红桤木遗传多样性研究
|
摘要
生物多样性是世界的自然财富,是人类赖以生存的基础,而遗传多样性又是生物多样性的基础,是遗传改良的源泉。红桤木是从北美引进的速生木本非豆科固氮树种,经济价值高,造林或栽培历史悠久,形成了优良的品系和相应的配套技术,适应性强,在特殊立地上可以造林,苗木生长量比国内同类树种或品种提高10%,红桤木成功引入中国,对造林及家具、制浆造纸等行业具有重要意义。因此,研究红桤木的遗传多样性具有重要意义。
本文选取了在六合基地和下蜀林场两个栽培居群60个样本进行多样性研究。运用方差分析、UPGMA聚类和RAPD标记等方法,从表型和分子水平等系统地研究了红桤木群体内、群体间的变异,进而为红桤木育种和遗传改良奠定基础。
1.通过对2个栽培居群60个样本进行表型特征分析,结果表明红桤木具有丰富的变异,各部位平均变异系数有明显差异,排序为：叶宽>叶长>叶脉>叶长宽比。
2.四个性状表型分化系数变异幅度为0.27%～24.68%。群体间平均表型分化系数为7.97%,群体内平均表型分化系数为92.03%,群体内变异为其主要的变异来源。
3.从45条随机引物中筛选出11条扩增谱带清晰且稳定的引物,用这些引物对60份种质进行扩增,扩增出位点81个,多态性位点81个,多态性比例为100%,具有较高的遗传多样性(Shannon多样性信息指数(Ⅰ)为0.509,Nei’s指数为0.3425),遗传分化Gst平均值为0.062,群体间基因多样性占总基因多样性的6.2%,群体内的基因多样性占总基因多样性的93.8%。红桤木的遗传多样性主要来自群体内部。
4.基于红桤木表型和RAPD分子标记的聚类分析,本文认为：从种质保存的角度,六合居群的33、37、40、41、42、43、44、45、46、48、55、57、58和59号种质,下蜀居群的1、2、3、4、9、10、11、12、16、17、19、20、21、22、23、24、25、26和27号种质是需要保存的种质,用作育种材料。
Bio-diversity is the natural wealth of the world and the foundation of human survial. The genetic diversity is the basis of bio-diversity and the source of genetic improvement. Alnus rubra is a fast-growing nitrogen-fixing species introduced from North America, high economic value, afforestation or cultivation has a long history,the formation of fine lines and corresponding supporting technology, adaptable, can be planted in special legislation on the ground, seedling growth compared with the similar species or varieties increased by 10%.The successful of Alnus rubra's introduction is important for forestation, furniture and paper making. Therefore, the research of Alnus rubra's genetic diversity has important meaning.
In this paper,60 individual samples were collected from two cultivated populations located in LiuHe and XiaShu. From the phenotypic and molecular levels to study systematically the variation patterns within and among of Alnus rubra by using variance analysis, UPGMA clustering and RAPD markers, It will lay the foundation of Alnus rubra's breeding and genetic improvement.
1. Analysis of phenotypic characteristics of 60 samples from two cultivated populations showed that Alnus rubra had abundant variations. The average coefficients of variation were significantly different in different parts. From highest to lowest:leaf width> leaf length> leaf venation> leaf ratio.
2. The variation range of four phenotypic differentiation coefficients was 0.27%～24.68%. The average coefficient of phenotypic differentiation among populations was 7.97%, while the average coefficient of phenotypic differentiation within groups was 92.03%, so the variation within groups was the main source of variation.
3.45 random primers were selected from Invitrogen Corporation for the amplification reaction in the experiment.11 of which showed highly efficient amplification and were used for 60 germplasm in amplification test. The bands were clear and more polymorphic. All the bands were polymorphic in 81 loci and the ratio of polymorphic was 100%. Genetic diversity was abundant that judged from the average value of Shannon index (0.509), Nei index (0.3425). Gst index (0.062). Genetic diversity within populations possessed 93.8% and that among populations possessed 6.2% of the total. We can conclude that genetic diversity of Alnus rubra mainly from within populations.
