小麦苗期抗旱相关形态指标的灰色关联度分析和Lr1基因在小麦骨干亲本衍生系的分布
|
摘要
干旱是世界范围内限制作物生产的主要非生物逆境之一,在我国北方,大部分小麦主产区干旱发生频繁,小麦生长发育进程中时常受到胁迫,严重影响粮食安全生产。培育和选择抗旱品种是解决干旱地区水资源不足和干旱频繁发生的主要途径之一,所以及时准确地鉴定、评价和筛选小麦品种的抗旱性是进行小麦抗旱育种和筛选抗旱小麦品种的基础。为了较系统研究小麦苗期形态指标与小麦抗旱性的关系,筛选有效的抗旱性指标,本研究使用聚乙二醇(PEG)-6000模拟干旱胁迫,采用灰色关联度分析法,分析了28份小麦品种苗期干旱胁迫下的苗高、最长根长、根数、地上部鲜重、地上部干重、根鲜重、根干重、叶片数、根冠比等9个形态指标与抗旱系数的关联程度,各指标与抗旱系数的关联度依次为：叶片数(0.817)>苗高(0.766)>地上部干重(0.746)>地上部鲜重(0.729)>根数(0.699)>根干重(0.688)>根鲜重(0.681)>根冠比(0.645)>最长根长(0.399)。分析表明,9个形态指标可分为叶片相关因子、地上部生物量因子和根部生物量因子三大类,地上部生物量因子的关联度大于根部生物量因子。同时利用主成分分析法对9个形态指标进行了分析,也得到相似结论。聚类结果能较好的反映各小麦品种的地域分布。
由小麦叶锈菌(Puccinia triticina)引起的小麦叶锈病是世界各产麦区一种普遍的病害,它会导致作物减产和作物品质的下降。利用抗病品种是一种最经济、有效和最安全的控制叶锈病流行的措施,小麦抗叶锈病Lr1基因是一种在世界小麦中分布较广的基因,含有Lr1抗性基因的小麦在接种后表现一种典型的超敏反应。本研究通过PCR分子标记技术检测四种小麦骨干亲本(南大2419,阿夫,燕大1817,碧玛4号)的328种衍生系品种的Lr1基因分布。结果表明：在所有测试品种中,Lr1基因的频率燕大1817(80%)>阿夫(78.1%)>南大2419(54.2%)>碧玛4号(32.4%)。Lr1基因在各个亲本衍生系的最高频率分别是南大2419子五代100%,燕大1817子三代100%,阿夫子三代90%,碧玛4号子三代45%。Lr1基因的分布频率呈现出从冬小麦区到春小麦区的上升趋势。
Grey Correlation Analysis of Morphological Traits Related to Drought Tolerance of Wheat at Seeding Stage and Distribution of Lrl gene in Derivatives from Core Chinese Wheat Parents
Drought is one of the main non-biological stress which is restricted production in the world. Drought is the main yield limiting factor in most parts of North Area, So the key road is choosing and breeding drought resistance variety. To cultivate and demonstrate good drought resistance variety has important meaning of saving water resource and evaluating wheat varieties drought resistance index exactly is the base of drought resistance wheat breeding and choosing. In this study,it aims at probing the relationship between morphological traits and drought tolerance of wheat cultivars and screening effective indexes for drought resistance. A total of 9 morphological traits including plant height, main root length, root number, shoot fresh weight, shoot dry weight, root fresh weight,root dry weight,leaf number,root/shoot ratio that measured in 28 wheat cultivars grown at laboratory drought stress by PEG-6000, were analyzed using grey relational grade. At the some time,cultivars were classified by using clustering analysis base on weight drought-tolerance coefficient. Grey correlation degrees were calculated between each trait and its drought coefficient and ranked as follows:leaf number (0.817)>plant height (0.766)>shoot dry weight (0.746)>shoot fresh weight (0.729)>root number (0.699) >root dry weight (0.688)> root fresh weight (0.681)>root/shoot ratio (0.645)>main root length (0.399). The result show that 9 morphological traits could be classified 3 categories. The cluster consequence dovetailed fundamentally with source and area distribution of cultivars.
Wheat leaf rust caused by fungal pathogen (Puccinia triticina) is a severe disease worldwide that is responsible for major crop damage and results in both yield losses and down grading in quality. The use of resistance variety is the most effective, economical and ecological method to control epidemics of leaf rust disease. The leaf rust disease resistance gene 1(Lrl) is present in a number of wheat cultivars. Plants containing the Lrl resistance gene show a typical hypersensitive reaction after infection with an avirulent pathogen race. In this study,The presence of Lrl in four core wheat parents (Mentana,Funo,Yan Da 1817 and Bi Ma 4) and 328 derivatives from these parents were determined using PCR based molecular method. Frequencies of the Lr1 genes in these derivatives were:Yan Da 1817 (80%)> Funo (78.1%)> Mentana (54.2%)> BiMa 4 (32.4%). The highest frequencies of Lr1 gene in Funo derivatives is 90%(Third of Funo),The highest frequencies of Lr1 gene in Yan Da 1817derivatives is 100%(Third of Yan Da 1817),The highest frequencies of Lr1 gene in Mentana derivatives is 100%(Fifth of Mentana),The highest frequencies of Lr1 gene in Bi Ma 4 derivatives is 45%(Third of BiMa 4). Frequencies of Lr1 loci increased in the derivatives from winter wheat zones to spring winter zones.
由小麦叶锈菌(Puccinia triticina)引起的小麦叶锈病是世界各产麦区一种普遍的病害,它会导致作物减产和作物品质的下降。利用抗病品种是一种最经济、有效和最安全的控制叶锈病流行的措施,小麦抗叶锈病Lr1基因是一种在世界小麦中分布较广的基因,含有Lr1抗性基因的小麦在接种后表现一种典型的超敏反应。本研究通过PCR分子标记技术检测四种小麦骨干亲本(南大2419,阿夫,燕大1817,碧玛4号)的328种衍生系品种的Lr1基因分布。结果表明：在所有测试品种中,Lr1基因的频率燕大1817(80%)>阿夫(78.1%)>南大2419(54.2%)>碧玛4号(32.4%)。Lr1基因在各个亲本衍生系的最高频率分别是南大2419子五代100%,燕大1817子三代100%,阿夫子三代90%,碧玛4号子三代45%。Lr1基因的分布频率呈现出从冬小麦区到春小麦区的上升趋势。
Grey Correlation Analysis of Morphological Traits Related to Drought Tolerance of Wheat at Seeding Stage and Distribution of Lrl gene in Derivatives from Core Chinese Wheat Parents
Drought is one of the main non-biological stress which is restricted production in the world. Drought is the main yield limiting factor in most parts of North Area, So the key road is choosing and breeding drought resistance variety. To cultivate and demonstrate good drought resistance variety has important meaning of saving water resource and evaluating wheat varieties drought resistance index exactly is the base of drought resistance wheat breeding and choosing. In this study,it aims at probing the relationship between morphological traits and drought tolerance of wheat cultivars and screening effective indexes for drought resistance. A total of 9 morphological traits including plant height, main root length, root number, shoot fresh weight, shoot dry weight, root fresh weight,root dry weight,leaf number,root/shoot ratio that measured in 28 wheat cultivars grown at laboratory drought stress by PEG-6000, were analyzed using grey relational grade. At the some time,cultivars were classified by using clustering analysis base on weight drought-tolerance coefficient. Grey correlation degrees were calculated between each trait and its drought coefficient and ranked as follows:leaf number (0.817)>plant height (0.766)>shoot dry weight (0.746)>shoot fresh weight (0.729)>root number (0.699) >root dry weight (0.688)> root fresh weight (0.681)>root/shoot ratio (0.645)>main root length (0.399). The result show that 9 morphological traits could be classified 3 categories. The cluster consequence dovetailed fundamentally with source and area distribution of cultivars.
