利用分子标记探讨赖草属植物的系统关系
|
摘要
赖草属Leymus Hochst.是禾本科Poaceae小麦族Triticeae Dumortier的一个重要多年生属,全世界大约有30个种和19个亚种,从北海的沿岸地区,越过中亚到东亚直至阿拉斯加和北美西部的广阔地域均有分布,多数种类集中于中亚和北美的高山。我国原记载约10种,加上近年来报道的新分类群,共25个种,4个亚种和3个变种,主要分布于西北、华北、东北及西南地区。该属的多数物种为草原和草甸的主要组成成分,许多种类是优良的牧草,具有较高的饲用价值。由于赖草属植物常生长在盐碱地和干早半干早的山坡、地埂,该属植物的有些物种对寒冷、干早、盐碱土等不良环境具有高度的适应性。同时,有些赖草属物种还具有抗病虫、穗大、粒多、粒大、高光效等优良特性。因此,作为麦类作物育种的重要三级基因源,赖草属植物对改良遗传基础日益狭窄的麦类作物有重要意义。然而,有关该属植物的起源、系统地位、物种界限、种间(内)亲缘关系等问题,仍然存在较大的分歧。
针对赖草属物种中Ns基因组的供体物种以及另一个基因组的来源问题,本研究利用同工酶、醇溶蛋白及RAPD标记对14种赖草属植物和10种近缘属二倍体物种的系统关系进行了分析。主要结果如下:
1.利用垂直平板聚丙烯酰胺凝胶电泳(PAGE)对赖草属、冰草属、偃麦草属、澳冰草属、拟鹅观草属、新麦草属和大麦属7个属24个物种的酯酶同工酶和超氧化物歧化酶同工酶进行比较分析。结果表明:①7个属24个物种共出现24种不同的同工酶酶谱,电泳分离出迁移率不同的45条酶带;②Leymus chinensis与Leymus其它13个种的亲缘关系较远,与Hordeum bogdanii的亲缘关系较近;Psathyrostachys juncea与Leymus的亲缘关系很近;③相似染色体组的物种聚类在一起;④两种同工酶的分析结果与细胞学及分子标记的研究结果基本一致。
2.利用酸性聚丙烯酰胺凝胶电泳(A-PAGE)对赖草属12种和冰草属、偃麦草属、澳冰草属、拟鹅观草属、新麦草属和大麦属7个属10个种共22份材料进行醇溶蛋白遗传分析。结果表明:①7个属22份材料共出现22种不同的醇溶蛋白图谱,22种图谱中分离出的52条带纹,多态性高达100%,说明属间和种间具有丰富的醇溶蛋白多态性;②Psathyrostachys juncea与Psa.huashanica的亲缘关系较远,而Psa.juncea与Leymus的亲缘关系很近;③含有相似染色体组的物种能各自聚类在一起,它们具有较近的亲缘关系;④醇溶蛋白的分析结果与细胞学及分子标记的研究结果基本一致,表明醇溶蛋白能够用于小麦族多年生物种系统亲缘关系分析。
3.对14个赖草属物种和10个近缘属物种进行基因组特异RAPD标记。选用小麦族中6个基本基因组(E、H、P、St、W、Ns)的特异RAPD引物进行PCR扩增检测,结果表明:①14个赖草属物种具有Ns基因组特异的RAPD标记,而没有P和E基因组的特异的RAPD标记;②Leymus ramosus具有W基因组特异的RAPD标记;L.triticoides、L.ramosus、L.paboanus、L.chinensis、L.pseudoracemosus具有St基因组特异的RAPD标记;L.angustus、L.arenarius、L.multicaulis、L.karelinii、L.tianshanicus、L.paboanus、L.cinereus、L.chinensis、L.racemosus具有H基因组特异的RAPD标记;③L.paboanus、L.chinensis不仅出现了H基因组的特异性条带,还出现了St基因组的特异性条带;L.ramosus不仅出现了W基因组的特异性条带,还出现了St基因组的特异性条带。
结合本实验的研究结果,赖草属植物为多重起源,Ns基因组来自新麦草属,Psathyrostachys juncea与赖草属系统关系较近。赖草属不含E、P基因组,St、W和H基因组参与了部分赖草属物种的起源。
Leymus Hochst.is an important perennial genus of Triticeae Dumortier(Poaceae). It includes about 30 species and 19 subspecies.They are distributed in the coastal areas of North Sea,Central Asia,East Asia,extending to Alaska and the western areas of North America.They are found in particularly large numbers on the mountains of Central Asia and North America.Formerly,it included about 10 species recorded in China.In addition to some new taxon reported in recent years,it includes about 25 species,4 subspecies and 3 varieties,which are mainly distributed in north-western, north,north-eastern and south-western regions in China.Many species of Leymus are not only main components of grasslands and grass marshlands,but also fine varieties of herbage and forge.Growing in saline or alkaline lands,and dry or semi-dry areas,some species are highly adaptable to coldness,dryness and saline or alkaline lands.And some species bear such desirable traits as disease and insect resistance,bigger spikes,more and bigger grains and efficient photosynthesis.Being an important genetic resource, Leymus is of significance to improvement of Triticeae cereal crops.However,the origin and definition of the genus,precise taxonomic ranks and relationships among the species in the genus have been under discussion.The study on genetic diversity of Leymus would provide theoretical foundation for using this genus plants to breed Triticeae crops and herbage.
The present study systematically evaluated the systematics relationships of Leymus on isoenzyme,gliadin and RAPD molecular level,and investigated the parental Psathyrostachys ancestries which offered the Ns genome to Leymus and to elucidate the origins of another base genome of Leymus.The main results showed as following:
1.The esterase isozyme(EST) and superoxide isozyme(SOD) of 24 species of Leymus,Australopyrum,Agropyron,Lophopyrum,Pseudoroegneria,Psathyrostachys and Hordeum in triticeae were analyzed by using polyacrylamid gel electrophoresis (PAGE) technique.The results show as follows:①All the 24 species have significant isozymes polymorphism,a total of 45 bands are separted;②The relationship between L.chinensis and the other 13 species of Leymus is remote,while it is close to Hordeum bogdanii,Psathyrostachys juncea and the species of Leymus have closer relationship;③The species of those which contain similar genomes are clustered together;④The results of isozymes analysis are basically comparable with those obtained from studies on cytology and morphology.
2.The gliadin of 22 materials of Leymus,Australopyrum,Agropyron, Lophopyrum,Pseudoroegneria,Psathyrostachys and Hordeum in Triticeae were analyzed by using acid-polyacrylamide gel electrophoresis(A-PAGE).The results show as follows:①All the 22 materials have significant gliadin polymorphism.A total of 52 bands are separated,and the gliadin ploymorphism amounts to 100%.It shows that there is plentiful polymorphism between these genera and species;②The relationship between Psathyrostachys juncea and Psa.huashanica is remote,while Psa.juncea and the species of Leymus have closer relationship;③The species of those which contain similar genomes are clustered together,which suggests they have closer relationship;④The results of gliadin analysis are basically comparable with those obtained from studies on cytology and morphology,which shows that the gliadin patterns is usefulness in the systematic studies of those perennial species in Triticeae.
