
61 魏岩,谭敦炎,尹林克.中国怪柳科植物叶解剖特征与分类关系的探讨[J].西北植物学报,1999, 19(1):113~118.
62 孙同兴,吴鸿,李秉滔,孙瑾,郑兴峰.中国假鹰爪属和皂帽花属叶的形态结构及其分类意义[J].植物分类学报,2002, 40(5):385~395
63 Adanson M.1763.Families des Plantes.2: 164.
64 宛涛,卫智军等.内蒙古草地现代植物花粉形态[M].北京:中国农业出版社,1999
65 谢成章编译.被子植物形态学[M].湖北:湖北科技出版社,1984: 178~ 187
66 高信曾.植物学(形态、解剖部分) [M].北京:人民教育出版社,1978: 149
67 蓝盛银等.植物花粉剥离观察扫描电镜图解[M].北京:科学出版社,1996: 5~10
68 李扬汉主编.植物学[M].上海:上海科学技术出版社,1995:I99~202
69 王开发等.孢粉学概论[M].北京:北京大学出版社,1983
70 张金谈等,现代花粉应用研究[M].北京;科学出版社,1990
71 蓝盛根,徐珍秀等. SEM Illustration of Strippde Pollen[M].北京:科学出版社, 1996.
72 张元明,潘伯荣等.中国柽柳科花粉形态研究及其分类意义的探讨[J].西北植物学报,2002,21(5):857~864
73 努沙力克,潘晓荣.我国西部白刺属及其近缘属的花粉形态与分类[J].干旱区研究,2003, (20) 20~23
74 张明理,田希娅.锦鸡儿属花粉形态及其分类意义[J].植物分类学报,1996, 34 (4) 397~409
75 李秀根,杨健.花粉形态数量化分析在中国梨属植物起源、演化和分类中的应用[J].果树学报,2002,19 (3):145~148
76 孙京田.松属植物花粉形态研究及其分类学意义[J].山东科学,2002, 15 (4): 35~40
77 C. A.斯特斯.植物分类学与生物系统学.北京:科学出版社,1986: 79
78 钱迎倩,马克平.生物多样性研究的原理与方法.北京:中国科学技术出版社,1994, 13~16
79 Grant, V. The Evolutionary Process: A Critical study of Evolutionary Theory.(2nded.) NewYork: Columbia University Press, 1991.
80 王牧兰.浑善达克沙地景观格局动态变化及其原因分析(硕士学位论文).内蒙古师范大学,2003.
81 马文红.浑善达克沙地的植被现状与沙漠化研究(硕士学位论文).内蒙古大学,2003.
82 杨淑宽,陈静生,郭蓄民.内蒙古小腾格里沙漠考察.治沙研究,北京:科学出版社 1962, 3: 144 ~ 151.
83 Zhong Decai&Qu Jianjun. Recent developmental trend and prediction of sand deserts in China. Journal of Arid Environments, 2003,53:317~329.
84 内蒙古自治区草原勘测设计院.内蒙锡林郭勒盟正蓝旗天然草场资源调查报告.1984.
85 马毓泉等. 内蒙古植物志(第二卷)[M]. 呼和浩特: 内蒙古人民出版社, 1990, 112~113.
86 中国科学院内蒙古宁夏考察队. 内蒙古植被[M]. 北京: 利学出版社, 1985, 785~786.
87 张红霞, 刘果厚, 崔秀萍. 干旱对浑善达克沙地榆叶片解剖结构的影响[J]. 植物研究, 2005, 25(1): 39~43.
88 李红丽, 董智, 王林和等. 浑善达克沙地榆树根系分努特征及生物量研究[J]. 干旱区资源与环境,2002, 16(4): 99~105.
89 刘建, 朱选伟等. 浑善达克沙地榆树疏林生态系统的空间异质性[J]. 环境科学, 2003, 24(4):35~45.