Wheat leaf rust caused by fungal pathogen (Puccinia triticina) is a severe disease worldwide that is responsible for major crop damage and results in both yield losses and down grading in quality. The use of resistance variety is the most effective, economical and ecological method to control epidemics of leaf rust disease. The leaf rust disease resistance gene 1(Lrl) is present in a number of wheat cultivars. Plants containing the Lrl resistance gene show a typical hypersensitive reaction after infection with an avirulent pathogen race. In this study,The presence of Lrl in four core wheat parents (Mentana,Funo,Yan Da 1817 and Bi Ma 4) and 328 derivatives from these parents were determined using PCR based molecular method. Frequencies of the Lr1 genes in these derivatives were:Yan Da 1817 (80%)> Funo (78.1%)> Mentana (54.2%)> BiMa 4 (32.4%). The highest frequencies of Lr1 gene in Funo derivatives is 90%(Third of Funo),The highest frequencies of Lr1 gene in Yan Da 1817derivatives is 100%(Third of Yan Da 1817),The highest frequencies of Lr1 gene in Mentana derivatives is 100%(Fifth of Mentana),The highest frequencies of Lr1 gene in Bi Ma 4 derivatives is 45%(Third of BiMa 4). Frequencies of Lr1 loci increased in the derivatives from winter wheat zones to spring winter zones.
引文
[1]Abbate P E,Dardanelli J L,Cantarero M G,et al.Climatic and water availability effects on water use efficiency in wheat[J].Crop Sci,2004,44(2):474～483
[2]赵济,陈传康.中国地理[M].北京：高等教育出版社,1999：9～10.
[3]倪胜利,张国宏,李兴茂.旱地小麦育种的研究机理与进展[J].现代农业科技,2007,24：102～105
[4]吴永成,周顺利,王志敏.小麦与抗旱性有关的根系遗传改良研究进展[J].麦类作物学报,2004,24(3)：101～104
[5]郝艳军,山仑.高粱抗旱机理的研究进展[J].中国农学通报,2004,20(3)：120～123
[6]欧巧明,倪建福,马瑞君.春小麦根系木质部导管与其抗旱性的关系[J].麦类作物学报,2005,25(3)：27～31
[7]肖玲,赵先贵.渭北旱塬小麦根系的再生能力及其意义[J].水土保持通报,1996,16(2)：55～57
[8]杨静慧,杨焕庭.苹果树植物叶片角质层厚度与植物抗旱性[J].天津农学院学报,1996,3(3)：27～28
[9]蒋高明.植物生理生态[M].北京,高等教育出版社,2004：191～192
[10]韦梅琴,李军乔.委陵菜属四种植物茎叶解剖结构的比较研究[J].青海师范学院学报,2003,(3)：48～50.
[11]Lupton FGH.小麦育种的理论基础[M].北京,北京农业大学出版社,1988,346～348
[12]张跃强,樊哲儒,王延军,等.小麦抗旱节水研究进展[J].新疆农业科学,2007,44(S3)：36～43
[13]卢振民.冬小麦近轴和远轴叶面气孔对土壤水分胁迫反应的敏感性[J],植物生理学报,1988,14(3)：223～227
[14]高彦萍,冯莹,马志军,等.水分胁迫下不同抗旱类型大豆叶片气孔特性变化研究[J].干旱地区农业研究,2007,25(2)：77～79
[15]王玮.植物对水分亏缺的某些生化反应[J].植物生理学通讯,1998,34(5)：388～393
[16]郑敏娜,李向林,万里强,等.水分胁迫对6种禾草叶绿体、线粒体超微结构及光合作用的影响[J].草地学报,2009,17(5)：643～649
[17]Singh B,Usha K.Salicylic acid induced physiological and biochemical changes in wheat seedlings under water stress[J].Plant Growth Regulation,2003,39(2):137～141
[18]Cohen I,Knopf J A,Irihimovitch V,et al.Aproposed mechanism for the inhibitory effects of oxidative stress on rubisco assembly and it s subunit expression[J].Plant Physiol,2005,137:376～387
[19]Shinozaki K,Yamaguchi-Shinazaki K.Gene expression and signal transduction in water-stress response[J].Plant Physiol,1997,115:327～334
[20]Mattioni C.Lacerenza N G,Troccoli A,et al.Water and salt stress induced alterations on praline metabolism of triticum durum seedlings[J].Physiol Plantarum,1997,101(4):787～792
[21]Nayyar H,Walia D P. Water stress induced proline accumulation in contrasting wheat genotypes as affected by calcium and abscisic acid[J].Biologia Plantarum,2003,46:275～279
[22]邵宏波,梁宗锁,邵明安.小麦抗旱生理生化和分子生物学研究进展与趋势[J].草业学报,2006,15(3)：5～17
[23]Takashi Hibino,Yu-Ling Meng,Yoshinobu Kawamitsu,et al.Molecular cloning and functional characterization of two kinds of betaine-aldehyde dehydrogenase in betaine-accumulating mangrove Avicennia marina (Forsk.) Vierh [J].Plant Mol Bio,2001,45:353～363
[24]Munns R,Bracely c J,Barlow E W R.Solute accumulation in the apex and leaves of wheat during water stress[J].Aus J Plant Physiol,1979 (6):379～389
[25]汤章城.逆境条件下植物脯氨酸的累积及其可能的意义[J].植物生理学通讯,1984(1)：15～17
[26]Bray E A.Stress induced gene expression in plants[M].Switzerland,Harwood Academic publishers GmbH Switzerland,1994,1～23
[27]刘春玲,陈慧萍,刘娥娥,等.水稻品种对几种逆境的多重耐性及与ABA的关系[J].作物学报,2003,29(5)：725～729
[28]Yang H Q,Jia W S.Water stress induced ABA accumulated by protein phosphorylation participating in roots of Malus rupenensis Rehd[J].Chinese Sci Bulletin,2001,46 (1):50～53
[29]余叔文,汤章城.植物生理与分子生物学[M].北京,科学出版社,1999：476～492
[30]Chen N,Goodwin PH,Hsiang T.The role of ethylene during the infection of Nicotiana tabacum by Colletotrichum destructivum[J]. J Exp Bot,2003,54(392):2449～2456
[31]郭丽红,王定康,杨晓虹,等.外源乙烯利对干旱胁迫过程中玉米幼苗某些抗逆生理指标的影响[J].云南大学学报(自然科学版),2004,6(4)：352～356
[32]Ronald Pierik,Rashmi Sasidharan,Laurentius A C J Voesenek.Growth control by ethylene: Adjusting phenotypes to the environment[J]. Plant Growth Regul,2007,26:188～200.