4. Based clustering analysis of phenotype and molecular makers of Alnus rubra, this paper suggested that the germplasm of No.33,37,40,41,42,43,44,45,46,48,55,57,58 and 59 from LiuHe population, as well as those of No 1,2,3,4,9,10,11,12,16,17,19,20,21,22,23, 24,25,26 and 27 from XiaShu population,were need to preserve as breeding material.,
本文选取了在六合基地和下蜀林场两个栽培居群60个样本进行多样性研究。运用方差分析、UPGMA聚类和RAPD标记等方法,从表型和分子水平等系统地研究了红桤木群体内、群体间的变异,进而为红桤木育种和遗传改良奠定基础。
1.通过对2个栽培居群60个样本进行表型特征分析,结果表明红桤木具有丰富的变异,各部位平均变异系数有明显差异,排序为：叶宽>叶长>叶脉>叶长宽比。
2.四个性状表型分化系数变异幅度为0.27%～24.68%。群体间平均表型分化系数为7.97%,群体内平均表型分化系数为92.03%,群体内变异为其主要的变异来源。
3.从45条随机引物中筛选出11条扩增谱带清晰且稳定的引物,用这些引物对60份种质进行扩增,扩增出位点81个,多态性位点81个,多态性比例为100%,具有较高的遗传多样性(Shannon多样性信息指数(Ⅰ)为0.509,Nei’s指数为0.3425),遗传分化Gst平均值为0.062,群体间基因多样性占总基因多样性的6.2%,群体内的基因多样性占总基因多样性的93.8%。红桤木的遗传多样性主要来自群体内部。
4.基于红桤木表型和RAPD分子标记的聚类分析,本文认为：从种质保存的角度,六合居群的33、37、40、41、42、43、44、45、46、48、55、57、58和59号种质,下蜀居群的1、2、3、4、9、10、11、12、16、17、19、20、21、22、23、24、25、26和27号种质是需要保存的种质,用作育种材料。
Bio-diversity is the natural wealth of the world and the foundation of human survial. The genetic diversity is the basis of bio-diversity and the source of genetic improvement. Alnus rubra is a fast-growing nitrogen-fixing species introduced from North America, high economic value, afforestation or cultivation has a long history,the formation of fine lines and corresponding supporting technology, adaptable, can be planted in special legislation on the ground, seedling growth compared with the similar species or varieties increased by 10%.The successful of Alnus rubra's introduction is important for forestation, furniture and paper making. Therefore, the research of Alnus rubra's genetic diversity has important meaning.
In this paper,60 individual samples were collected from two cultivated populations located in LiuHe and XiaShu. From the phenotypic and molecular levels to study systematically the variation patterns within and among of Alnus rubra by using variance analysis, UPGMA clustering and RAPD markers, It will lay the foundation of Alnus rubra's breeding and genetic improvement.
1. Analysis of phenotypic characteristics of 60 samples from two cultivated populations showed that Alnus rubra had abundant variations. The average coefficients of variation were significantly different in different parts. From highest to lowest:leaf width> leaf length> leaf venation> leaf ratio.
2. The variation range of four phenotypic differentiation coefficients was 0.27%～24.68%. The average coefficient of phenotypic differentiation among populations was 7.97%, while the average coefficient of phenotypic differentiation within groups was 92.03%, so the variation within groups was the main source of variation.
3.45 random primers were selected from Invitrogen Corporation for the amplification reaction in the experiment.11 of which showed highly efficient amplification and were used for 60 germplasm in amplification test. The bands were clear and more polymorphic. All the bands were polymorphic in 81 loci and the ratio of polymorphic was 100%. Genetic diversity was abundant that judged from the average value of Shannon index (0.509), Nei index (0.3425). Gst index (0.062). Genetic diversity within populations possessed 93.8% and that among populations possessed 6.2% of the total. We can conclude that genetic diversity of Alnus rubra mainly from within populations.
4. Based clustering analysis of phenotype and molecular makers of Alnus rubra, this paper suggested that the germplasm of No.33,37,40,41,42,43,44,45,46,48,55,57,58 and 59 from LiuHe population, as well as those of No 1,2,3,4,9,10,11,12,16,17,19,20,21,22,23, 24,25,26 and 27 from XiaShu population,were need to preserve as breeding material.,
引文
蔡从利,王建波,朱英国,等.山羊草属二倍体物种亲缘关系的RAPD分析[J].遗传,2001,23(3)：229
曹后男,高光出.桃属植物随机扩增多态性DNA分析的最适反应条件[J].延边大学农学学报,1999,21(4)：249-253.
陈益泰,李桂,王惠雄.桤木自然分布区内表型变异的研究[J].林业科学研究,1999,12(4)：
陈之端,路安民.桦木科植物的起源和早期演化[J].中国科学院研究生院学报,1995,12(2)：200-205.
陈之端,路安民.桦木科植物的系统发育和演化[J].中国科学院院刊,2001,(3)：188-191.
陈之端.桦木科植物的系统发育与地理分布(续)[J].植物分类学报,1994,32(2)：101-153
陈志伟,官华忠,吴为人,等.稻瘟病抗性基因Pi-1连锁SSR标记的筛选和应用.福建农林大学学报(自然科学版),2005,34(1)：74-77.
成俊卿.木材学[M].北京：中国林业出版社,1985.
崔成日,徐启江,崔崇士,等.洋葱RAPD遗传多样性分析[J].园艺学报,2006,33(4)：863-865.
单雪,王秀利,仇雪梅,等.分子标记及其在海洋动物遗传研究中的应用[J].生物技术通讯,2005,16(4)：463-466.