90 李永庚, 蒋高明等. 人为干扰对浑善达克沙地榆树疏林的影响[J]. 植物生态学报, 2003,27(6): 829~834.
91 李胜功,赵爱芬,常学礼.科努沁沙地植被演替的几个问题.中国沙漠,1997,1(17):25
92 郭建华,李玉珊,李晶华.白榆一新变种.植物研究,1988,8(1):107
93 马毓泉等. 内蒙古植物志(第二卷)[M]. 呼和浩特: 内蒙古人民出版社, 1990, 112~113.
94 刘果厚,贾宝丽.浑善达克沙地榆的遗传多样性研究[J].干旱区资源与环境.2003,17(5):123~128
95 王伏雄著.中国植物花粉形态(第二版)[M].科学出版社 1997,3~5
96 王战,中国森林带,农学,1936:41~66.
97 郑艳.安徽景天属 12 种植物的花粉形态研究[J].植物研究,1997,17(2) :15 8~161
98 辛益群,张玉龙等.中国榆属药粉形态研究及其分类意义[J].植物学报,1993,35(2):91~95
99 L. A. Kuprianova.论花粉和袍子的进化阶段[J].植物学通报,1974, 28~34
100 Wang Y.Contrasts and observations of dissecting construction of three drought resistant plant leaf slices.Sichuan Forest Science,2003,24(1):64~47
101 Chen Y M.Chen H B, Chen G J.et al. A study on leaf structure and drought-resistanrt of banana. Chinese Journal of'Tropical Agriculture, 2001,92( 4): 14~16.
102 Li G Y,Gao G X, Lu Y L, et al. Observation on the morphology and anatomical structure of three bushes. Research of Soil andl Water Conservation, 1995,2( 2):141~145.
103 Li Y H. Botany. The Second Press. Shanghai: Shanghai Science and TechnologyPress. 1995.
104 Li Z L. Morphology and structure of drought vegetation. Biology Journal, 1981(4):9~12.
105 Zhou G L, Da L X T,An Z X, et al. Study on the morphology of leaf epidemis and comparative anatomy of leaf of Atriplex in Xinjiamg. Arid Zone Research. 1995,12( 3):34~37.
106 邹喻苹,葛颂,王晓东等.系统与进化植物学中的分子标记[M].科学出版社,2002, 161~167
107 汪小泉,邹喻苹,张大明等.银杉遗传多样性的 RAPD 分析[J].中国科学(C 辑 1996.26 (5):436~441
108 SHENG Yan, ZHENG Wei-Hong, PEI Ke-Quanl, MA Ke-Pingl. Population Genetic Structure of a Dominant Desert Tree, Haloxylon ammodendron (Chenopodiaceae), in the Southeast Gurbantunggut Desert Detected by RAPD and ISSR Markers [J].植物学报,2004, 46 (6): 675~681
109 祖元刚,孙梅康乐主编.生态适应与生态进化的分子机理[M].高等教育出版社,施普林格出版社,2000 ,193~201
110 府宇雷,钱吉,张美云等.金华地区野生大豆小种群的分子生态学研究[J].复旦学报(自然科学版),1999, 38(5):584~586
111 武耀廷,张天真,殷剑美.利用分子标记和形态学性状检测的陆地棉栽培品种遗传多样性[J] .遗传学报,2001, 28(11):1040~1050
112 王经源,黄儒珠.用于 DNA 提取的刺桫椤叶片组织保存方法研究[J].2002,18(1):82~85
113 钱伟,葛颂.居群遗传结构研究中显性标记数据分析方法初探[J].遗传学报.2001, 28 (3):244～255.
114 Hamrick J.L.Godt M. J. W.Allozyme Diversity in Plant Species. In A.H. D. Brown, M.T. Clegg, A. L.Kahler&B.S.Wier(eds), Plant Population Genetics,Breeding and Genetic Resources. Sinauer,Associates,Inc.Sunderland, MA. pp.1989,43~63.