[33]段喜华,孙莲慧,郭晓瑞,等.拟南芥乙烯突变体在干旱胁迫下的形态学差异[J].植物研究,2009,29(1)：39～42
[34]Fan T L,Balta M,Rudd J,Payne W A.Canopy temperature depression as a potential selection criterion for drought resistance in wheat[J].Agri Sci in China,2005,4:793～800
[35]Zhang J X,Kirham M B.Drought stress induced changes in activities of superoxide dismutase,catalase,and perexidase in wheat species [J].Plant Cell Physiol,1994,35 (5):785～791
[36]Hieng B,Ugrinovic K,Sustar-Vozlic J,et al.Different classes of proteases are involved in the response to drought of Phaseolus vulgaris L.cultivars differing in sensitivity [J].Plant Physiol,2004,161:519～530
[37]Badger M R.Photosynthetic oxygen exchanges[J].Plant Physiol,1985,36:27～53
[38]张喜英,裴东,曲懋正.几种作物的生理指标对土壤水分变动的阈值反应[J].植物生态学报,2000,24(3)：280～283
[39]景蕊莲,作物抗旱研究的现状与思考[J].干旱地区农业研究,1999,17(2)：79～85
[40]周桂莲,杨慧霞.小麦抗旱性鉴定的生理生化指标及其分析评价[J].干旱地区农业研究,1996,14(2)：65～69
[41]张林刚,邓西平.小麦抗旱性生理生化研究进展[J].干旱地区农业研究,2000,18(3)：125～132
[42]张娟,张正斌,谢惠民,等.小麦叶片水分利用效率及相关生理性状的关系[J].作物学报,2005,31(12)：1593～1599
[43]蔡晓莉,吴振录,樊哲儒,等.新疆春小麦主栽品种和新品种的抗旱性研究初报[J].新疆农业科学,2005,42(2)：70～76
[44]林关石.谷子品种抗旱性观察[J].陕西农业科学,1984,4：17～19
[45]刘学义.大豆抗旱性评定方法探讨[J].中国油料,1986,4：23～26
[46]兰巨生.作物抗旱指数的概念和统计方法[J].华北农学报,1990,5(2)：20～25
[47]兰巨生.农作物综合抗旱性评价方法的研究[J].西北农业学报,1998,7(3)：85～87
[48]栗雨勤.高产抗旱冬小麦新品种鉴定筛选方法研究[J].华北农学报,1998,13：45～49
[49]路贵和,安海润.作物抗旱性鉴定方法与指标研究进展[J].山西农业科学,1999,27(4)：39～43
[50]冀天会,张灿军,杨子光,等.小麦抗旱性鉴定方法及评价指标研究Ⅲ-综合评价指标的比较研究[J].中国农学通报,2007,23(9)：226～230
[51]张灿军,冀天会,杨子光,等.小麦抗旱性鉴定方法及评价指标研究Ⅱ-抗旱性鉴定评价技术规程[J].中国农学通报,2007,23(10)：418～421
[52]胡荣海.农作物资源的抗旱筛选技术及其应用[J].农牧情报研究,1989(1)：36～41
[53]胡荣海,昌小屏.反复干旱法的生理基础及其应用[J].华北农学报,1996,11(3)：51～56
[54]黎裕.植物抗旱育种及其进展[J].世界农业,1994,3：18～20
[55]周该生.水稻抗旱性苗期鉴定指标筛选与节水机理研究[D].武汉,华中农业大学,2006
[56]刘思峰.灰色系统理论及其应用(第三版)[M].北京,2004,科学出版社
[57]扬子光,王书子,吴少辉,等.灰色关联分析在小麦品种抗旱性鉴定上的应用[J].中国农学通报,2002,18(2)：42～43
[58]冀天会,张灿军,谢惠民.小麦品种抗旱性研究的产量鉴定指标的比较研究[J].中国农学通报,2006,22(1)：103～106
[59]王士强,胡银岗,佘奎军,等.小麦抗旱相关农艺性状和生理生化性状的灰色关联度分析[J].中国农业科学,2007,40(11)：2452～2459
[60]李玉发,何中国,李淑芳,等.东北地区春小麦主要性状与产量间的灰色关联分析[J].麦类作物学报,2005,25(1)：139～141
[61]裘敏,魏亦勤,刘旺清,等.小麦5个产量性状与产量的灰色关联度分析[J].安徽农业科学,2007,35(2)：390
[62]韩启秀.运用灰色关联度对山东小麦新品种(系)综合表现的评价[J].中国农学通报,2005,21(2)：312～314
[63]高素玲,曹雯梅.应用灰色系统理论综合评价春性小麦新品种[J].河南农业科学,2006(4)：31～33
[64]Komer J A.Genetics of resistance to wheat leaf rust[J].Annual Review Phytopathology,1996, 34:435～455
[65]Kloppers F J,Pretorius Z A.Effects of combinations among genes Lrl3,Lr34 and Lr37 on components of resistance in wheat to leafrust[J].Plant Pathology,1997,46:737～750
[66]Knott D R.The wheat rust breeding for resistance[J].Theor Appl Genet,1989:191～198
[67]Friebc B,Zeller F J, Mukai Y, et al.Characterization of rust resistant wheat Agropyrom intermedium derivatives by C-banding in situ hybridization and isozyme analysis[J].Theor Appl Genet,1992,83:775～782
[68]R A.McIntosh,J Dubcovsky,W J Rogers,et al.Catalogue of gene symbols for wheat:2009 supplement[R].Annual Wheat Newsl,2009,55
[69]Kilpatrick RA.New wheat cultivars and longevity of rust resistance,1971～1975[M].Beltsvill e,MD:US Department of Agriculture,Agricultural Research Service,1975
[70]Etienne Duveiller, Ravi P. Singh, Julie M. Nicol. The challenges of maintaining wheat productivity:pests, diseases, and potential epidemics[J]. Euphytica,2007,157(3):417～430
[71]Eriksen L,Afshari F,Christiansen M J,el al.Yr32 for resistance to stripe rust present in the wheat cultivar Carstens V[J].Theor Appl Genet,2004,108:567～575
[72]McIntosh R A,Lagudah E S.Cytogenetical studies in wheat Ⅷ.Gene Yr24 for resistance to stripe rust[J].Plant Breed,2000,119:81～83
[73]Wan A M,Zhao Z H,Chen X M,et al,Wheat stripe rust epidemic and virulence of Puccinia striiformis f.sp.tritici in China in 2002[J].Plant Dis,2004,88:896～904
[74]Oelke L M, Kolmer J A.Genetics of leaf rust resistance in spring wheat cultivars Alsen and Norm [J].Phytopathology,2005,95:773～778
[75]曹张军,井金学,王美南,等.小麦品种贵农22号抗条锈基因遗传分析[J].西北植物学报,2004,24(6)：991～996
[76]代君丽,牛永春.小麦农家品种大籽糙抗条锈性的遗传分析[J].遗传,2003,25(3)：311～313
[77]徐世昌,张敬原,赵文生,等.小麦京核891-1抗条锈病主效、微效基因的遗传分析[J].中国农业科学,2001,34(3)：272～276
[78]谢皓,牛永春,陈孝,等.小麦新品系YW243抗条锈性鉴定和遗传分析[J].植物病理学报,2003,33(3)：243～247
[79]陈万权,胡长程,晏思自,等.冬小麦品种北京837抗叶锈病基因的染色体定位研究[J].植物病理学报,1995,25(4)：297～300
[80]薛秀庄,吉万全,王秋英,等.小麦性状的缺体分析[J].西北农业学报,1995,4(4)：1～5
[81]Flor H H. The complementary genetics systems in flax and flax rust[J].Advances in Genetics, 1956,8:29～54
[82]Loegering W Q,McIntosh R A,Burton C H.Computer analysis of disease data to derive hy pothetical genotypes for reaction of host varieties to pathogens[J].Can J Genet Cytni,1971,13:742～747
[83]Browder L E.Designation of two genes for resistance to Puccinia recondite in triticum aestivum[J].Crop Sci,1972,12:705～706
[84]Long D L,Schafer J F,Roelfs A P,et al. Specifici virulence of Puccinia Recondite f sp.Trifici in the United States for 1979～1983[J].Plant Dis,1985,9:343～347
[85]Dubin H J, Johnson R,Stubbs R W.Postulated genes for resistance to strip rust in selected CIMMYT and related wheats [J].Plant Dis,1989,73(6):472～475