葛颂,王明庥,陈岳武.用同工酶研究马尾松群体的遗传结构[J]林业科学,1988,24(4)：399-409.
辜云杰,王启和,罗建勋.四川桤木天然群体果实表型多样性研究[J].四川林业科技,2009,30(2)：19-22.
顾万军,刘和林,岳德智,等.桤木优树子代苗期性状变异的研究[J].四川林业科技,1995,16(3)：52-57.
韩斌,巩合德.哀牢山桤木次生林根瘤固氮能力研究[J].安徽农业研究,2009,37(17)：8272-8273,8283.
黄福平,梁月荣,陆建良,等.应用RAPD和ISSR分子标记构建茶树回交1代部分遗传图谱[J].茶叶科学,2006,26(3)：171-176.
黄志宏,黄维南.豆科茎瘤植物的结瘤、固氮和应用[J].亚热带植物通讯,1996,25(2)：40-46.
季天祜.第二林业与林纸结合[M].北京：中国轻工业出版社,1997.
季维智,宿兵.遗传多样性的原理与方法[M].杭州：浙江科学技术出版社,1999.
贾继增.分子标记种质资源鉴定和分子育种[J].中国农业科学,1996,29(4)：1-10.
姜建国,严媛,宋冬林.盐藻Psy基因保守序列的克隆及同源性分析[J].华南理工大学学报：自然科学版,2004,32(10)：76-79.
姜静,杨传平,刘桂丰,等.应用RAPD技术对东北地区白桦种源遗传变异的分析[J].东北林业大学学报,2001,29(2)：30-34.
蒋建德,赵振英,朱宝琴.非豆科树木根瘤内生菌的分离[J].林业科学，1987,23(1)：85-89.
李斌,顾万春,卢宝铭.白皮松天然群体种实性状表型多样性研究[J].生物多样性,2002,10(2)：181-187.
李洁,熊智.云南尼泊尔桤木遗传多样性研究[J].浙江林学院学报,2008,25(1)：16-21.
梁月荣,田中村一,武田善行.应用RAPD分子标记分析“晚绿”品种的杂交亲本[J].茶树科学,2000,20(1)：22.
林清洪,黄维南.福建省主要固氮树木VA菌根真菌的分离鉴定[J].亚热带植物讯,1999,28(1)：65.
刘士平,李信,汪朝阳,等.利用分子标记辅助改良珍汕97的稻瘟病抗性[J].植物学报,2003,45(11)：1346-1350.
刘延吉,耿书,田晓艳.辽宁地区五种野生软枣猕猴桃RAPD遗传多样性分析[J].北方园艺.2010,(18)：130-132
吕梅.红桤木生理生态、固氮活性及根瘤放线菌研究[D].南京林业大学学位论文,2008,3-24.
罗建勋.云杉天然群体遗传多样性研究[D].中国林业科学研究院,2004.
罗美娟.短枝木麻黄种源群体遗传多样性与遗传变异规律研究[D].福州：福建农林大学图书馆,2005.
M. Lalonde.桤木根瘤共生固氮研究技术与观测[J].亚热带植物通讯,1990,1：59-62.
潘建蓄,陈启锋,黄世贞.诱导固氮菌与稻苗结瘤共生的研究[J].福建农业大学学报,1998,27(4)：405-410.
潘远智,庞博,孙振元,等.雅江报春天然居群RAPD遗传多样性[J].四川农业大学学报.2008.26(1)：28-31.
庞广昌,姜冬梅.群体遗传多样性和数据分析[J].林业科学,1995,31(6)：543-550.
任旭琴.遗传多样性及其研究方法[J].淮阴工学学报,2002,11(5)：6-8.
沈向,郭卫东,吴燕民,等.杏43个品种资源的RAPD分类[J].园艺学报,2000,27(1)：55-56.
税珺,黄少伟,陈炳铨.火炬松原生种源和引种群体RAPD遗传多样性[J].华南农业大学学报,2005,26(3)：74-76,81.
宋丛文,张新叶,胡兴宜.秃杉种内遗传多样性的RAPD分析[J].湖北林业科技,2004(4)：1-4.
汪小全,邹喻苹,张大明,等.RAPD应用于遗传多样性和系统学研究中的问题[J].植物学报,1996,38(12)：954-962.
王晨光,阮继生,石彦林.木麻黄根瘤中分离的四株弗兰克氏菌的比较研究[J].微生物学通报,1993,20(3)：80-83.
王璐璐.贵州省白菜型油菜的RAPD遗传多样性分析[J].贵州农业科学,2009,37(10)：5-7.
王祖良,丁丽霞,赵明水,等.濒危植物天目铁木遗传多样性的RAPD分析[J].浙江林学院学报,2008,25(3)：304-308.
吴捷,陈华癸,周平贞.尼泊尔马桑放线菌共生固氮根瘤的感染和发育[J].武汉植物学研究,1991,9(1)：1-4.