115 李巧霞.青藏高原东部篙草属五种植物的遗传多样性研究[D].西北师范大学硕士毕业论文,2005,34~37
116 张林静.新疆阜康荒漠植物群落物种多样性与梭梭的遗传多样性研究[D].北京林业大学硕士毕业论文,2003 , 45~51
117 郭柯 , 刘海江 . 浑善达克沙地四种生境中榆树天然更新幼苗发育的比较 [J]. 生态学报,2004,24(9):2024~2028
118 刘占林,赵桂仿.种群遗传学原理及其在珍惜濒危植物保护中的应用[J].生物多样性,1999,7(4):340~346
119 Hamrick J.L.,Godt M. J. W.Allozyme Diversity in Plant Species. In A.H. D. Brown, M.T. Clegg, A. L.Kahler&B.S。Wier(eds), Plant Population Genetics, Breeding and Genetic Resources. Sinauer, Associates, Inc.Sunderland, MA. pp.1989,43~63.
120 Loveless M P, Hamrick J L. Ecological determinant of genetic structure in plant populations. Ann Rev Ecol Syst., 1984, 15:65~95
121 Yam L. L He X, et al. Ecological anatomy of nine priority species in A La Sam Area. Journal of Inner Mongolia Agriculture University,2000, 21(3):65~71.
122 张日清,谭晓风,吕芳德.林木 RAPD 标记技术研究进展[J],吉首大学学报,2001,22(3):16~22.
123 谭晓风,胡芳名,张启发.银杏主要栽培品种的分子鉴别[J].中南林学院学报,1998,18(3):3~10.
124 Thorp, J P. The molecular dock hypothesis:biochemical evolution, genetic differentiation, and systematica[J]. Annual Review lf Ecology Systematic, 1982, 13( I): 139~168.
2 陈灵芝.中国的生物多样性现状及其保护对策[M].科学出版社.1993, 1～3
3 McNeely J. A. et al. Conserving the World's Biological Diversity 1990,(生物多样性译从(一),中国科学技术出版社,1992).
4 马克平,钱迎倩.生物多样性保护及其研究进展,应用与环境生物学报[J]1998,4(1):95~99
5 董玉深,生物多样性及作物遗传多样性检测.作物品种资源[J],1995, 3:1~5
6 钱迎倩,马克平.生物多样性研究的几个国际热点.广西植物[J],1996,16 (4):295~299
7 钱迎倩,马克平.生物技术与生物安全.自然资源学报[J].1995, 10 (4) :322~331
8 王英杰.中国的生物多样性及其发展.科学对社会的影响[J].1995, 4:2~3
9 World Resources Institute(WRI)等.马克平等译.全球生物多样性策略.北京:中国标准出版社.1993
10 Solos P S,Soltis D E. Genetic variation in endemic and widespread plant species:exarnples from: Saxifragceae and polystichum, Aliso, 1991, 13:215~223
11 蒋志刚等.保护生物学(生物多样性研究丛书).杭州:浙江科学技术出版社.1997
12 沈浩,刘登义.遗传多样性的概述.生物学杂志,2001,16(3): 5～7
13 Templeton A R. Biodiversity at the genetic level: experiences form disparate microorganisms. In:Hawkks W D L. Biodiversity measurement and estimation. London: Chapman&Hall. 1996:140~141.