[86]Johnson R,Stubbs RW,Kirmani MAS,et al.Discussion of a method resulting in erroneous postulation of the gene Yr8 for resistance to Puccinia striiformis in Pakistan wheat cultivars[J]. Cereal Rusts and Powdery Mildews Bulletin,1987,15:13～19
[87]Zoldan S M,Barcellos A L.Postulation of genes (Lr) for resistance to leaf rust in Brazilian wheat cultivars[J].Fitopatol Bras,2002,27(5):508～516
[88]陈万权,王剑雄.76个小麦种质资源抗叶锈及秆锈基因初步分析[J].作物学报,1997,23(6)：655～663
[89]王奎荣,陈万权,冯国华,等.黄淮麦区小麦品种抗叶锈病基因分析[J].江苏农业学报,1997,13(3)：167～170
[90]Zhao Huan-huan, Singh R P,Huerta-Espino Julio,et al. Resistance to leaf rust in forty-three Chinese wheat cultivars and line[J].ActaPhytopathologica Sinica,2006,36(6):539～549
[91]伍少云,奉有壁,汤翠凤.云南小麦资源创新种质的抗锈性与生态适应性鉴定研究[J].麦类作物学报.2002,22(3)：25～30
[92]Singh R P,Gupta A K.Genes for leaf rust resistance in Indian and Pakistan wheat resistance tested with Mexican pathotypes of Puccinia recondita f.sp. tritici[J].Euphytica,1991,5 7:27～36
[93]Winzeler M,Winzeler H,Keller B,et al.Endopestidase polymorphism and linkage of the EP-Dlc null allele with the Lr19 leaf rust resistance gene in hexaploid wheat[J].Plant Breeding,1995,114:24～28
[94]Feuillet C,Messmer M,Schachenmayr G,et al.Genetic and physical characterization of the Lrl leaf rust resistance locus in wheat(Triticum aestivum L.) [J].Mol Gen Genet,1995,248:553～562
[95]Qiu JW,Schurch AC,Yahiaoui N,et al.Physical mapping and identification of a candidate for the leaf rust resistance gene Lrl of wheat[J].Theor Appl Genet,2007,115:159～168
[96]张娜,闫红飞,张英春,等.小麦抗叶锈病基因Lr2c的SSR标记[J].农业生物技术学报,2009,17(1)：148～152
[97]Sacco F, Suilrez E Y, Naranjo T. Mapping of the leaf rust resistance gene Lr3 on chromosome 6B of Sinvalocho MA wheat[J].Genome,1998,41(5):686～690
[98]Gupta S K, Charpe A, Koul S, et al.Development and validation of molecular markers linked to an Aegilops umbellulata-derived leaf rust resistance gene, Lr9, for marker assisted selection in bread wheat[J].Genome,2005,48(5):823～830
[99]Schachermayr G,Feuillet C,Keller B.Molecular markers for the detection of the wheat leaf rust resistance gene Lr10 in diverse genetic backgrounds[J].Mol Breeding,1997,9(3):65～74
[100]Feuillet C,Schachermayr G,Keller B,et al.Molecular cloning of a new receptor-like kinase gene encoded at the Lr10 disease resistance locus of wheat[J].The Plant Journal,1995,11(1):45～52
[101]Nils S,Catherine F,Thomas W,et al.Subgenome chromosome walking in wheat:A 450-kb physical contig in Triticum monococcum L.spans the Lr10 resistance locus in hexaploid wheat(Triticum astivum L.)[J].PNAS,2000,97:13436-13441
[102]Seyfarth R,Feuillet C,Schachermayr G,et al.Molecular mapping of the adult-plant leaf rust resistance gene Lr13 in wheat(Triticum astivum L.) [J].Genet Breed,2000,54:193～198
[103]Prins R J,Groene wald Z G,Marais F,et al.AFLP and STS tagging of Lr19, a gene conferring resistance to leaf rust in wheat[J].Theor Appl Genet,2001,103:618-624
[104]李星,杨文香,李亚宁,等.小麦抗叶锈病基因Lr19的SSR标记[J].中国农业科学,2005,38(6)：1156-1159
[105]刘雅辉,闫红飞,杨文香,等.小麦抗叶锈病基因Lr19的SRAP标记[J].华北农学报,2007,2007,22(4)：193～196
[106]闫红飞,杨文香,陈云芳,等.偃麦草属E染色体组特异SCAR标记对Lr19的特异性和稳定性研究[J].植物病理学报,2009,39(1)：76～81
[107]Neu C H,Stein N,Keller B.Genetic mapping of the Lr20-pml resistance locus reveals suppressed recombination on chromosome arm 7AL in hexaploid wheat[J].Genome,2002,45:737～744
[108]Huang L,Gill B S.An RGA-like marker detects all known Lr21 leaf rust resistance gene family members in Aegilops tauschii and wheat[J].Theor Appl Genet,2001,103:1007～1013
[109]Schachermayr G M,Messmer M M, Feuillet C,et al.Identification of molecular markers linked to the Agropyron elongatum-derived leaf resistance gene Lr24 in wheat[J].Theor Appl Genet,1995,90:982～990
[110]Dedryver F,Jubier M F,Thouvenin J,et al.Molecular markers linked to the rust resistance genes Lr24 in different wheat cultivars[J].Genome,1996,39(5):830～835
[111]张娜,陈玉婷,李亚宁,等.小麦抗叶锈病基因Lr24的一个新STS标记[J].作物学报,2008,34(2)：212～216
[112]Procunier J D,Townley-Smith T,Fox S,et al.PCR-based RAPD/DGGE markers linked to leaf rust resistance genes Lr29 and Lr25 in wheat(Triticum aestivum L.)[J].Genet Breeding,1995,49:87～ 92
[113]Chai J F,Zhou R H,Jia J Z,et al.Development and application of a new codominant PCR marker for detecting 1BL/1RS wheat-rye chromosome translocations[J].Plant Breed,2006,125:302～304
[114]Nelson J C,Singh R P,Autrique E,et al.Mapping genes conferring and suppressing leaf rust resistance in wheat[J].Crop Science,1997,37:1928～1935
[115]Naik S,Gill K S,Prakasa Rao V S,et al.Identification of a STS marker linked to the Aegilops speltoides-derived leaf rust resistance gene Lr28 in wheat[J].Theor Appl Genet,1998,97:535～540
[116]Yogesh V,Parveen C,Rippy S,et al.Tagging of Aegilops speltoides-derived leaf rust resistance gene Lr28 with a microsatellite marker in wheat[J].Plant Biochemistry Biotechnology, 2004,13:47～49
[117]Tar M, Punnhauser L, Csosz L, et al. Identification of molecular markers for an efficient leaf rust resistance gene Lr29 in wheat[J]. Acta Biologica Szegediensis,2002,46:133～134
[118]Autrique E,Singh R P, Tanksley S D,et al.Molecular markers for four leaf rust resistance genes introgressed into wheat from wild relatives[J].Genome,1995,38:75～83