吴晓雷,贺超英,陈受益,等.用SSR分子标记研究大豆属种间亲缘进化关系[J].遗传学报,2001,28(4)：359-366.
吴晓雷,贺超英等.大豆遗传图谱的构建和分析[J].遗传学报,2001,28(11)：1051-1061.
吴晓丽.浙北夏季干旱对不同土壤上档木生长和结瘤固氮的影响[J]林业科学研究,1992,5(2)：225-230.
吴祖洪,周国璋.四川恺木根瘤主要氮素同化酶活力及其超微结构研究[J].林业科学研究,1994,7(6)：629-633.
肖兴翠,王海风,张国君,等.桤木纸浆材造林密度实验[J].湖南林业科技,2008,35(4)：27-29.
徐清乾,许忠坤,周小玲,等.四川桤木木材密度和纤维特性研究[J].湖北林业科技,2008,152(4)：12-15.
徐世凤,吴立勋.江南桤木光合特性研究[J].湖南林业科技,1994,21(4)：10-14.
宣继萍,章镇.苹果品种ISSR指纹图谱构建[J].果树学报,2002,19(6)：421-423.
杨志成,叶长青,封建文等.四川桤木幼林施用磷肥效应[J].林业科学研究,1995,8(1)：112-
易克,徐向利,卢向阳,等.利用SSR和ISSR标记技术构建西瓜分子遗传连锁图谱[J].湖南农业大学学报(自然科学版),2003,29(4)：333-337.
尤勇,洪菊生.RAPD标记在杉木种源遗传变异上的应用[J]林业科学,1998,34(4)：32-37.
曾杰,王中仁,周世良,等.广西区西南桦天然居群遗传多样性研究[J].植物生态学报2003,27(1)：66-72.
张立平,林伯年,林德绪,等.葡萄属RAPD分类研究[J].园艺学报,1998,25(2)：191-193.
张绮纹,苏晓华,李金华.杨树优良纸浆材和生态防护林新品种欧美杨107号[J].林业科学,1999,35(3)：31-37.
张颖娟,杨持.中国特有种四合木种群遗传多样性的RAPD分析[J].生态学报,2002(11)：1917-1922.
赵之伟,赵志坚,杜丽涛.欧洲赤杨根瘤及其内生菌的形态研究[J].微生物学通报,1995,22(1)：3-4.
赵之伟.蒙自桤木根瘤内生菌的分离培养[J].微生物学通报,1996,23(5)：259-261.
周连第,兰彦平,曹庆昌,等.板栗叶片性状表型多样性研究[J].中国农学通报,2005,21(9)：136-139.
朱骏义,陆静梅.东北桤木和辽东桤木生物学特性研究.2008
朱万泽,王金锡,薛建辉,等.台湾桤木引种的光合生理特性研究[J].西北植物学报,2004,24(11)：2012--2019.
卓仁英,陈益泰.四川桤木不同群体间遗传分化研究[J].浙江林业科技,2005,25(1)：13-16.
宗成文,曹后男,赵成日,等.桃属植物RAPD反应体系优化[J].延边大学农学学报,2005,27(3)：153-158.
Ager,A.A.1987.Genetic variation in red alder(Alnus rubra.Bong.)in relation to climate and geography in the Pacific Northwest.Ph.D.thesis,Univ.of Washington,Seattle.
Agrawal A A. Optimal foraging and phenotypic plasticity in plants [J].Trends in Ecology and Evolution,2002,17(7):305
Arthur F.Noskowiak.Distribution of aggregate rays in red alder[J]. Wood and Fiber,1978,10(1):58-68.
Borghetti M,et al.Response to water stress of Italian alder seedlings from diverse geographic origins[J]. Canadian Journal of Forest research,1989,19(8):1071-1076.
Briggs,D.G., D.S.DeBell, W. A. Atkinson (Compilers).Utilization and management of Alder[M]. USDA Forest Services Genetic Technology Report. PNW 70,1978,379pp.
Dang,Q.L.C.Y. Xie et al.Genetic variation of ecophysiological traits in red alder(Alnus rubra Bong) [J].Canadian Journal of Forest research,1994,24(11):2150-2156.
DeBell,D.D.,M.A.Radwan and J.M.Kraft.1983.Influence of red alder on chemical properties of a clay loam soil in western Washingtong[J]. USDA For.Serv.,Res.Pap.,PNW-313.
DeBell, Dean S.Turpin, Thomas C.1983. Red Alder. In:Burns, Russel M, compiler. Silvicultural systems for the major forest types of the United States. ed. Agricultural Handbook No.445. Washington:U.S.Department of Agriculture:26-28
De la Rosa,A Angiolillo,C Guerrero.A first linkage map of olive(Olea europaea L.)cultivars using RAPD,AFLP,RFLP and SSR markers[J]. Theor Appl Genet,2003,106:965-970.