14 Weising K, Nybom H, Wolff K, Meyer W. DNA fingerprinting in plants and fungi. CRC Press, Inc., Bota Raton, 1995
15 Wright, S. The genetical structure of populations. Ann Eugen.1951,15:323～354
16 Brown A H. Isozymes,plant population genetic structure,genetic conservation. Theor Appl Genet.1978,52:145~157
17 Levins R. Theory of fitness in a heterogeneous environment(II).Developmental flexibility, niche selection. Amer Natur.1963, 97:75~90
18 Loveless M P, Hamrick J L. Ecological determinant of genetic structure in plant populations. Ann Rev Ecol Syst.1984, 15:65~95
19 Grant V. The evolutionary Process: A Critical Study of Evolutionary Theory(2nd ed). New York: Columbia University Press,1991
20 Hamrick J L. Isozymes ananysis of genetic structure of plant population:In Soltis P.(eds) Isozymes in Plant Biology. Washington: Dioscorides Press,1989, 87~105
21 Hamrick J L, Allard R W. Microgeographical variation in allozyme frequencies in Avena barbata. Proc Nat Sci., 1972,69:2100~2104
22 McNeilly T, Bradshaw A D. Evolutionary processes in population of copper tolerant Agrostis tenuis. Evolution,1968,22:108~118
23 Antonovics J,Bradshaw A D et al. Heavy metal tolerance in plants. Adv Ecol Res.,1971,7:1~58
24 Bradshaw A D.Evolution.Plant evolution in extreme environments.1n:Creed R ed. Ecological Genetics and Qxford: Blackwell
25 Barrett S C, Eckert C G. Variation, eolution of mating systems in seed plants.In:Kawano Sed.Bi-ological Approaches, Evolutionary Trends in Plants. London:Academic Press,1990:229~254
26 Mayr E. Animal Species, Evolution. Cambridge: Harvard University Press, 1963
27 Ehrlich P R,Raven P H. Differentiation of populations.Science,1969,165:1228~1232
28 Levin D A, Kerster H W. Gene flow in seed plants. Evol Biol., 1974,7:139~220
29 Merrell D J.黄瑞复等译.生态遗传学.北京:科学出版社,1991
30 Hamrick J L. Gene flow, distribution of genetic variation in plant populations. In: K. Urbanska(ed) Differentiation Patterns in Higher Plants. New York:Academic Press, 1987,53~67
31 Slatkin M. Rare alleles as indicators of gene flow. Evolution,1985,39:53~65
32 Wright S. Evolution mendelian population. Genetics, 1931,16:97
33 李难.进化论教程.北京:高等教育出版社.1999: 271
34 Wright, S. The genetical structure of populations. Ann Eugen., 1951,15:323~354
35 Welsh J ,Peterson C,Mcclelland M,et al. Genomic fingerprinting using arbitary 州 med PCRvand a matrix of pairwise combinations of primere[J].Nucleic Acids Research, 1990, 19:5275~5279
36 Williams J K,Kubelik A R,Livak K L, et al. DNA polymorphisms amplified by arbitrary primers are useful as genetic markers[J]. Nucleic Acids Research, 1990,18(22):6531~6535
37 Wu L,Lin H. IdenLifving huffalograss( Buchloe dactyloides (Nutt)Ebgekm) cultivar breeding lines using ramdom amplifide polymorphic DNARAPD markers [J].Journal of the American Society for Horticultural Science,1994,119(1):126~130
38 罗正荣,米森敬三,杨浦明.应用 PARD 技术进行柿种类和品种鉴定[J].日本园艺学会杂志,1995(64):535~541
39 傅荣昭,邵鹏柱,高文远.DNA 分子标记技术及其在药用植物研究上的应用前景[J].生物工程进展,1998, 18( 4) : 14~ 17
40 贾继增.分子标记种质资源鉴定和分子标记育种[J].中国农业科学,1996, 29( 4) : 1~10
41 Takimori N. Shinoda K. Kadotam N. et a .RAPD markers for confirmation of somatic hybrids in the dihaploed breeding of potato[J].Plant Cell Rep,1994,13:367~371
42 史水忠郭文武邓秀新.柑桔 RAPD 技术体系建立与体细胞杂种鉴定[J].园艺学报 1995,25(2): 15~110.