3.Six genome-specific RAPD markers representing Ns,E,W,P,St,H genome respectively were utilized to determine the genome relationships among Leymus and its related species.The results indicated that:the Ns genome-specific RAPD marker presented in 14 polyploid species of Leymus,while the E and P genome-specific RAPD markers absent in 14 polyploid species of Leymus;②the W genome-specific RAPD marker presented in L.ramosus;the St genome-specific RAPD marker presented in L.triticoides,L.ramosus,L.paboanus,L.chinensis,L. pseudoracemosuss;the H genome-specific RAPD marker presented in L.angustus,L. arenarius,L.multicaulis,L.karelinii,L.tianshanicus,L.paboanus,L.cinereus,L. chinensis,L.racemosus;③L.paboanus and L.chinensis had the specific bands of Ns genome,H genome and St genome;L.ramosus had the specific bands of Ns genomes, St genomes and W genomes.
Based on these results from this study,we can draw conclusions that:Leymus species have multiple origins.Ns genome arose from the genus Psathyrostachys, While Psa.juncea and the species of Leymus have closer relationship.E and P genome absent in Leymus,While the partial species of Leymus had close phylogenetic relationship with St,W and H genome.
针对赖草属物种中Ns基因组的供体物种以及另一个基因组的来源问题,本研究利用同工酶、醇溶蛋白及RAPD标记对14种赖草属植物和10种近缘属二倍体物种的系统关系进行了分析。主要结果如下:
1.利用垂直平板聚丙烯酰胺凝胶电泳(PAGE)对赖草属、冰草属、偃麦草属、澳冰草属、拟鹅观草属、新麦草属和大麦属7个属24个物种的酯酶同工酶和超氧化物歧化酶同工酶进行比较分析。结果表明:①7个属24个物种共出现24种不同的同工酶酶谱,电泳分离出迁移率不同的45条酶带;②Leymus chinensis与Leymus其它13个种的亲缘关系较远,与Hordeum bogdanii的亲缘关系较近;Psathyrostachys juncea与Leymus的亲缘关系很近;③相似染色体组的物种聚类在一起;④两种同工酶的分析结果与细胞学及分子标记的研究结果基本一致。
2.利用酸性聚丙烯酰胺凝胶电泳(A-PAGE)对赖草属12种和冰草属、偃麦草属、澳冰草属、拟鹅观草属、新麦草属和大麦属7个属10个种共22份材料进行醇溶蛋白遗传分析。结果表明:①7个属22份材料共出现22种不同的醇溶蛋白图谱,22种图谱中分离出的52条带纹,多态性高达100%,说明属间和种间具有丰富的醇溶蛋白多态性;②Psathyrostachys juncea与Psa.huashanica的亲缘关系较远,而Psa.juncea与Leymus的亲缘关系很近;③含有相似染色体组的物种能各自聚类在一起,它们具有较近的亲缘关系;④醇溶蛋白的分析结果与细胞学及分子标记的研究结果基本一致,表明醇溶蛋白能够用于小麦族多年生物种系统亲缘关系分析。
3.对14个赖草属物种和10个近缘属物种进行基因组特异RAPD标记。选用小麦族中6个基本基因组(E、H、P、St、W、Ns)的特异RAPD引物进行PCR扩增检测,结果表明:①14个赖草属物种具有Ns基因组特异的RAPD标记,而没有P和E基因组的特异的RAPD标记;②Leymus ramosus具有W基因组特异的RAPD标记;L.triticoides、L.ramosus、L.paboanus、L.chinensis、L.pseudoracemosus具有St基因组特异的RAPD标记;L.angustus、L.arenarius、L.multicaulis、L.karelinii、L.tianshanicus、L.paboanus、L.cinereus、L.chinensis、L.racemosus具有H基因组特异的RAPD标记;③L.paboanus、L.chinensis不仅出现了H基因组的特异性条带,还出现了St基因组的特异性条带;L.ramosus不仅出现了W基因组的特异性条带,还出现了St基因组的特异性条带。
结合本实验的研究结果,赖草属植物为多重起源,Ns基因组来自新麦草属,Psathyrostachys juncea与赖草属系统关系较近。赖草属不含E、P基因组,St、W和H基因组参与了部分赖草属物种的起源。
Leymus Hochst.is an important perennial genus of Triticeae Dumortier(Poaceae). It includes about 30 species and 19 subspecies.They are distributed in the coastal areas of North Sea,Central Asia,East Asia,extending to Alaska and the western areas of North America.They are found in particularly large numbers on the mountains of Central Asia and North America.Formerly,it included about 10 species recorded in China.In addition to some new taxon reported in recent years,it includes about 25 species,4 subspecies and 3 varieties,which are mainly distributed in north-western, north,north-eastern and south-western regions in China.Many species of Leymus are not only main components of grasslands and grass marshlands,but also fine varieties of herbage and forge.Growing in saline or alkaline lands,and dry or semi-dry areas,some species are highly adaptable to coldness,dryness and saline or alkaline lands.And some species bear such desirable traits as disease and insect resistance,bigger spikes,more and bigger grains and efficient photosynthesis.Being an important genetic resource, Leymus is of significance to improvement of Triticeae cereal crops.However,the origin and definition of the genus,precise taxonomic ranks and relationships among the species in the genus have been under discussion.The study on genetic diversity of Leymus would provide theoretical foundation for using this genus plants to breed Triticeae crops and herbage.
The present study systematically evaluated the systematics relationships of Leymus on isoenzyme,gliadin and RAPD molecular level,and investigated the parental Psathyrostachys ancestries which offered the Ns genome to Leymus and to elucidate the origins of another base genome of Leymus.The main results showed as following:
1.The esterase isozyme(EST) and superoxide isozyme(SOD) of 24 species of Leymus,Australopyrum,Agropyron,Lophopyrum,Pseudoroegneria,Psathyrostachys and Hordeum in triticeae were analyzed by using polyacrylamid gel electrophoresis (PAGE) technique.The results show as follows:①All the 24 species have significant isozymes polymorphism,a total of 45 bands are separted;②The relationship between L.chinensis and the other 13 species of Leymus is remote,while it is close to Hordeum bogdanii,Psathyrostachys juncea and the species of Leymus have closer relationship;③The species of those which contain similar genomes are clustered together;④The results of isozymes analysis are basically comparable with those obtained from studies on cytology and morphology.