Fowells, H. A.,compiler.1965. Silvics of forest trees of the United States. Agric. Handb.271. Washington, DC:U.S. Department of Agriculture, Forest Service.762 p.
Franco J, Crossa J, Ribaut J M, et al. A method for combining molecular markers and phenotypic attributes for classifying p lant genotypes[J].Theor Appl Genet,2000,103:944-952.
Goncalves,P-de-S,R.C.Kellison.Potential of black alder in the South.Technical Report,School of Forest Resources,North Carolina State University,1980,No.62
Haeussler,S.1986.Germination and first-year survival of red alder seedlings in the central Coast Range of Oregon.M..S.thesis,Oregon State Univ,Corvallis.
Hagidimitrou M,Kastsiotis A,Menexes G,Pontikis C,Loukas M.Genetic diversity of major Greek olive cultivars using molecular (AFLPs and RAPDs) markers and morphological traits[J]. JAmer Soc Hort Sci,2005,130:211-217.
Han,Y.C.K.O.Ryu et.al.Provenance test of 53 provenances of European alder(Alnus glutinosa (L.) Gaertn) [J].Research Report Institute Forest of genetics,1990, No.26,55-62.
Lester,D.D.,Murray,M.D.,Debell,D.S.1989.Geographic variation in red alder. USDA For.Serv.Pacific Northwest Res. Stn. Res. Pap. PNW-409
Martin K.J.,Y.Tanaka and D.D.Myrold.1991.Peat carrier increases inoculation success with Frankia on red alder(Alnus rubra Bong.)in fumigated nursery beds[J].New For.5:43-50.
McDeill,J.D.,M.K.Hollingsworth,W.L.Peterson,A.J.Moffat,L.J.Sheppard,and C.T.Wheppard,and C.T.Wheeler.1989.Inoculation of Alnus rubra seedlings ton improve seedling growth and forest performance[J].Great Britain For.Comm.Res.Div.,Res.Inf.Note 144.
Murillo O.and H.H.Hattemer.1997.Inheritance of isozyme variants of Alnus acuminata ssp.arguta (Schlectendal) Furlow[J].Silvae Genetica,46(1):51-55
O. Murillo,H.H.Hattemer.lnheritance of Isozvme variants of Anus acuminat.assp.arguta(Schlectendal)Furlow.Silvae Genetica, 1997,46(1):51-55.
Perinet P,Tremblay F M.Commercial micropropagation of actinorhizal alder.New Forests,1987,1(3):225-230.
Pieri,S.,S.Teri.Variation height:collar-diameter ratio in young Italian alder(Alnus cordata)families[J].Annali Accademia Italinan di Scienze Forestli,1989,38:391-400
Radwan M A, Max T A, Johnson D W. Soft wood cuttings for propagation of red alder[J]. New Forests,1989,3(1):21-30.
Roger E.Hernandez,Guillermo Restrepo. Natural variation in wood properties of Alnus acuminata H. B. K. grown in Colombia[J]. Wood and Fiber Science,1995,27(1):41-48.
Sinons A M,Johnston M O.Variation in seed traits of Lobelia inflate (Campanulaceace):sources and fitness consequences [J]. American Journal of Botany,2000,87(1):124-132.
Stettler.R.F.1978.Biological aspects of red alder pertinent to potentiaol breeding programs.In Utilization and management of alder[M].Compiled by D.GBriggs.,D.S.DeBell,and W.A.Atkinson.USDA For.Serv.,Gen.Tech.Rep.PNW-70.209-222.
Usman K. J.Suharjo.John D.Tjepkema.Occurrence of hemoglobin in the nitrogen-fixing root nodules of Alnus glutinosa[J]. Physiologic Plantarum,1995,95:247-252.
Wheeler,C.T.,M.K.Hollingsworth,J.E.Hooker,J.D.McNeill,W.L.Mason,A.J.Moffat,and L.J.Sheppard.l991
Worthington,N.P.,R.H.Ruth,and E.E.Matson,Red alder:its management and utilization.USDA For.Serv.,1962,Misc.Publ.881.
Zhang Jian-hua Variation and allometry of seed weight in Aeschynomene America[J]. Annals of Botany,1998,82:843-847.
曹后男,高光出.桃属植物随机扩增多态性DNA分析的最适反应条件[J].延边大学农学学报,1999,21(4)：249-253.
陈益泰,李桂,王惠雄.桤木自然分布区内表型变异的研究[J].林业科学研究,1999,12(4)：
陈之端,路安民.桦木科植物的起源和早期演化[J].中国科学院研究生院学报,1995,12(2)：200-205.
陈之端,路安民.桦木科植物的系统发育和演化[J].中国科学院院刊,2001,(3)：188-191.