43 Xu Y-S. Clark M S. Pehu E. et al .Use of RAPD marker to screen somatic hybrids between Solanum cuberosum and S.brevidens[J].Nature,1993(322):275~279
44 史水忠郭文武邓秀新.柑桔 RAPD 技术体系建立与体细胞杂种鉴定[J].园艺学报 1995,25(2): 15~110
45 邹喻苹.RAPD 技术在检测遗传多样性中的应用[Z].北京: 中国科学院植物研究所系统与进化植物开放研究实验室,1994, 1~32
46 汪小荃,邹喻苹,张大明等.银杉遗传多样性的 RAPD 分析[J].中国科学(C 辑 1996.) , 26 (5): 436~441
47 李春香,杨群等.水杉自然居群遗传多样性的 RAPD 研究[J].中山大学学报(自然科学版),1999,38(1):59～63
48 李宽锐,黄敏仁等.1997.青杨的遗传分化.植物学报,39 (8 ): 753~758
49 梁红健.刘敏. 应用 RAPD 技术对 19 种中国兰花品种的分类学研究[J].园艺学报.1996, 23( 4) : 365~ 370.
50 汪小全,邹喻苹,张大明等. 1996. RAPD 应用于遗传多样性和系统学研究中的问题.杭物学报,38(12):954~962
51 乔爱民,刘佩瑛,雷建军.1998.芥菜 16 个变种的 RAPD 研究.植物学报,40 (10 ): 915~921
52 惠东威,庄炳吕,陈受宜.1996.RAPD 重建的大显属植物的亲缘关系.遗传学报,23 ( 6 ): 460~468
53 徐克学著.数量分类学[M].科学出版社,1994 .1~14
54 马骥,李俊祯,孔红.我国沙区 6 种篙属植物的种子微形态特征[J].中国沙漠,2002, 22~28
55 于海秋,徐克章等.土壤干旱对玉米叶片膜透性及叶绿体超微结构的影响[J].吉林农业大学学报,2002,(24) 3:17~23
56 敖成齐,陈功锡,张宏达.山茶属的叶表皮形态及其分类学意义[J].云南植物 2002, 24 (1) :68~74
57 蔡淑华.植物解剖学[M].台湾:国立编译馆
58 刘捷平,植物形态解剖学[M].北京:北京师范学院出版社,1991
59 郑湘如.植物解剖结构显微图谱[M].北京:农业出版社,1983
60 胡正海.结构植物学的现状和发展趋向的探[J].西北植物学报 2000,20 (6) : 899~906
61 魏岩,谭敦炎,尹林克.中国怪柳科植物叶解剖特征与分类关系的探讨[J].西北植物学报,1999, 19(1):113~118.
62 孙同兴,吴鸿,李秉滔,孙瑾,郑兴峰.中国假鹰爪属和皂帽花属叶的形态结构及其分类意义[J].植物分类学报,2002, 40(5):385~395
63 Adanson M.1763.Families des Plantes.2: 164.
64 宛涛,卫智军等.内蒙古草地现代植物花粉形态[M].北京:中国农业出版社,1999
65 谢成章编译.被子植物形态学[M].湖北:湖北科技出版社,1984: 178~ 187
66 高信曾.植物学(形态、解剖部分) [M].北京:人民教育出版社,1978: 149
67 蓝盛银等.植物花粉剥离观察扫描电镜图解[M].北京:科学出版社,1996: 5~10
68 李扬汉主编.植物学[M].上海:上海科学技术出版社,1995:I99~202
69 王开发等.孢粉学概论[M].北京:北京大学出版社,1983
70 张金谈等,现代花粉应用研究[M].北京;科学出版社,1990
71 蓝盛根,徐珍秀等. SEM Illustration of Strippde Pollen[M].北京:科学出版社, 1996.