[119]E S Lagudah, H. McFadden, R P Singh,et al. Molecular genetic characterization of the Lr34/Yr18 slow rusting resistance gene region in wheat[J].Theor Appl Genet,2006,114(1):21～ 30
[120]Evans S. Lagudah, Simon G. Krattinger, Sybil Herrera-Foessel, et al.Gene-specific markers for the wheat gene Lr34|Yr18|Pm38 which confers resistance to multiple fungal pathogens[J]. Theor Appl Genet,2009,119(5):889～898
[121]Seyfarth R, Feuillet C, Schachermayr Q,et al.Development of a molecular for the adult plant leaf rust resistance gene Lr35 in wheat[J]. Theor Appl Genet,1999,99:554～560
[122]Gold J,Harder D,Townley Smith F,et al.Development of a molecular marker for rust resistance genes Sr39 and Lr35 in wheat breeding lines[J].Electronic Joural of Biotechnology, 1999,2(1):45～52
[123]李丽娜.小麦抗叶锈基因Lr35的AFLP和SSR分子标记[D].保定,河北农业大学,2005
[124]Robert O, Abelard C, Dedryver F,et al.Identification of molecular markers for the wheat of the yellow rust resistance gene Yr17 in wheat[J].Mol Breed,1999,5:167～175
[125]Seah S,Bariana H,Jahier J,et al.The introgressed segment carry rust resistance genes Yrl7, Lr37 and Sr38 in wheat can be assayed by a cloned disease resistance gene-like sequence[J]. Theor Appl Genet,2001,102:600～605
[126]张立荣,徐大庆,杨文香,等.小麦抗叶锈基因Lr37 ISSR分子标记[J].农业生物技术学 报,2004,1：86～89
[127]杨文香Lr37,Lr44 AFLP分子标记及124个小麦品种(系)抗叶锈基因鉴定[D].保定,河北农业大学,2003
[128]闫红飞,杨文香,褚栋,等.小麦抗叶锈基因Lr38的一个新标记[J].中国农业科学,2008,41(11)：3604～3609
[129]Mebrate SA,Oerke EC, Dehne HW, et al.Mapping of the leaf rust resistance gene Lr38 on wheat chromosome arm 6DL using SSR markers[J].Euphytica,2008,162:457～466
[130]Raupp W J,Sukhwinder-Singh G L,Brown-Guedira B S,et al.Cytogenetic and molecular mapping of the rust resistance gene Lr39 in wheat[J].Theor Appl Genet,2001,102:347～352
[131]Huang L, Korzun V, Gill B S.Molecular markers linked to the rust resistance genes Lr39 and Lr40 of wheat introgressed from Aegilops tauschii[D].Plant & Animal Genome Ⅶ Conference, Town & Country Hotel, San Diego,CA:1999～2001,383
[132]Lottering J M,Botha A M, Klopper R F.AFLP and RAPD markers linked to leaf rust resistance gene Lr4l in wheat[J].Plant and Soil,2002,19:1～6
[133]N. ZHANG,W. YANG,H. YAN,et al.Molecular markers for leaf rust resistance gene Lr45 in wheat based on AFLP analysis[J].Agri Sci in China,2006,5(12):938～943
[134]William M,Singh R P,Huerta Espino J,et al.Molecular marker mapping of leaf rust resistance gene Lr46 and its association with stripe rust resistance gene Yr29 in wheat[J]. Phytopatholgy,2003,93(2):153～159
[135]Helguera M,Khan I A,Dubcovsky J.Development of PCR markers for the wheat leaf rust resistance gene Lr47[J].Theor Appl Genet,2000,100(1):137～143
[136]Bansal U K, Hayden M J,Venkata B P, et al.Genetic mapping of adult plant leaf rust resistance genes Lr48 and Lr49 in common wheat[J].Theor Appl Genet,2008,117:307～312
[137]Brown Guedira G L,Singh S, Fritz A K.Performance and mapping of leaf rust resistance transferred to wheat from Triticum armeniacum[J].Phytopathol,2003,93(7):784～789
[138]Helguem M,Vanzetti L,Soria M,et al.PCR markers for Triticum Speltoides leaf rust resistance gene Lr51 and their use to develop isogenic hard red spring wheat lines[J].Crop Sci,2005, 45:728-734
[139]Colin H,Julian T,Brent M.Locating the broad spectrum wheat leaf rust resistance gene Lr52(LrW)to chromosome 5B by a new cytogenetic method [J].Theor Appl Genet,2005, 110:1453-1457
[140]Hiebert CW,Thomas JB,McCallum B D,et al.Genetic mapping of the wheat leaf rust resistance gene Lr60 (LrW2)[J].Crop Sci,2008,48:1020～1026
[141]Herrera-Foessel SA,Singh RP,Huerta-Espino J,et al.Molecular mapping of a leaf rust resistance gene on the short arm of chromosome 6B of durum wheat[J].Plant Dis,2008,92:1650～1654
[142]刘志勇,王小玲,倪中福,等.小麦抗叶锈基因Lr9,Lr24的分子标记辅助选择研究[J].农业生物技术学报,2000,8(1)：13～16
[143]陈云芳,刘莉,杨文香,等.33个小麦品种(系)抗叶锈基因Lr19分子检测[J].分子植物育种,2008,6(5)：1015～1018
[144]Lukasz S,Lidia GJerzy C,et al.Leaf rust resistance genes of wheat-identification in cultivars and resistance sources[J].J Appl Genet,2003,44(2):139～149
[145]L. G. Tyryshkin,E.I.Gul'tyaeva,N. V.Alpat'eva,et al. Identification of effective leaf rust resistance genes in wheat (Triticum aestivum) using STS markers[J].Russian Journal of Genetics,2006,42(6):662～666
[146]张利军,李在峰,Mortcn Lillcmo,等.CIMMYT小麦品种Saar的叶锈成株抗性QTL分析[J].中国农业科学,2009,42(2)：388～397
[147]Li Huang.High-resolution mapping and molecular cloning of the leaf rust resistance gene Lr21 in wheat[D]. Ph.D Thesis,2002
[148]王海燕TcLr35小麦抗病相关基因的克隆及分析[D].保定,河北农业大学,2006
[149]Simon G K,Evans S L,Wolfgang S,et al.A Putative ABC Transporter Confers Durable Resistance to Multiple Fungal Pathogens in Wheat[J]. Science,2009,323(5919):1360～1363
[150]Roebner R M D,Gennaro A, Ceoloni C.A candidate for Lr19,an exotic gene conditioning leaf rust resistance in wheat[J].Funct Integr Genomics,2009,9(3):325～334
[151]McIntosh RA,Wellings CR,Park RF. Wheat rusts:an atlas of resistance genes (ed. A. Cloud-Guest & K. Jeans)[M]. Dordrecht:Kluwer,1995:1～200.