2.The gliadin of 22 materials of Leymus,Australopyrum,Agropyron, Lophopyrum,Pseudoroegneria,Psathyrostachys and Hordeum in Triticeae were analyzed by using acid-polyacrylamide gel electrophoresis(A-PAGE).The results show as follows:①All the 22 materials have significant gliadin polymorphism.A total of 52 bands are separated,and the gliadin ploymorphism amounts to 100%.It shows that there is plentiful polymorphism between these genera and species;②The relationship between Psathyrostachys juncea and Psa.huashanica is remote,while Psa.juncea and the species of Leymus have closer relationship;③The species of those which contain similar genomes are clustered together,which suggests they have closer relationship;④The results of gliadin analysis are basically comparable with those obtained from studies on cytology and morphology,which shows that the gliadin patterns is usefulness in the systematic studies of those perennial species in Triticeae.
3.Six genome-specific RAPD markers representing Ns,E,W,P,St,H genome respectively were utilized to determine the genome relationships among Leymus and its related species.The results indicated that:the Ns genome-specific RAPD marker presented in 14 polyploid species of Leymus,while the E and P genome-specific RAPD markers absent in 14 polyploid species of Leymus;②the W genome-specific RAPD marker presented in L.ramosus;the St genome-specific RAPD marker presented in L.triticoides,L.ramosus,L.paboanus,L.chinensis,L. pseudoracemosuss;the H genome-specific RAPD marker presented in L.angustus,L. arenarius,L.multicaulis,L.karelinii,L.tianshanicus,L.paboanus,L.cinereus,L. chinensis,L.racemosus;③L.paboanus and L.chinensis had the specific bands of Ns genome,H genome and St genome;L.ramosus had the specific bands of Ns genomes, St genomes and W genomes.
Based on these results from this study,we can draw conclusions that:Leymus species have multiple origins.Ns genome arose from the genus Psathyrostachys, While Psa.juncea and the species of Leymus have closer relationship.E and P genome absent in Leymus,While the partial species of Leymus had close phylogenetic relationship with St,W and H genome.
引文
[1]Bothmer R von,Salmon B.Triticeae:a tribe for food,feed and fun.in:Wang R R-C,K B Jensen,C Jaussi(eds),Proc 2nd Inern Triticeae Symp Logan.USA,Logan:Utah State University,1994,1-12.
[2]Dewey D R.The genomic system of classification as a guide to intergeneric hybridization with the perennial Triticeae.In:Gustafson J P(eds),Gene Manipulation in plant improvement.NewYork:Plenum,1984,209-280.
[3]杨瑞武.赖草属植物的系统与进化研究.雅安:四川农业大学,2003.
[4]董玉琛.小麦野生近缘植物的研究和利用.李胜琳,马缘生主编,植物优异种质资源及其开拓利用.北京:中国科学技术出版社.1992,39-44.
[5]刘大钧.向小麦转移外源抗病性的回顾与展望.南京农业大学学报,1994a,17(3):1-7.
[6]刘大钧.外源基因在小麦育种中的利用.作物杂志,1994b,(6):1-7.
[7]Riley R,Chapman V,Johnson R.Introduction of yellow rust resistance of Aegilops comosa into wheat by genetically induced homoeologous combination.Nature,1968,217:383-384.
[8]Feldman M..Gene transfer from wild species into cultivated plants.Genetika,1983,15:145-161.
[9]Feldman M.Cytogenetics and molecular approaches to alien gene transfer in wheat.In:Miller T E,Koebner R M D(eds),Proc 7th Int Wheat Genet Symp.Cambridge,England,Cambridge:Cambridge University Press,1988,23-32.
[10]Sharma H C,Gill B S.Current Status of wide hybridization in wheat.Euphytica,1983,32:17-31.
[11]Dvorak J,Meguire P E.Triticeae,the gene pool for wheat breeding.Genome mapping of wheat and related species.Publ Workshop.Westsaerment,California,1990,3-8.
[12]Jiang J M,Friebe B,Gill B S.Recent advances in alien gene tranfer in wheat.Euphytica,1994,73:199-212.
[13]Hochstetter C F.Nachtraglicher Commentar zu meiner Abhandlung."Aufbau der Graspflanze etc."Flora 1848,7:105-118.
[14]Pilger R.Addimenta Agrostologica.Ⅰ.Triticeae(Hordeae).Bot Jahrb,1949,74:1-27.
[15]Pilger R.Das System der Graminaea.Bot Jahrb,1954,76:281-284.
[16]耿以礼.中国主要植物图说.禾本科.北京:科学出版社,1959,342-409.
[17]Tzvelev N N.Despeciebus nonnulis novis vel minus cognitise pamur.Bot Mater Gerb Bot Inst Komarova Akad Nauk SSR(Leningrad),1960,20:412-439.
[18] Tzvelev N N. Tribe 3. Triticeae Dum. Poaceae URSS. Leningrad: Nauka, 1976. 176-189.
[19] Love A, D Love. Some nomenclatural changes in the European flora. I .Species and supraspecific categories. Bot Not, 1961, 114: 33-47.
[20] Melderis A. Leymus. In: T G Tutin et al., (eds). Flora Eueopaea. vol. 5. Cambridge: Cambridge University Press, 1980, 190-192.
[21] Bentham G.. Notes on Gramineae. Bot J Linn Soc, 1881,18:14-134.
[22] Bentham G and J D Hooker. Genera plantarum. vol.3. London. 1881.
[23] Boissier E. Flora Orientalis. vol.5. H Georg Geneva. 1884
[24] Hackel E. Hordeae. In: A Engler and K Prantl (eds). Die naturlichen pflanzentamilien. W Engelmann. Leipzig. 1887, 76-89.
[25] Coste H. Flore descriptive et illustree de la France 3.1906.
[26] Hitchcock C L. Gramineae. In: Hitchcock C. L. et al., (eds.)Vascular Plants of the Pacific Northwest, vol. 1. Seattle: University of Washington Press, 1969, 384-725.
[27] Holmgren A H and N H Holmgren. Triticeae. In: Cronquist et al., (eds). Intermountain Flora, vol. 6. New York: Columbia University Press, 1977, 293-336.
[28] Estes J R, R J Tyrl. The generic concept and generic circumscription in the triticeae: an end paper. In: J R Estes et al., (eds). Grasses and Grasslands. Norman: University of Oklahoma Press, 1982, 145-164.
[29] Nevski S A. Uber das system der tribe Hordeae Benth. Flora et Systematica Plantae vasculares. Ser 1. Fasc 1. Leningrad, 1933, 9-32.
[30] Nevski S A. Tribe XIV Hordeae Benth. In: Komarov V L, Roshevits R Y, Shishkin B K (eds). Flora URSS 2, Leningrad, 1934, 590-720.
[31] Stebbins G L, Walters M S. Artificial and natural hybrids in the Gramineae, tribe Hordeae III Hybrids involving Elymus condensatus and E. triticoides. Amer J Bot, 1949, 36: 291-301.
[32] Dewey D R. Genome relations among diploid Elymus juncea and certain tetraploid and octoploid Elymus species. Amer J Bot, 1970a, 57: 633-639.
[33] Dewey D R. Cytogenetics of tetraploid Elymus cinereus, E. triticoides, E. multicaulis, E. karataviensis, and their F_1 hybrids. Bot Gaz, 1972,133: 51-57.