72 张元明,潘伯荣等.中国柽柳科花粉形态研究及其分类意义的探讨[J].西北植物学报,2002,21(5):857~864
73 努沙力克,潘晓荣.我国西部白刺属及其近缘属的花粉形态与分类[J].干旱区研究,2003, (20) 20~23
74 张明理,田希娅.锦鸡儿属花粉形态及其分类意义[J].植物分类学报,1996, 34 (4) 397~409
75 李秀根,杨健.花粉形态数量化分析在中国梨属植物起源、演化和分类中的应用[J].果树学报,2002,19 (3):145~148
76 孙京田.松属植物花粉形态研究及其分类学意义[J].山东科学,2002, 15 (4): 35~40
77 C. A.斯特斯.植物分类学与生物系统学.北京:科学出版社,1986: 79
78 钱迎倩,马克平.生物多样性研究的原理与方法.北京:中国科学技术出版社,1994, 13~16
79 Grant, V. The Evolutionary Process: A Critical study of Evolutionary Theory.(2nded.) NewYork: Columbia University Press, 1991.
80 王牧兰.浑善达克沙地景观格局动态变化及其原因分析(硕士学位论文).内蒙古师范大学,2003.
81 马文红.浑善达克沙地的植被现状与沙漠化研究(硕士学位论文).内蒙古大学,2003.
82 杨淑宽,陈静生,郭蓄民.内蒙古小腾格里沙漠考察.治沙研究,北京:科学出版社 1962, 3: 144 ~ 151.
83 Zhong Decai&Qu Jianjun. Recent developmental trend and prediction of sand deserts in China. Journal of Arid Environments, 2003,53:317~329.
84 内蒙古自治区草原勘测设计院.内蒙锡林郭勒盟正蓝旗天然草场资源调查报告.1984.
85 马毓泉等. 内蒙古植物志(第二卷)[M]. 呼和浩特: 内蒙古人民出版社, 1990, 112~113.
86 中国科学院内蒙古宁夏考察队. 内蒙古植被[M]. 北京: 利学出版社, 1985, 785~786.
87 张红霞, 刘果厚, 崔秀萍. 干旱对浑善达克沙地榆叶片解剖结构的影响[J]. 植物研究, 2005, 25(1): 39~43.
88 李红丽, 董智, 王林和等. 浑善达克沙地榆树根系分努特征及生物量研究[J]. 干旱区资源与环境,2002, 16(4): 99~105.
89 刘建, 朱选伟等. 浑善达克沙地榆树疏林生态系统的空间异质性[J]. 环境科学, 2003, 24(4):35~45.
90 李永庚, 蒋高明等. 人为干扰对浑善达克沙地榆树疏林的影响[J]. 植物生态学报, 2003,27(6): 829~834.
91 李胜功,赵爱芬,常学礼.科努沁沙地植被演替的几个问题.中国沙漠,1997,1(17):25
92 郭建华,李玉珊,李晶华.白榆一新变种.植物研究,1988,8(1):107
93 马毓泉等. 内蒙古植物志(第二卷)[M]. 呼和浩特: 内蒙古人民出版社, 1990, 112~113.
94 刘果厚,贾宝丽.浑善达克沙地榆的遗传多样性研究[J].干旱区资源与环境.2003,17(5):123~128
95 王伏雄著.中国植物花粉形态(第二版)[M].科学出版社 1997,3~5
96 王战,中国森林带,农学,1936:41~66.
97 郑艳.安徽景天属 12 种植物的花粉形态研究[J].植物研究,1997,17(2) :15 8~161
98 辛益群,张玉龙等.中国榆属药粉形态研究及其分类意义[J].植物学报,1993,35(2):91~95
99 L. A. Kuprianova.论花粉和袍子的进化阶段[J].植物学通报,1974, 28~34
100 Wang Y.Contrasts and observations of dissecting construction of three drought resistant plant leaf slices.Sichuan Forest Science,2003,24(1):64~47
101 Chen Y M.Chen H B, Chen G J.et al. A study on leaf structure and drought-resistanrt of banana. Chinese Journal of'Tropical Agriculture, 2001,92( 4): 14~16.
102 Li G Y,Gao G X, Lu Y L, et al. Observation on the morphology and anatomical structure of three bushes. Research of Soil andl Water Conservation, 1995,2( 2):141~145.