陈之端.桦木科植物的系统发育与地理分布(续)[J].植物分类学报,1994,32(2)：101-153
陈志伟,官华忠,吴为人,等.稻瘟病抗性基因Pi-1连锁SSR标记的筛选和应用.福建农林大学学报(自然科学版),2005,34(1)：74-77.
成俊卿.木材学[M].北京：中国林业出版社,1985.
崔成日,徐启江,崔崇士,等.洋葱RAPD遗传多样性分析[J].园艺学报,2006,33(4)：863-865.
单雪,王秀利,仇雪梅,等.分子标记及其在海洋动物遗传研究中的应用[J].生物技术通讯,2005,16(4)：463-466.
葛颂,王明庥,陈岳武.用同工酶研究马尾松群体的遗传结构[J]林业科学,1988,24(4)：399-409.
辜云杰,王启和,罗建勋.四川桤木天然群体果实表型多样性研究[J].四川林业科技,2009,30(2)：19-22.
顾万军,刘和林,岳德智,等.桤木优树子代苗期性状变异的研究[J].四川林业科技,1995,16(3)：52-57.
韩斌,巩合德.哀牢山桤木次生林根瘤固氮能力研究[J].安徽农业研究,2009,37(17)：8272-8273,8283.
黄福平,梁月荣,陆建良,等.应用RAPD和ISSR分子标记构建茶树回交1代部分遗传图谱[J].茶叶科学,2006,26(3)：171-176.
黄志宏,黄维南.豆科茎瘤植物的结瘤、固氮和应用[J].亚热带植物通讯,1996,25(2)：40-46.
季天祜.第二林业与林纸结合[M].北京：中国轻工业出版社,1997.
季维智,宿兵.遗传多样性的原理与方法[M].杭州：浙江科学技术出版社,1999.
贾继增.分子标记种质资源鉴定和分子育种[J].中国农业科学,1996,29(4)：1-10.
姜建国,严媛,宋冬林.盐藻Psy基因保守序列的克隆及同源性分析[J].华南理工大学学报：自然科学版,2004,32(10)：76-79.
姜静,杨传平,刘桂丰,等.应用RAPD技术对东北地区白桦种源遗传变异的分析[J].东北林业大学学报,2001,29(2)：30-34.
蒋建德,赵振英,朱宝琴.非豆科树木根瘤内生菌的分离[J].林业科学，1987,23(1)：85-89.
李斌,顾万春,卢宝铭.白皮松天然群体种实性状表型多样性研究[J].生物多样性,2002,10(2)：181-187.
李洁,熊智.云南尼泊尔桤木遗传多样性研究[J].浙江林学院学报,2008,25(1)：16-21.
梁月荣,田中村一,武田善行.应用RAPD分子标记分析“晚绿”品种的杂交亲本[J].茶树科学,2000,20(1)：22.
林清洪,黄维南.福建省主要固氮树木VA菌根真菌的分离鉴定[J].亚热带植物讯,1999,28(1)：65.
刘士平,李信,汪朝阳,等.利用分子标记辅助改良珍汕97的稻瘟病抗性[J].植物学报,2003,45(11)：1346-1350.
刘延吉,耿书,田晓艳.辽宁地区五种野生软枣猕猴桃RAPD遗传多样性分析[J].北方园艺.2010,(18)：130-132
吕梅.红桤木生理生态、固氮活性及根瘤放线菌研究[D].南京林业大学学位论文,2008,3-24.
罗建勋.云杉天然群体遗传多样性研究[D].中国林业科学研究院,2004.
罗美娟.短枝木麻黄种源群体遗传多样性与遗传变异规律研究[D].福州：福建农林大学图书馆,2005.
M. Lalonde.桤木根瘤共生固氮研究技术与观测[J].亚热带植物通讯,1990,1：59-62.
潘建蓄,陈启锋,黄世贞.诱导固氮菌与稻苗结瘤共生的研究[J].福建农业大学学报,1998,27(4)：405-410.
潘远智,庞博,孙振元,等.雅江报春天然居群RAPD遗传多样性[J].四川农业大学学报.2008.26(1)：28-31.
庞广昌,姜冬梅.群体遗传多样性和数据分析[J].林业科学,1995,31(6)：543-550.
任旭琴.遗传多样性及其研究方法[J].淮阴工学学报,2002,11(5)：6-8.
沈向,郭卫东,吴燕民,等.杏43个品种资源的RAPD分类[J].园艺学报,2000,27(1)：55-56.
税珺,黄少伟,陈炳铨.火炬松原生种源和引种群体RAPD遗传多样性[J].华南农业大学学报,2005,26(3)：74-76,81.
宋丛文,张新叶,胡兴宜.秃杉种内遗传多样性的RAPD分析[J].湖北林业科技,2004(4)：1-4.
汪小全,邹喻苹,张大明,等.RAPD应用于遗传多样性和系统学研究中的问题[J].植物学报,1996,38(12)：954-962.