[34] Dewey D R. The genome constitution and phylogeny of Elymus ambiguous. Amer J Bot, 1976, 63: 626-634.
[35]Dewey D R..Genomic and phylogenetic relationships among North American Perennial Triticeae.In:J R Estes et al.,(eds).Grasses and Grasslands.Norman:University of Oklahoma Press,1982,51-88.
[36]Jaaska V.Enzyme variability and phylogenetic relationships in the grass genera Agropyron Gaertn.And Elymus L.Izv Akad Nauk Estonsk SSR Ser Biol,1974,23:3-18.
[37]Stebbins G L.Taxonomy and the evolution of genera,with special reference to the family Gramineae.Evolution,1956,10:235-245.
[38]Bowden W M.Cytotaxonomy of the species and interspecific hybrids of genus Agropyron in Canada and neighbouring areas.Can J Bot,1965,43:1421-1448.
[39]Love A.Generic evaluation of the wheatgrass.Biol Zentralbl,1982,101:199-212.
[40]Baum B R.Taxonomy of the tribe Triticeae(Poaceae) using various numerical techniques,Ⅰ.Historical perspectives,dta accumulation,and character analysis.Can J Bot,1977,55:1712-1740.
[41]Baum B R.The genus Elymus in Canada-Bowden's generic concept and key reappraised and relectotypification of E.canadenssi.Can J Bot,1979,57:946-951.
[42]Baum B R.The generic problem in the Triticeae:numerical taxonomy and related concepets,In:J R Estes et al.,(eds).Grasses and grasslands.Norman:University of oklahoma Press,1982,109-144.
[43]Barkworth M E,Atkins R J.Leymus Hochst.(Gramineae:Triticeae) in North Ameriea:Taxonomy and distribution.Amer J Bot,1984,71(5):609-625.
[44]Tzvelev N.N.Grasses of the Soviet Union.New Delhi:Oxonian Press PVT.Ltd.,1976b,253-276.
[45]Anamthawat-Jonsson K,Bodvarsdottir S K.Genomic and genetic relationships among species of Leymus(Poaceae:Triticeae) inferred from 18S-26S ribosomal genes.Amer J Bot,2001,88(4):553-559.
[46]Bodvarsdottir and Anamthawat-Jonsson.Isolation,characterization,and analysis of Leymus-specific DNA sequences.Genome,2003,46:673-682.
[47]候宽昭.中国种子植物科属辞典.北京:科学出版社,1958.
[48]候宽昭.中国种子植物科属辞典(修订版).北京:科学出版社,1982.
[49]郭本兆.中国植物志.北京:科学出版社,1987,9(3):15-26.
[50]颜济,杨俊良.中国赖草属新植物.云南植物研究,1983.5(3):275-276.
[51]崔乃然,钟骏平.新疆谷类作物的野生近缘植物.新疆八一农学院学报,1984,(1):11-35.
[52]吴玉虎.新疆赖草属二新种.植物研究,1992,12(4):343-345.
[53]孙根楼,颜济,杨俊良.新分布于中国的一种赖草形态学和细胞学观察[J].四川农业大学学报,1992,10(4):574-550.
[54]蔡联炳.国产赖草属新分类群.植物分类学报,1995,33(5):491-496.
[55]蔡联炳.赖草属资料.植物研究,1997,17(1):28-32.
[56]蔡联炳.青海赖草属一新种和一新变种.植物分类学报,2001,39(1):75-77.
[57]崔大方.新疆赖草属的新分类群.植物研究,1998,18(2):144-148.
[58]赵可夫,李法曾.中国盐生植物.北京:科学出版社,1999,331-334.
[59]Wang R R-C,Hsiao C.Morphology and cytology of interspecific hybrid of Leymus mollis,d Hered,1984,75:488-492.
[60]Zhang H B,Dvorak J.The genome origin of tetraploid of Leymus(Poaceae:triticeae) inferred from variation in repeated nucleotide sequences.Amer J Bot,1991,78:871-884.
[61]Shiotan 1.Species differentiation in Agropyron,Elymusu,Hystrix,and Sitanion.Proceedings of the 12th International Congress of Genetics.The Science Council of Japan,Tokyo.1968,184.
[62]Love A.Conspectus of the Triticeae.Feddes Repert,1984,95:425-521.
[63]Wang R R C,Jensen K B.Absence of the J genome in species(Poaceae:Triticeae):evidence from DNA hybridization and meiotic pairing.Genome,1994,37:231-235.
[64]φrgaard M.and Heslop-Harrison J.S.,Investigations of genome relationships between Leymus,Psathyrostachys and Hordeum inferred by genome DNA:DNA in situ hybridization.Annals of Botany,1994,73:195-203.
[65]Wang R.R-C,Bother V.,Dvorak J.,Linde-Laursen 1.,and Muramatsu M.Genome symbols in the Triticeae(Poaceae).In:Wang R R-C et al.(eds.),Proceedings of the 2nd International Triticeae Symposium.Utah State University Press,Logan,Utah,USA.1994,29-34.
[66]Sun G L,Wu B H,and Liu F.Cytogenetic and genomic relationships of Thinopyrurn elongatum with two Psathyrostachys species and with Leymus secaIinus Poaceae.Plant Systematics and Evolution,1995a,197:225-231.
[67]Wu X L,Larson,Hu Z M,Palazzo A J,Jones T A,Wang R R-C,Jensen K B,and Chatterton N J.Molecular genetic linkage maps for alloteterap1oid Leymus wildryes(Gramineae:Triticeae).Genome,2003,46:627-646.
[68]Kishii M,Nagak K,Tsujimoto H.Sasakuma T.Exclusive localization of tandem repetitive sequences in subtelomeric heterochromatin regions of Leymus racemosus(Poaceae,Triticeae).Chromosome Research,1999,7:519-529.
[69]Kishii M,Tsujimoto H,Genus-specific localization of the Tail family of tandem-repetitive sequences in either the centromeric or subtelomeric regions in Triticeae species(Poaceae) and its evolution in wheat.Genome,2002,45:946-955.
[70]Kishii M,Wang R R-C,Tsujimoto H.Characteristics and behaviour of the chromosomes of Leymus mollis and L.racemosus(Poaceae,Triticeae) during mitosis and meiosis.Chromosome Research,2003,11:741-748.
[71]周新成.我国西北地区赖草的遗传多样性及赖草属8个物种基因组来源的分子进化分析.北京:中国农业科学作物科学研究所,2006.
[72]孙根楼,颜济,杨俊良.新疆小麦族植物染色体数的观察.广西植物,1990,10(2):143-148.
[73]孙振雷,黄凤兰.赖草的染色体组型及非整倍体.哲里木畜牧学院学报,1998,8(2):10-13.
[74]段晓刚等.羊草染色体组型分析.中国草原,1984,(1):63-65.
[75]孙义凯,赵毓棠,董玉琛等.东北地区小麦族11种植物的核型报道.植物分类学报,1992,30(4):342-345.