103 Li Y H. Botany. The Second Press. Shanghai: Shanghai Science and TechnologyPress. 1995.
104 Li Z L. Morphology and structure of drought vegetation. Biology Journal, 1981(4):9~12.
105 Zhou G L, Da L X T,An Z X, et al. Study on the morphology of leaf epidemis and comparative anatomy of leaf of Atriplex in Xinjiamg. Arid Zone Research. 1995,12( 3):34~37.
106 邹喻苹,葛颂,王晓东等.系统与进化植物学中的分子标记[M].科学出版社,2002, 161~167
107 汪小泉,邹喻苹,张大明等.银杉遗传多样性的 RAPD 分析[J].中国科学(C 辑 1996.26 (5):436~441
108 SHENG Yan, ZHENG Wei-Hong, PEI Ke-Quanl, MA Ke-Pingl. Population Genetic Structure of a Dominant Desert Tree, Haloxylon ammodendron (Chenopodiaceae), in the Southeast Gurbantunggut Desert Detected by RAPD and ISSR Markers [J].植物学报,2004, 46 (6): 675~681
109 祖元刚,孙梅康乐主编.生态适应与生态进化的分子机理[M].高等教育出版社,施普林格出版社,2000 ,193~201
110 府宇雷,钱吉,张美云等.金华地区野生大豆小种群的分子生态学研究[J].复旦学报(自然科学版),1999, 38(5):584~586
111 武耀廷,张天真,殷剑美.利用分子标记和形态学性状检测的陆地棉栽培品种遗传多样性[J] .遗传学报,2001, 28(11):1040~1050
112 王经源,黄儒珠.用于 DNA 提取的刺桫椤叶片组织保存方法研究[J].2002,18(1):82~85
113 钱伟,葛颂.居群遗传结构研究中显性标记数据分析方法初探[J].遗传学报.2001, 28 (3):244～255.
114 Hamrick J.L.Godt M. J. W.Allozyme Diversity in Plant Species. In A.H. D. Brown, M.T. Clegg, A. L.Kahler&B.S.Wier(eds), Plant Population Genetics,Breeding and Genetic Resources. Sinauer,Associates,Inc.Sunderland, MA. pp.1989,43~63.
115 李巧霞.青藏高原东部篙草属五种植物的遗传多样性研究[D].西北师范大学硕士毕业论文,2005,34~37
116 张林静.新疆阜康荒漠植物群落物种多样性与梭梭的遗传多样性研究[D].北京林业大学硕士毕业论文,2003 , 45~51
117 郭柯 , 刘海江 . 浑善达克沙地四种生境中榆树天然更新幼苗发育的比较 [J]. 生态学报,2004,24(9):2024~2028
118 刘占林,赵桂仿.种群遗传学原理及其在珍惜濒危植物保护中的应用[J].生物多样性,1999,7(4):340~346
119 Hamrick J.L.,Godt M. J. W.Allozyme Diversity in Plant Species. In A.H. D. Brown, M.T. Clegg, A. L.Kahler&B.S。Wier(eds), Plant Population Genetics, Breeding and Genetic Resources. Sinauer, Associates, Inc.Sunderland, MA. pp.1989,43~63.
120 Loveless M P, Hamrick J L. Ecological determinant of genetic structure in plant populations. Ann Rev Ecol Syst., 1984, 15:65~95
121 Yam L. L He X, et al. Ecological anatomy of nine priority species in A La Sam Area. Journal of Inner Mongolia Agriculture University,2000, 21(3):65~71.
122 张日清,谭晓风,吕芳德.林木 RAPD 标记技术研究进展[J],吉首大学学报,2001,22(3):16~22.
123 谭晓风,胡芳名,张启发.银杏主要栽培品种的分子鉴别[J].中南林学院学报,1998,18(3):3~10.
124 Thorp, J P. The molecular dock hypothesis:biochemical evolution, genetic differentiation, and systematica[J]. Annual Review lf Ecology Systematic, 1982, 13( I): 139~168.