[152]Ling HQ, Zhu YX, Keller B.High-resolution mapping of the leaf rust disease resistance gene Lrl in wheat and characterization of BAC clones from the Lrl locus[J].Theor Appl Genet,2003,106: 875-882
[153]Ling HQ,Qiu JW,Singh RP,et al.Identification and genetic characterization of an Aegilops tauschii ortholog of the wheat leaf rust disease resistance gene Lr1[J].Theor Appl Genet,2004, 109:1133～1138
[154]Cloutier S, McCallum BD, Loutre C, et al.Leaf rust resistance gene Lrl, isolated from bread wheat(Triticum aestivum L.) is a member of the large psr567 gene family[J].Plant Mol Biol,2007, 65:93～106
[155]刘仙俊.小麦抗叶锈病基因Lr1的进化分析[D].雅安,四川农业大学,2008
[156]山仑.我国西北地区植物水分研究与旱地农业增产[J].植物学通讯,1983(5)：7～11
[157]吴永成,周顺利,王志敏.小麦与抗旱性有关的根系遗传改良研究进展[J].麦类作物学报,2004,24(3)：101～104
[158]Wang W,Vmocur B, Altman A.Plant responses to drought, salinity and extreme temperatures: towards genetics engineering for stress tolerance[J].Planta,2003,21(8):1～14
[159]武仙山,昌小平,景蕊莲.小麦灌浆期抗旱性鉴定指标的综合评价[J].麦类作物学报,2008,28(4)：626～632
[160]王贺正,李艳,马均,等.水稻苗期抗旱性指标的筛选[J].作物学报,2007,33(9)：1523～1529
[161]王春平,张伟,张改生,等.三种分析方法对小麦新品种的综合评价[J].麦类作物学报,2006,26(5)：45～48
[162]刘春芳,罗湘宁,李冬.灰色关联度分析法在亚麻品种评价中的应用[J].中国油料,1997,19(1)：23～26
[163]阎洁治,白春华.灰色关联度分析在大豆引种规律研究中的应用[J].农业系统科学与综合研究,1997,13(2)：88～90
[164]庞欣,李春俭,张福锁.部分根系供磷对小麦幼苗生长及同化物分配的影响[J].作物学报,2000,26(6)：719～724
[165]杨子光,张灿军,冀天会,等.小麦抗旱性鉴定方法及评价指标研究Ⅳ-萌发期抗旱指标的比较研究[J].中国农学通报,2007,23(12)：173～176
[166]曹宏,张希彪,陈靠山.“天达2116”浸种对小麦种子萌发与幼苗抗旱性的影响[J].中国农学通报,2001,17(6)：24～27
[167]张尧庭等.多元统计分析引论[M].北京,科学出版社,1993
[168]张玉梅,林琪,姜雯,等.渗透胁迫条件下不同抗旱性小麦品种萌发期生理生化指标的变化[J].麦类作物学报,2006,26(6)：125～130.
[169]王谨,刘桂茹,杨学举.PEG胁迫下小麦再生植株根系特性与抗旱性的关系[J].麦类作物学报,2006,26(3)：117～119
[170]黄清华,景蕊莲,吴新元,等.人工模拟水分胁迫条件下小麦幼苗抗旱性鉴定指标的分析[J].麦类作物学报,2007,27(4)：661～666
[171]周广生,周竹青,朱旭彤,等.用隶属函数法评价小麦的耐湿性[J].麦类作物学报,2001,21(4)：34～37
[172]李志博,尚勋武.灰色关联度分析法在兰州拉面粉质量评价中的应用[J].麦类作物学报,2007,27(1)：59～62
[173]成雪峰,柴守玺,张凤云.河西绿洲灌区不同灌溉模式下春小麦主要农艺性状与产量和WUE的灰色关联分析[J].麦类作物学报,2007,27(4)：699～704
[174]张灿军,姚宇卿,王育红,等.早稻抗旱性鉴定方法与指标研究-Ⅰ鉴定方法与评价指标[J].干旱地区农业研究,2005,23(3)：33～36
[175]郜俊红,廖先静,李晓青,等.水分胁迫对不同小麦品种种子萌发的影响[J].安徽农业科学,2008,36(8)：3106～3107
[176]Higuchi R,Dollinger Q Walsh P S,et al.Simultaneous amplification and detection of specific DNA sequences[J].Nature Biotechnology,1992,10:413～417
[177]庄巧生.中国小麦品种改良及系谱分析[M].北京,中国农业出版社,2003
[178]金善宝.中国小麦品种及其系谱[M].北京,中国农业出版社,1983
[179]冯晶,淦爱华,徐世昌,等.中国小麦条锈菌鉴别寄主南大2419抗条锈性遗传分析[J].中国农业大学学报,2009,14(3)：42～48
[180]张知仪,张阳,李俊,等.春化基因VRN-1在骨干亲本南大2419衍生系中的分布[J].西南农业学报,2009,22(3)：568～571
[181]司清林,刘新伦,刘智奎,等.阿夫及其衍生小麦品种(系)的SSR分析[J].作物学报,2009,35(4)：615～619
[182]李红琴,相吉山,郭青云,等.小麦骨干亲本阿夫及其衍生品种(系)的高分子量麦谷蛋白亚基演变分析[J].植物遗传资源学报,2009,10(1)：37～41
[183]张利军.中国小麦品种(系)抗叶锈性鉴定以及CIMMYT小麦Saar的成株抗叶锈QTL分析[D].保定,河北农业大学.2008
[2]赵济,陈传康.中国地理[M].北京：高等教育出版社,1999：9～10.
[3]倪胜利,张国宏,李兴茂.旱地小麦育种的研究机理与进展[J].现代农业科技,2007,24：102～105
[4]吴永成,周顺利,王志敏.小麦与抗旱性有关的根系遗传改良研究进展[J].麦类作物学报,2004,24(3)：101～104
[5]郝艳军,山仑.高粱抗旱机理的研究进展[J].中国农学通报,2004,20(3)：120～123
[6]欧巧明,倪建福,马瑞君.春小麦根系木质部导管与其抗旱性的关系[J].麦类作物学报,2005,25(3)：27～31
[7]肖玲,赵先贵.渭北旱塬小麦根系的再生能力及其意义[J].水土保持通报,1996,16(2)：55～57
[8]杨静慧,杨焕庭.苹果树植物叶片角质层厚度与植物抗旱性[J].天津农学院学报,1996,3(3)：27～28
[9]蒋高明.植物生理生态[M].北京,高等教育出版社,2004：191～192
[10]韦梅琴,李军乔.委陵菜属四种植物茎叶解剖结构的比较研究[J].青海师范学院学报,2003,(3)：48～50.