[76]阎贵兴,张素贞,云锦凤等.33种禾本科饲用植物的染色体核型研究.中国草地,1991,(5):1-13.
[77]卫俊智.新疆大赖草的细胞学初步研究.遗传,1990,12(1):12-14.
[78]段晓刚,王丽,卜秀玲.大赖草染色体组型分析.中国草地,1991,(1):45-47.
[79]智力,蔡联炳.赖草属5个种的核型与进化.西北植物学报,2000,20(5):876-878.
[80]段晓刚,卜秀玲,山蓝等.五种赖草属植物染色体组型分析.中国草地,1993,(2):44-47.
[81]潘国富,孙振雷.羊草PMC减数分裂、花粉育性及结实性的研究.中国草地,1986,(3):7-14.
[82]Allokas H.Hybrids,partial hybrids and monoploids from interspecific crosses of the Triticeae species attained by in vitro embryo culture.Yliopistopaino,Helsinki,1997,1-15.
[83]Anarnthawat-Jonsson K,Bodvarsdottir S K.Meiosis of wheat×lymegrass hybrids.Chromosome Research,1998,6:339-343.
[84]Jensen K B,Redinbaugh M,Blood M,Horton W H,Asay K H.Natural hybrids of Elymus elymoides,and Leymus salinus subsp.Salmonis(Poaceae:Triticeae).Crop Science,1999,39:976-982.
[85]王苏玲,齐莉莉,陈佩度,等.大赖草及近缘种染色体C-分带的研究.植物学报,1999,41(3):258-262.
[86]刘芳,孙根楼.赖草属七个种同工酶研究.广西植物,1997,17(1):169-173.
[87]赵茂林,贾继增,董玉琛.赖草属物种生化遗传标记的初步研究.作物品种资源,1997,(3):1-3.
[88]卢萍.羊草服酶同工酶分析.内蒙古师范大学学报,1990,(2):63-65.
[89]张丽萍,陈丽颖,李建东.东北松嫩草原羊草的过氧化物酶同工酶及酯酶同工酶的研究.草业科学,1992,9(2):49-51.
[90]陈世龙.内蒙古东部地区不同类型草原中羊草的同工酶研究.中国草地,1994,(6):47-50.
[91]汪敏,钱吉,郑师章.羊草不同地理种群的同工酶研究.应用生态学报,1998,9(3):269-272.
[92]任晓琴,陈佩度,刘大均.大赖草染色体组构成的生化分析.遗传,1996,18(4):4-7.
[93]胡志昂,王洪新.蛋白质多样性和品种鉴定.植物学报,1991,33(7):556.
[94]鲍晓明,黄百渠.小麦-冰草异附加系种子醇溶蛋白基因表达的分析.作物学报,1993,19(3):233-238.
[95]刘杰,刘公社,等.用微卫星序列构建羊草遗传指纹图谱.植物学报,2000,42(9):985-987.
[96]柴守诚,刘大均,陈佩度,等.大赖草和新麦草物种专化DNA重复序列研究Ⅱ在小麦族中分布的多态性.西北农业大学学报,1999,27(4):1-6.
[97]Dubcovsky J,Schlatter A R,Echaide M.Genome analysis of South American Elymus(Triticeae)and Leymus(Triticeae) species based on variation in repeated nucleotide sequences.Genome,1997,40(4):505-520.
[98]Anamthawat-Jonsson K,Bragason B T H,Bodvarsdottir S K.Molecular variation in Leymus species and population.Molecular Evolution,1999,8:309-315.
[99]Hole D J,Jensen K B,Wang R R-C,Clawson S M.Molecular marker analysis of Leymus flavescens and chromosome pairing in Leymus flavescens hybrids(Poaceae:Triticeae).International Journal of Plant Science,1999,160:371-376.
[100]Wang Y S,Teng X H,Huang D M,Nakamura M,Hong R M.Genetic variation and clonal diversity of the two divergent types of clonal populations of Leymus chinensis Tzvel on the Song Liao steppe in the west of northeastern China.Journal of Integrative Plant Biology,2005a,47(7):811-822.
[101]Wang Y S,Zhao L P,Wang H,Wang J,Huang D M,Hong R M,Tent X H,Nakamura M. Molecular genetic variation in a clonal plant variation in a clonal plant population of Leymus chinensis(Trin.) Tzvel.Journal of Integrative Plant Biology,2005b,47(9):1055-1064.
[102]刘惠芬,高玉葆,王丹,任安芝,阮维斌,陈磊,赵念席.内蒙古典型草原种群遗传分化的RAPD分析.生态学报,2004a,24(3):423-431.
[103]刘惠芬,高玉葆,阮维斌,陈磊,李长林,赵念席,王丹.内蒙古中东部不同草原地带羊草种群遗传分化.生态学报,2004b,24(10):2157-2164.
[104]Kuhns U,Frctz T A.Distinguishing rose culfivars by polyacrylamide gelelectrophoresis Ⅱ isoenzyme variation among cultivars.J Am Soc Hortic Sci,1978,103:509-516.
[105]段晓刚,卜秀玲.布顿大麦草Giemsa-C带研究.中国草地,1994,4:45-46,50.
[106]Lu B R,Yen C,Yang J L.Cytogenetic studies of hybrid between Psathyrostachys juncea and Psa.Huashanica(Poaceae).Nord J Bot,1989,9(1):11-14.
[107]王秀娥,李万隆,刘大均.新麦草属两个物种的C-带研究.南京农业大学学报,1998,21(1):10-13.
[108]Kesara A J.Genomic and genetic relationships among species of Leymus(Poaceae:Triticeae)inferred from 18S-26S ribosomal genes.American Journal of Botany,2001,88(4):553-559.
[109]刘静,杨瑞武,周永红,等.小麦族猬草属和近缘属物种醇溶蛋白分析.四川农业大学学报,2006,24(3):266-268.
[110]Hsiao C,Chatterton N J,Asay K H,Jensen K B.Phylogenetic relationships of the monogenomic species of the wheat tribe,Triticeae(Poacae),inferred from nuclear rDNA(internal transenbed spacer) sequences.Genome,1995,38:221-223.
[111]Hsiao C,Wang,R R C,Dewey,D R.Karyotype analysis and genome relationships of 22 diploid species in the tribe Triticeae.Can.J.Genet.Cytol,1986,287:109-120.
[112]Endo T R,Gill B S.The heterochromatin distribution and genome evolution in diploid species of Elymus and Agropyron.Can.J.Genet.Cytol,1984,26:669-678.
[113]张海琴.小麦族猬草属植物基因组组成及其分类地位研究.雅安:四川农业大学,2006.
[114]Jaccard P.Nowvelles researches ser la distribution florale.Bull Soc Vaud Sci Nat,1908,44:223-270.
[115]Wang L,Guo J,Zhao G F,Genetic diversity of the endangered and endemic species Psathyrostach.ys huashanica natural populations using simple sequence repeats(SSRs) markers.Biochemical Systematics and Ecology,2006,34:310-318.