王晨光,阮继生,石彦林.木麻黄根瘤中分离的四株弗兰克氏菌的比较研究[J].微生物学通报,1993,20(3)：80-83.
王璐璐.贵州省白菜型油菜的RAPD遗传多样性分析[J].贵州农业科学,2009,37(10)：5-7.
王祖良,丁丽霞,赵明水,等.濒危植物天目铁木遗传多样性的RAPD分析[J].浙江林学院学报,2008,25(3)：304-308.
吴捷,陈华癸,周平贞.尼泊尔马桑放线菌共生固氮根瘤的感染和发育[J].武汉植物学研究,1991,9(1)：1-4.
吴晓雷,贺超英,陈受益,等.用SSR分子标记研究大豆属种间亲缘进化关系[J].遗传学报,2001,28(4)：359-366.
吴晓雷,贺超英等.大豆遗传图谱的构建和分析[J].遗传学报,2001,28(11)：1051-1061.
吴晓丽.浙北夏季干旱对不同土壤上档木生长和结瘤固氮的影响[J]林业科学研究,1992,5(2)：225-230.
吴祖洪,周国璋.四川恺木根瘤主要氮素同化酶活力及其超微结构研究[J].林业科学研究,1994,7(6)：629-633.
肖兴翠,王海风,张国君,等.桤木纸浆材造林密度实验[J].湖南林业科技,2008,35(4)：27-29.
徐清乾,许忠坤,周小玲,等.四川桤木木材密度和纤维特性研究[J].湖北林业科技,2008,152(4)：12-15.
徐世凤,吴立勋.江南桤木光合特性研究[J].湖南林业科技,1994,21(4)：10-14.
宣继萍,章镇.苹果品种ISSR指纹图谱构建[J].果树学报,2002,19(6)：421-423.
杨志成,叶长青,封建文等.四川桤木幼林施用磷肥效应[J].林业科学研究,1995,8(1)：112-
易克,徐向利,卢向阳,等.利用SSR和ISSR标记技术构建西瓜分子遗传连锁图谱[J].湖南农业大学学报(自然科学版),2003,29(4)：333-337.
尤勇,洪菊生.RAPD标记在杉木种源遗传变异上的应用[J]林业科学,1998,34(4)：32-37.
曾杰,王中仁,周世良,等.广西区西南桦天然居群遗传多样性研究[J].植物生态学报2003,27(1)：66-72.
张立平,林伯年,林德绪,等.葡萄属RAPD分类研究[J].园艺学报,1998,25(2)：191-193.
张绮纹,苏晓华,李金华.杨树优良纸浆材和生态防护林新品种欧美杨107号[J].林业科学,1999,35(3)：31-37.
张颖娟,杨持.中国特有种四合木种群遗传多样性的RAPD分析[J].生态学报,2002(11)：1917-1922.
赵之伟,赵志坚,杜丽涛.欧洲赤杨根瘤及其内生菌的形态研究[J].微生物学通报,1995,22(1)：3-4.
赵之伟.蒙自桤木根瘤内生菌的分离培养[J].微生物学通报,1996,23(5)：259-261.
周连第,兰彦平,曹庆昌,等.板栗叶片性状表型多样性研究[J].中国农学通报,2005,21(9)：136-139.
朱骏义,陆静梅.东北桤木和辽东桤木生物学特性研究.2008
朱万泽,王金锡,薛建辉,等.台湾桤木引种的光合生理特性研究[J].西北植物学报,2004,24(11)：2012--2019.
卓仁英,陈益泰.四川桤木不同群体间遗传分化研究[J].浙江林业科技,2005,25(1)：13-16.
宗成文,曹后男,赵成日,等.桃属植物RAPD反应体系优化[J].延边大学农学学报,2005,27(3)：153-158.
Ager,A.A.1987.Genetic variation in red alder(Alnus rubra.Bong.)in relation to climate and geography in the Pacific Northwest.Ph.D.thesis,Univ.of Washington,Seattle.
Agrawal A A. Optimal foraging and phenotypic plasticity in plants [J].Trends in Ecology and Evolution,2002,17(7):305
Arthur F.Noskowiak.Distribution of aggregate rays in red alder[J]. Wood and Fiber,1978,10(1):58-68.
Borghetti M,et al.Response to water stress of Italian alder seedlings from diverse geographic origins[J]. Canadian Journal of Forest research,1989,19(8):1071-1076.
Briggs,D.G., D.S.DeBell, W. A. Atkinson (Compilers).Utilization and management of Alder[M]. USDA Forest Services Genetic Technology Report. PNW 70,1978,379pp.
Dang,Q.L.C.Y. Xie et al.Genetic variation of ecophysiological traits in red alder(Alnus rubra Bong) [J].Canadian Journal of Forest research,1994,24(11):2150-2156.