[11]Lupton FGH.小麦育种的理论基础[M].北京,北京农业大学出版社,1988,346～348
[12]张跃强,樊哲儒,王延军,等.小麦抗旱节水研究进展[J].新疆农业科学,2007,44(S3)：36～43
[13]卢振民.冬小麦近轴和远轴叶面气孔对土壤水分胁迫反应的敏感性[J],植物生理学报,1988,14(3)：223～227
[14]高彦萍,冯莹,马志军,等.水分胁迫下不同抗旱类型大豆叶片气孔特性变化研究[J].干旱地区农业研究,2007,25(2)：77～79
[15]王玮.植物对水分亏缺的某些生化反应[J].植物生理学通讯,1998,34(5)：388～393
[16]郑敏娜,李向林,万里强,等.水分胁迫对6种禾草叶绿体、线粒体超微结构及光合作用的影响[J].草地学报,2009,17(5)：643～649
[17]Singh B,Usha K.Salicylic acid induced physiological and biochemical changes in wheat seedlings under water stress[J].Plant Growth Regulation,2003,39(2):137～141
[18]Cohen I,Knopf J A,Irihimovitch V,et al.Aproposed mechanism for the inhibitory effects of oxidative stress on rubisco assembly and it s subunit expression[J].Plant Physiol,2005,137:376～387
[19]Shinozaki K,Yamaguchi-Shinazaki K.Gene expression and signal transduction in water-stress response[J].Plant Physiol,1997,115:327～334
[20]Mattioni C.Lacerenza N G,Troccoli A,et al.Water and salt stress induced alterations on praline metabolism of triticum durum seedlings[J].Physiol Plantarum,1997,101(4):787～792
[21]Nayyar H,Walia D P. Water stress induced proline accumulation in contrasting wheat genotypes as affected by calcium and abscisic acid[J].Biologia Plantarum,2003,46:275～279
[22]邵宏波,梁宗锁,邵明安.小麦抗旱生理生化和分子生物学研究进展与趋势[J].草业学报,2006,15(3)：5～17
[23]Takashi Hibino,Yu-Ling Meng,Yoshinobu Kawamitsu,et al.Molecular cloning and functional characterization of two kinds of betaine-aldehyde dehydrogenase in betaine-accumulating mangrove Avicennia marina (Forsk.) Vierh [J].Plant Mol Bio,2001,45:353～363
[24]Munns R,Bracely c J,Barlow E W R.Solute accumulation in the apex and leaves of wheat during water stress[J].Aus J Plant Physiol,1979 (6):379～389
[25]汤章城.逆境条件下植物脯氨酸的累积及其可能的意义[J].植物生理学通讯,1984(1)：15～17
[26]Bray E A.Stress induced gene expression in plants[M].Switzerland,Harwood Academic publishers GmbH Switzerland,1994,1～23
[27]刘春玲,陈慧萍,刘娥娥,等.水稻品种对几种逆境的多重耐性及与ABA的关系[J].作物学报,2003,29(5)：725～729
[28]Yang H Q,Jia W S.Water stress induced ABA accumulated by protein phosphorylation participating in roots of Malus rupenensis Rehd[J].Chinese Sci Bulletin,2001,46 (1):50～53
[29]余叔文,汤章城.植物生理与分子生物学[M].北京,科学出版社,1999：476～492
[30]Chen N,Goodwin PH,Hsiang T.The role of ethylene during the infection of Nicotiana tabacum by Colletotrichum destructivum[J]. J Exp Bot,2003,54(392):2449～2456
[31]郭丽红,王定康,杨晓虹,等.外源乙烯利对干旱胁迫过程中玉米幼苗某些抗逆生理指标的影响[J].云南大学学报(自然科学版),2004,6(4)：352～356
[32]Ronald Pierik,Rashmi Sasidharan,Laurentius A C J Voesenek.Growth control by ethylene: Adjusting phenotypes to the environment[J]. Plant Growth Regul,2007,26:188～200.
[33]段喜华,孙莲慧,郭晓瑞,等.拟南芥乙烯突变体在干旱胁迫下的形态学差异[J].植物研究,2009,29(1)：39～42
[34]Fan T L,Balta M,Rudd J,Payne W A.Canopy temperature depression as a potential selection criterion for drought resistance in wheat[J].Agri Sci in China,2005,4:793～800
[35]Zhang J X,Kirham M B.Drought stress induced changes in activities of superoxide dismutase,catalase,and perexidase in wheat species [J].Plant Cell Physiol,1994,35 (5):785～791
[36]Hieng B,Ugrinovic K,Sustar-Vozlic J,et al.Different classes of proteases are involved in the response to drought of Phaseolus vulgaris L.cultivars differing in sensitivity [J].Plant Physiol,2004,161:519～530
[37]Badger M R.Photosynthetic oxygen exchanges[J].Plant Physiol,1985,36:27～53
[38]张喜英,裴东,曲懋正.几种作物的生理指标对土壤水分变动的阈值反应[J].植物生态学报,2000,24(3)：280～283
[39]景蕊莲,作物抗旱研究的现状与思考[J].干旱地区农业研究,1999,17(2)：79～85
[40]周桂莲,杨慧霞.小麦抗旱性鉴定的生理生化指标及其分析评价[J].干旱地区农业研究,1996,14(2)：65～69
[41]张林刚,邓西平.小麦抗旱性生理生化研究进展[J].干旱地区农业研究,2000,18(3)：125～132
[42]张娟,张正斌,谢惠民,等.小麦叶片水分利用效率及相关生理性状的关系[J].作物学报,2005,31(12)：1593～1599
[43]蔡晓莉,吴振录,樊哲儒,等.新疆春小麦主栽品种和新品种的抗旱性研究初报[J].新疆农业科学,2005,42(2)：70～76
[44]林关石.谷子品种抗旱性观察[J].陕西农业科学,1984,4：17～19
[45]刘学义.大豆抗旱性评定方法探讨[J].中国油料,1986,4：23～26
[46]兰巨生.作物抗旱指数的概念和统计方法[J].华北农学报,1990,5(2)：20～25
[47]兰巨生.农作物综合抗旱性评价方法的研究[J].西北农业学报,1998,7(3)：85～87
[48]栗雨勤.高产抗旱冬小麦新品种鉴定筛选方法研究[J].华北农学报,1998,13：45～49
[49]路贵和,安海润.作物抗旱性鉴定方法与指标研究进展[J].山西农业科学,1999,27(4)：39～43
[50]冀天会,张灿军,杨子光,等.小麦抗旱性鉴定方法及评价指标研究Ⅲ-综合评价指标的比较研究[J].中国农学通报,2007,23(9)：226～230
[51]张灿军,冀天会,杨子光,等.小麦抗旱性鉴定方法及评价指标研究Ⅱ-抗旱性鉴定评价技术规程[J].中国农学通报,2007,23(10)：418～421
[52]胡荣海.农作物资源的抗旱筛选技术及其应用[J].农牧情报研究,1989(1)：36～41
[53]胡荣海,昌小屏.反复干旱法的生理基础及其应用[J].华北农学报,1996,11(3)：51～56
[54]黎裕.植物抗旱育种及其进展[J].世界农业,1994,3：18～20
[55]周该生.水稻抗旱性苗期鉴定指标筛选与节水机理研究[D].武汉,华中农业大学,2006
[56]刘思峰.灰色系统理论及其应用(第三版)[M].北京,2004,科学出版社
[57]扬子光,王书子,吴少辉,等.灰色关联分析在小麦品种抗旱性鉴定上的应用[J].中国农学通报,2002,18(2)：42～43
[58]冀天会,张灿军,谢惠民.小麦品种抗旱性研究的产量鉴定指标的比较研究[J].中国农学通报,2006,22(1)：103～106
[59]王士强,胡银岗,佘奎军,等.小麦抗旱相关农艺性状和生理生化性状的灰色关联度分析[J].中国农业科学,2007,40(11)：2452～2459