[116]Wang R R-C,Dewey D R,Hsaio C.Intergeneric hybrids of Agropyron and Pseudoroegneria.Bot Gaz,1985,146(2):268-274.
[117]Wang R R-C.Diploid perennial intergeneric hybrids in the tribe Triticeae.1.Agropyron cristatum ×Pseudoroegneria libanotica and Critesion violaceum×Psathyrostachys juncea.Crop Sci,26:75-79.
[118]Wei J Z,Wang R R C.Genome and specific markers and genome relationship of diploid perennial species in Triticeae based on RAPD analyses.Genome,1995,38:1230-1236.
[119]Jensen K B,Wang R R C.Cytological and molecular evidence for transferring Elymus coreanus from the genus Elymus to Leymus and molecular evidence for Elymus californicus(Poaceae:Triticeae).International Journal of Plant Science 1997,158:872-877.
[2]Dewey D R.The genomic system of classification as a guide to intergeneric hybridization with the perennial Triticeae.In:Gustafson J P(eds),Gene Manipulation in plant improvement.NewYork:Plenum,1984,209-280.
[3]杨瑞武.赖草属植物的系统与进化研究.雅安:四川农业大学,2003.
[4]董玉琛.小麦野生近缘植物的研究和利用.李胜琳,马缘生主编,植物优异种质资源及其开拓利用.北京:中国科学技术出版社.1992,39-44.
[5]刘大钧.向小麦转移外源抗病性的回顾与展望.南京农业大学学报,1994a,17(3):1-7.
[6]刘大钧.外源基因在小麦育种中的利用.作物杂志,1994b,(6):1-7.
[7]Riley R,Chapman V,Johnson R.Introduction of yellow rust resistance of Aegilops comosa into wheat by genetically induced homoeologous combination.Nature,1968,217:383-384.
[8]Feldman M..Gene transfer from wild species into cultivated plants.Genetika,1983,15:145-161.
[9]Feldman M.Cytogenetics and molecular approaches to alien gene transfer in wheat.In:Miller T E,Koebner R M D(eds),Proc 7th Int Wheat Genet Symp.Cambridge,England,Cambridge:Cambridge University Press,1988,23-32.
[10]Sharma H C,Gill B S.Current Status of wide hybridization in wheat.Euphytica,1983,32:17-31.
[11]Dvorak J,Meguire P E.Triticeae,the gene pool for wheat breeding.Genome mapping of wheat and related species.Publ Workshop.Westsaerment,California,1990,3-8.
[12]Jiang J M,Friebe B,Gill B S.Recent advances in alien gene tranfer in wheat.Euphytica,1994,73:199-212.
[13]Hochstetter C F.Nachtraglicher Commentar zu meiner Abhandlung."Aufbau der Graspflanze etc."Flora 1848,7:105-118.
[14]Pilger R.Addimenta Agrostologica.Ⅰ.Triticeae(Hordeae).Bot Jahrb,1949,74:1-27.
[15]Pilger R.Das System der Graminaea.Bot Jahrb,1954,76:281-284.
[16]耿以礼.中国主要植物图说.禾本科.北京:科学出版社,1959,342-409.
[17]Tzvelev N N.Despeciebus nonnulis novis vel minus cognitise pamur.Bot Mater Gerb Bot Inst Komarova Akad Nauk SSR(Leningrad),1960,20:412-439.
[18] Tzvelev N N. Tribe 3. Triticeae Dum. Poaceae URSS. Leningrad: Nauka, 1976. 176-189.
[19] Love A, D Love. Some nomenclatural changes in the European flora. I .Species and supraspecific categories. Bot Not, 1961, 114: 33-47.
[20] Melderis A. Leymus. In: T G Tutin et al., (eds). Flora Eueopaea. vol. 5. Cambridge: Cambridge University Press, 1980, 190-192.
[21] Bentham G.. Notes on Gramineae. Bot J Linn Soc, 1881,18:14-134.
[22] Bentham G and J D Hooker. Genera plantarum. vol.3. London. 1881.
[23] Boissier E. Flora Orientalis. vol.5. H Georg Geneva. 1884
[24] Hackel E. Hordeae. In: A Engler and K Prantl (eds). Die naturlichen pflanzentamilien. W Engelmann. Leipzig. 1887, 76-89.
[25] Coste H. Flore descriptive et illustree de la France 3.1906.
[26] Hitchcock C L. Gramineae. In: Hitchcock C. L. et al., (eds.)Vascular Plants of the Pacific Northwest, vol. 1. Seattle: University of Washington Press, 1969, 384-725.
[27] Holmgren A H and N H Holmgren. Triticeae. In: Cronquist et al., (eds). Intermountain Flora, vol. 6. New York: Columbia University Press, 1977, 293-336.
[28] Estes J R, R J Tyrl. The generic concept and generic circumscription in the triticeae: an end paper. In: J R Estes et al., (eds). Grasses and Grasslands. Norman: University of Oklahoma Press, 1982, 145-164.
[29] Nevski S A. Uber das system der tribe Hordeae Benth. Flora et Systematica Plantae vasculares. Ser 1. Fasc 1. Leningrad, 1933, 9-32.
[30] Nevski S A. Tribe XIV Hordeae Benth. In: Komarov V L, Roshevits R Y, Shishkin B K (eds). Flora URSS 2, Leningrad, 1934, 590-720.
[31] Stebbins G L, Walters M S. Artificial and natural hybrids in the Gramineae, tribe Hordeae III Hybrids involving Elymus condensatus and E. triticoides. Amer J Bot, 1949, 36: 291-301.
[32] Dewey D R. Genome relations among diploid Elymus juncea and certain tetraploid and octoploid Elymus species. Amer J Bot, 1970a, 57: 633-639.
[33] Dewey D R. Cytogenetics of tetraploid Elymus cinereus, E. triticoides, E. multicaulis, E. karataviensis, and their F_1 hybrids. Bot Gaz, 1972,133: 51-57.
[34] Dewey D R. The genome constitution and phylogeny of Elymus ambiguous. Amer J Bot, 1976, 63: 626-634.
[35]Dewey D R..Genomic and phylogenetic relationships among North American Perennial Triticeae.In:J R Estes et al.,(eds).Grasses and Grasslands.Norman:University of Oklahoma Press,1982,51-88.
[36]Jaaska V.Enzyme variability and phylogenetic relationships in the grass genera Agropyron Gaertn.And Elymus L.Izv Akad Nauk Estonsk SSR Ser Biol,1974,23:3-18.
[37]Stebbins G L.Taxonomy and the evolution of genera,with special reference to the family Gramineae.Evolution,1956,10:235-245.
[38]Bowden W M.Cytotaxonomy of the species and interspecific hybrids of genus Agropyron in Canada and neighbouring areas.Can J Bot,1965,43:1421-1448.