DeBell,D.D.,M.A.Radwan and J.M.Kraft.1983.Influence of red alder on chemical properties of a clay loam soil in western Washingtong[J]. USDA For.Serv.,Res.Pap.,PNW-313.
DeBell, Dean S.Turpin, Thomas C.1983. Red Alder. In:Burns, Russel M, compiler. Silvicultural systems for the major forest types of the United States. ed. Agricultural Handbook No.445. Washington:U.S.Department of Agriculture:26-28
De la Rosa,A Angiolillo,C Guerrero.A first linkage map of olive(Olea europaea L.)cultivars using RAPD,AFLP,RFLP and SSR markers[J]. Theor Appl Genet,2003,106:965-970.
Fowells, H. A.,compiler.1965. Silvics of forest trees of the United States. Agric. Handb.271. Washington, DC:U.S. Department of Agriculture, Forest Service.762 p.
Franco J, Crossa J, Ribaut J M, et al. A method for combining molecular markers and phenotypic attributes for classifying p lant genotypes[J].Theor Appl Genet,2000,103:944-952.
Goncalves,P-de-S,R.C.Kellison.Potential of black alder in the South.Technical Report,School of Forest Resources,North Carolina State University,1980,No.62
Haeussler,S.1986.Germination and first-year survival of red alder seedlings in the central Coast Range of Oregon.M..S.thesis,Oregon State Univ,Corvallis.
Hagidimitrou M,Kastsiotis A,Menexes G,Pontikis C,Loukas M.Genetic diversity of major Greek olive cultivars using molecular (AFLPs and RAPDs) markers and morphological traits[J]. JAmer Soc Hort Sci,2005,130:211-217.
Han,Y.C.K.O.Ryu et.al.Provenance test of 53 provenances of European alder(Alnus glutinosa (L.) Gaertn) [J].Research Report Institute Forest of genetics,1990, No.26,55-62.
Lester,D.D.,Murray,M.D.,Debell,D.S.1989.Geographic variation in red alder. USDA For.Serv.Pacific Northwest Res. Stn. Res. Pap. PNW-409
Martin K.J.,Y.Tanaka and D.D.Myrold.1991.Peat carrier increases inoculation success with Frankia on red alder(Alnus rubra Bong.)in fumigated nursery beds[J].New For.5:43-50.
McDeill,J.D.,M.K.Hollingsworth,W.L.Peterson,A.J.Moffat,L.J.Sheppard,and C.T.Wheppard,and C.T.Wheeler.1989.Inoculation of Alnus rubra seedlings ton improve seedling growth and forest performance[J].Great Britain For.Comm.Res.Div.,Res.Inf.Note 144.
Murillo O.and H.H.Hattemer.1997.Inheritance of isozyme variants of Alnus acuminata ssp.arguta (Schlectendal) Furlow[J].Silvae Genetica,46(1):51-55
O. Murillo,H.H.Hattemer.lnheritance of Isozvme variants of Anus acuminat.assp.arguta(Schlectendal)Furlow.Silvae Genetica, 1997,46(1):51-55.
Perinet P,Tremblay F M.Commercial micropropagation of actinorhizal alder.New Forests,1987,1(3):225-230.
Pieri,S.,S.Teri.Variation height:collar-diameter ratio in young Italian alder(Alnus cordata)families[J].Annali Accademia Italinan di Scienze Forestli,1989,38:391-400
Radwan M A, Max T A, Johnson D W. Soft wood cuttings for propagation of red alder[J]. New Forests,1989,3(1):21-30.
Roger E.Hernandez,Guillermo Restrepo. Natural variation in wood properties of Alnus acuminata H. B. K. grown in Colombia[J]. Wood and Fiber Science,1995,27(1):41-48.
Sinons A M,Johnston M O.Variation in seed traits of Lobelia inflate (Campanulaceace):sources and fitness consequences [J]. American Journal of Botany,2000,87(1):124-132.
Stettler.R.F.1978.Biological aspects of red alder pertinent to potentiaol breeding programs.In Utilization and management of alder[M].Compiled by D.GBriggs.,D.S.DeBell,and W.A.Atkinson.USDA For.Serv.,Gen.Tech.Rep.PNW-70.209-222.
Usman K. J.Suharjo.John D.Tjepkema.Occurrence of hemoglobin in the nitrogen-fixing root nodules of Alnus glutinosa[J]. Physiologic Plantarum,1995,95:247-252.
Wheeler,C.T.,M.K.Hollingsworth,J.E.Hooker,J.D.McNeill,W.L.Mason,A.J.Moffat,and L.J.Sheppard.l991
Worthington,N.P.,R.H.Ruth,and E.E.Matson,Red alder:its management and utilization.USDA For.Serv.,1962,Misc.Publ.881.
Zhang Jian-hua Variation and allometry of seed weight in Aeschynomene America[J]. Annals of Botany,1998,82:843-847.