[39]Love A.Generic evaluation of the wheatgrass.Biol Zentralbl,1982,101:199-212.
[40]Baum B R.Taxonomy of the tribe Triticeae(Poaceae) using various numerical techniques,Ⅰ.Historical perspectives,dta accumulation,and character analysis.Can J Bot,1977,55:1712-1740.
[41]Baum B R.The genus Elymus in Canada-Bowden's generic concept and key reappraised and relectotypification of E.canadenssi.Can J Bot,1979,57:946-951.
[42]Baum B R.The generic problem in the Triticeae:numerical taxonomy and related concepets,In:J R Estes et al.,(eds).Grasses and grasslands.Norman:University of oklahoma Press,1982,109-144.
[43]Barkworth M E,Atkins R J.Leymus Hochst.(Gramineae:Triticeae) in North Ameriea:Taxonomy and distribution.Amer J Bot,1984,71(5):609-625.
[44]Tzvelev N.N.Grasses of the Soviet Union.New Delhi:Oxonian Press PVT.Ltd.,1976b,253-276.
[45]Anamthawat-Jonsson K,Bodvarsdottir S K.Genomic and genetic relationships among species of Leymus(Poaceae:Triticeae) inferred from 18S-26S ribosomal genes.Amer J Bot,2001,88(4):553-559.
[46]Bodvarsdottir and Anamthawat-Jonsson.Isolation,characterization,and analysis of Leymus-specific DNA sequences.Genome,2003,46:673-682.
[47]候宽昭.中国种子植物科属辞典.北京:科学出版社,1958.
[48]候宽昭.中国种子植物科属辞典(修订版).北京:科学出版社,1982.
[49]郭本兆.中国植物志.北京:科学出版社,1987,9(3):15-26.
[50]颜济,杨俊良.中国赖草属新植物.云南植物研究,1983.5(3):275-276.
[51]崔乃然,钟骏平.新疆谷类作物的野生近缘植物.新疆八一农学院学报,1984,(1):11-35.
[52]吴玉虎.新疆赖草属二新种.植物研究,1992,12(4):343-345.
[53]孙根楼,颜济,杨俊良.新分布于中国的一种赖草形态学和细胞学观察[J].四川农业大学学报,1992,10(4):574-550.
[54]蔡联炳.国产赖草属新分类群.植物分类学报,1995,33(5):491-496.
[55]蔡联炳.赖草属资料.植物研究,1997,17(1):28-32.
[56]蔡联炳.青海赖草属一新种和一新变种.植物分类学报,2001,39(1):75-77.
[57]崔大方.新疆赖草属的新分类群.植物研究,1998,18(2):144-148.
[58]赵可夫,李法曾.中国盐生植物.北京:科学出版社,1999,331-334.
[59]Wang R R-C,Hsiao C.Morphology and cytology of interspecific hybrid of Leymus mollis,d Hered,1984,75:488-492.
[60]Zhang H B,Dvorak J.The genome origin of tetraploid of Leymus(Poaceae:triticeae) inferred from variation in repeated nucleotide sequences.Amer J Bot,1991,78:871-884.
[61]Shiotan 1.Species differentiation in Agropyron,Elymusu,Hystrix,and Sitanion.Proceedings of the 12th International Congress of Genetics.The Science Council of Japan,Tokyo.1968,184.
[62]Love A.Conspectus of the Triticeae.Feddes Repert,1984,95:425-521.
[63]Wang R R C,Jensen K B.Absence of the J genome in species(Poaceae:Triticeae):evidence from DNA hybridization and meiotic pairing.Genome,1994,37:231-235.
[64]φrgaard M.and Heslop-Harrison J.S.,Investigations of genome relationships between Leymus,Psathyrostachys and Hordeum inferred by genome DNA:DNA in situ hybridization.Annals of Botany,1994,73:195-203.
[65]Wang R.R-C,Bother V.,Dvorak J.,Linde-Laursen 1.,and Muramatsu M.Genome symbols in the Triticeae(Poaceae).In:Wang R R-C et al.(eds.),Proceedings of the 2nd International Triticeae Symposium.Utah State University Press,Logan,Utah,USA.1994,29-34.
[66]Sun G L,Wu B H,and Liu F.Cytogenetic and genomic relationships of Thinopyrurn elongatum with two Psathyrostachys species and with Leymus secaIinus Poaceae.Plant Systematics and Evolution,1995a,197:225-231.
[67]Wu X L,Larson,Hu Z M,Palazzo A J,Jones T A,Wang R R-C,Jensen K B,and Chatterton N J.Molecular genetic linkage maps for alloteterap1oid Leymus wildryes(Gramineae:Triticeae).Genome,2003,46:627-646.
[68]Kishii M,Nagak K,Tsujimoto H.Sasakuma T.Exclusive localization of tandem repetitive sequences in subtelomeric heterochromatin regions of Leymus racemosus(Poaceae,Triticeae).Chromosome Research,1999,7:519-529.
[69]Kishii M,Tsujimoto H,Genus-specific localization of the Tail family of tandem-repetitive sequences in either the centromeric or subtelomeric regions in Triticeae species(Poaceae) and its evolution in wheat.Genome,2002,45:946-955.
[70]Kishii M,Wang R R-C,Tsujimoto H.Characteristics and behaviour of the chromosomes of Leymus mollis and L.racemosus(Poaceae,Triticeae) during mitosis and meiosis.Chromosome Research,2003,11:741-748.
[71]周新成.我国西北地区赖草的遗传多样性及赖草属8个物种基因组来源的分子进化分析.北京:中国农业科学作物科学研究所,2006.
[72]孙根楼,颜济,杨俊良.新疆小麦族植物染色体数的观察.广西植物,1990,10(2):143-148.
[73]孙振雷,黄凤兰.赖草的染色体组型及非整倍体.哲里木畜牧学院学报,1998,8(2):10-13.
[74]段晓刚等.羊草染色体组型分析.中国草原,1984,(1):63-65.
[75]孙义凯,赵毓棠,董玉琛等.东北地区小麦族11种植物的核型报道.植物分类学报,1992,30(4):342-345.
[76]阎贵兴,张素贞,云锦凤等.33种禾本科饲用植物的染色体核型研究.中国草地,1991,(5):1-13.
[77]卫俊智.新疆大赖草的细胞学初步研究.遗传,1990,12(1):12-14.
[78]段晓刚,王丽,卜秀玲.大赖草染色体组型分析.中国草地,1991,(1):45-47.
[79]智力,蔡联炳.赖草属5个种的核型与进化.西北植物学报,2000,20(5):876-878.
[80]段晓刚,卜秀玲,山蓝等.五种赖草属植物染色体组型分析.中国草地,1993,(2):44-47.
[81]潘国富,孙振雷.羊草PMC减数分裂、花粉育性及结实性的研究.中国草地,1986,(3):7-14.
[82]Allokas H.Hybrids,partial hybrids and monoploids from interspecific crosses of the Triticeae species attained by in vitro embryo culture.Yliopistopaino,Helsinki,1997,1-15.
[83]Anarnthawat-Jonsson K,Bodvarsdottir S K.Meiosis of wheat×lymegrass hybrids.Chromosome Research,1998,6:339-343.
[84]Jensen K B,Redinbaugh M,Blood M,Horton W H,Asay K H.Natural hybrids of Elymus elymoides,and Leymus salinus subsp.Salmonis(Poaceae:Triticeae).Crop Science,1999,39:976-982.
[85]王苏玲,齐莉莉,陈佩度,等.大赖草及近缘种染色体C-分带的研究.植物学报,1999,41(3):258-262.
[86]刘芳,孙根楼.赖草属七个种同工酶研究.广西植物,1997,17(1):169-173.
[87]赵茂林,贾继增,董玉琛.赖草属物种生化遗传标记的初步研究.作物品种资源,1997,(3):1-3.
[88]卢萍.羊草服酶同工酶分析.内蒙古师范大学学报,1990,(2):63-65.
[89]张丽萍,陈丽颖,李建东.东北松嫩草原羊草的过氧化物酶同工酶及酯酶同工酶的研究.草业科学,1992,9(2):49-51.
[90]陈世龙.内蒙古东部地区不同类型草原中羊草的同工酶研究.中国草地,1994,(6):47-50.
[91]汪敏,钱吉,郑师章.羊草不同地理种群的同工酶研究.应用生态学报,1998,9(3):269-272.
[92]任晓琴,陈佩度,刘大均.大赖草染色体组构成的生化分析.遗传,1996,18(4):4-7.
[93]胡志昂,王洪新.蛋白质多样性和品种鉴定.植物学报,1991,33(7):556.
[94]鲍晓明,黄百渠.小麦-冰草异附加系种子醇溶蛋白基因表达的分析.作物学报,1993,19(3):233-238.
[95]刘杰,刘公社,等.用微卫星序列构建羊草遗传指纹图谱.植物学报,2000,42(9):985-987.
[96]柴守诚,刘大均,陈佩度,等.大赖草和新麦草物种专化DNA重复序列研究Ⅱ在小麦族中分布的多态性.西北农业大学学报,1999,27(4):1-6.
[97]Dubcovsky J,Schlatter A R,Echaide M.Genome analysis of South American Elymus(Triticeae)and Leymus(Triticeae) species based on variation in repeated nucleotide sequences.Genome,1997,40(4):505-520.
[98]Anamthawat-Jonsson K,Bragason B T H,Bodvarsdottir S K.Molecular variation in Leymus species and population.Molecular Evolution,1999,8:309-315.
[99]Hole D J,Jensen K B,Wang R R-C,Clawson S M.Molecular marker analysis of Leymus flavescens and chromosome pairing in Leymus flavescens hybrids(Poaceae:Triticeae).International Journal of Plant Science,1999,160:371-376.
[100]Wang Y S,Teng X H,Huang D M,Nakamura M,Hong R M.Genetic variation and clonal diversity of the two divergent types of clonal populations of Leymus chinensis Tzvel on the Song Liao steppe in the west of northeastern China.Journal of Integrative Plant Biology,2005a,47(7):811-822.
[101]Wang Y S,Zhao L P,Wang H,Wang J,Huang D M,Hong R M,Tent X H,Nakamura M. Molecular genetic variation in a clonal plant variation in a clonal plant population of Leymus chinensis(Trin.) Tzvel.Journal of Integrative Plant Biology,2005b,47(9):1055-1064.
[102]刘惠芬,高玉葆,王丹,任安芝,阮维斌,陈磊,赵念席.内蒙古典型草原种群遗传分化的RAPD分析.生态学报,2004a,24(3):423-431.
[103]刘惠芬,高玉葆,阮维斌,陈磊,李长林,赵念席,王丹.内蒙古中东部不同草原地带羊草种群遗传分化.生态学报,2004b,24(10):2157-2164.
[104]Kuhns U,Frctz T A.Distinguishing rose culfivars by polyacrylamide gelelectrophoresis Ⅱ isoenzyme variation among cultivars.J Am Soc Hortic Sci,1978,103:509-516.
[105]段晓刚,卜秀玲.布顿大麦草Giemsa-C带研究.中国草地,1994,4:45-46,50.
[106]Lu B R,Yen C,Yang J L.Cytogenetic studies of hybrid between Psathyrostachys juncea and Psa.Huashanica(Poaceae).Nord J Bot,1989,9(1):11-14.
[107]王秀娥,李万隆,刘大均.新麦草属两个物种的C-带研究.南京农业大学学报,1998,21(1):10-13.
[108]Kesara A J.Genomic and genetic relationships among species of Leymus(Poaceae:Triticeae)inferred from 18S-26S ribosomal genes.American Journal of Botany,2001,88(4):553-559.
[109]刘静,杨瑞武,周永红,等.小麦族猬草属和近缘属物种醇溶蛋白分析.四川农业大学学报,2006,24(3):266-268.
[110]Hsiao C,Chatterton N J,Asay K H,Jensen K B.Phylogenetic relationships of the monogenomic species of the wheat tribe,Triticeae(Poacae),inferred from nuclear rDNA(internal transenbed spacer) sequences.Genome,1995,38:221-223.
[111]Hsiao C,Wang,R R C,Dewey,D R.Karyotype analysis and genome relationships of 22 diploid species in the tribe Triticeae.Can.J.Genet.Cytol,1986,287:109-120.
[112]Endo T R,Gill B S.The heterochromatin distribution and genome evolution in diploid species of Elymus and Agropyron.Can.J.Genet.Cytol,1984,26:669-678.
[113]张海琴.小麦族猬草属植物基因组组成及其分类地位研究.雅安:四川农业大学,2006.
[114]Jaccard P.Nowvelles researches ser la distribution florale.Bull Soc Vaud Sci Nat,1908,44:223-270.
[115]Wang L,Guo J,Zhao G F,Genetic diversity of the endangered and endemic species Psathyrostach.ys huashanica natural populations using simple sequence repeats(SSRs) markers.Biochemical Systematics and Ecology,2006,34:310-318.
[116]Wang R R-C,Dewey D R,Hsaio C.Intergeneric hybrids of Agropyron and Pseudoroegneria.Bot Gaz,1985,146(2):268-274.
[117]Wang R R-C.Diploid perennial intergeneric hybrids in the tribe Triticeae.1.Agropyron cristatum ×Pseudoroegneria libanotica and Critesion violaceum×Psathyrostachys juncea.Crop Sci,26:75-79.
[118]Wei J Z,Wang R R C.Genome and specific markers and genome relationship of diploid perennial species in Triticeae based on RAPD analyses.Genome,1995,38:1230-1236.
[119]Jensen K B,Wang R R C.Cytological and molecular evidence for transferring Elymus coreanus from the genus Elymus to Leymus and molecular evidence for Elymus californicus(Poaceae:Triticeae).International Journal of Plant Science 1997,158:872-877.