[60]李玉发,何中国,李淑芳,等.东北地区春小麦主要性状与产量间的灰色关联分析[J].麦类作物学报,2005,25(1)：139～141
[61]裘敏,魏亦勤,刘旺清,等.小麦5个产量性状与产量的灰色关联度分析[J].安徽农业科学,2007,35(2)：390
[62]韩启秀.运用灰色关联度对山东小麦新品种(系)综合表现的评价[J].中国农学通报,2005,21(2)：312～314
[63]高素玲,曹雯梅.应用灰色系统理论综合评价春性小麦新品种[J].河南农业科学,2006(4)：31～33
[64]Komer J A.Genetics of resistance to wheat leaf rust[J].Annual Review Phytopathology,1996, 34:435～455
[65]Kloppers F J,Pretorius Z A.Effects of combinations among genes Lrl3,Lr34 and Lr37 on components of resistance in wheat to leafrust[J].Plant Pathology,1997,46:737～750
[66]Knott D R.The wheat rust breeding for resistance[J].Theor Appl Genet,1989:191～198
[67]Friebc B,Zeller F J, Mukai Y, et al.Characterization of rust resistant wheat Agropyrom intermedium derivatives by C-banding in situ hybridization and isozyme analysis[J].Theor Appl Genet,1992,83:775～782
[68]R A.McIntosh,J Dubcovsky,W J Rogers,et al.Catalogue of gene symbols for wheat:2009 supplement[R].Annual Wheat Newsl,2009,55
[69]Kilpatrick RA.New wheat cultivars and longevity of rust resistance,1971～1975[M].Beltsvill e,MD:US Department of Agriculture,Agricultural Research Service,1975
[70]Etienne Duveiller, Ravi P. Singh, Julie M. Nicol. The challenges of maintaining wheat productivity:pests, diseases, and potential epidemics[J]. Euphytica,2007,157(3):417～430
[71]Eriksen L,Afshari F,Christiansen M J,el al.Yr32 for resistance to stripe rust present in the wheat cultivar Carstens V[J].Theor Appl Genet,2004,108:567～575
[72]McIntosh R A,Lagudah E S.Cytogenetical studies in wheat Ⅷ.Gene Yr24 for resistance to stripe rust[J].Plant Breed,2000,119:81～83
[73]Wan A M,Zhao Z H,Chen X M,et al,Wheat stripe rust epidemic and virulence of Puccinia striiformis f.sp.tritici in China in 2002[J].Plant Dis,2004,88:896～904
[74]Oelke L M, Kolmer J A.Genetics of leaf rust resistance in spring wheat cultivars Alsen and Norm [J].Phytopathology,2005,95:773～778
[75]曹张军,井金学,王美南,等.小麦品种贵农22号抗条锈基因遗传分析[J].西北植物学报,2004,24(6)：991～996
[76]代君丽,牛永春.小麦农家品种大籽糙抗条锈性的遗传分析[J].遗传,2003,25(3)：311～313
[77]徐世昌,张敬原,赵文生,等.小麦京核891-1抗条锈病主效、微效基因的遗传分析[J].中国农业科学,2001,34(3)：272～276
[78]谢皓,牛永春,陈孝,等.小麦新品系YW243抗条锈性鉴定和遗传分析[J].植物病理学报,2003,33(3)：243～247
[79]陈万权,胡长程,晏思自,等.冬小麦品种北京837抗叶锈病基因的染色体定位研究[J].植物病理学报,1995,25(4)：297～300
[80]薛秀庄,吉万全,王秋英,等.小麦性状的缺体分析[J].西北农业学报,1995,4(4)：1～5
[81]Flor H H. The complementary genetics systems in flax and flax rust[J].Advances in Genetics, 1956,8:29～54
[82]Loegering W Q,McIntosh R A,Burton C H.Computer analysis of disease data to derive hy pothetical genotypes for reaction of host varieties to pathogens[J].Can J Genet Cytni,1971,13:742～747
[83]Browder L E.Designation of two genes for resistance to Puccinia recondite in triticum aestivum[J].Crop Sci,1972,12:705～706
[84]Long D L,Schafer J F,Roelfs A P,et al. Specifici virulence of Puccinia Recondite f sp.Trifici in the United States for 1979～1983[J].Plant Dis,1985,9:343～347
[85]Dubin H J, Johnson R,Stubbs R W.Postulated genes for resistance to strip rust in selected CIMMYT and related wheats [J].Plant Dis,1989,73(6):472～475
[86]Johnson R,Stubbs RW,Kirmani MAS,et al.Discussion of a method resulting in erroneous postulation of the gene Yr8 for resistance to Puccinia striiformis in Pakistan wheat cultivars[J]. Cereal Rusts and Powdery Mildews Bulletin,1987,15:13～19
[87]Zoldan S M,Barcellos A L.Postulation of genes (Lr) for resistance to leaf rust in Brazilian wheat cultivars[J].Fitopatol Bras,2002,27(5):508～516
[88]陈万权,王剑雄.76个小麦种质资源抗叶锈及秆锈基因初步分析[J].作物学报,1997,23(6)：655～663
[89]王奎荣,陈万权,冯国华,等.黄淮麦区小麦品种抗叶锈病基因分析[J].江苏农业学报,1997,13(3)：167～170
[90]Zhao Huan-huan, Singh R P,Huerta-Espino Julio,et al. Resistance to leaf rust in forty-three Chinese wheat cultivars and line[J].ActaPhytopathologica Sinica,2006,36(6):539～549
[91]伍少云,奉有壁,汤翠凤.云南小麦资源创新种质的抗锈性与生态适应性鉴定研究[J].麦类作物学报.2002,22(3)：25～30
[92]Singh R P,Gupta A K.Genes for leaf rust resistance in Indian and Pakistan wheat resistance tested with Mexican pathotypes of Puccinia recondita f.sp. tritici[J].Euphytica,1991,5 7:27～36
[93]Winzeler M,Winzeler H,Keller B,et al.Endopestidase polymorphism and linkage of the EP-Dlc null allele with the Lr19 leaf rust resistance gene in hexaploid wheat[J].Plant Breeding,1995,114:24～28
[94]Feuillet C,Messmer M,Schachenmayr G,et al.Genetic and physical characterization of the Lrl leaf rust resistance locus in wheat(Triticum aestivum L.) [J].Mol Gen Genet,1995,248:553～562
[95]Qiu JW,Schurch AC,Yahiaoui N,et al.Physical mapping and identification of a candidate for the leaf rust resistance gene Lrl of wheat[J].Theor Appl Genet,2007,115:159～168
[96]张娜,闫红飞,张英春,等.小麦抗叶锈病基因Lr2c的SSR标记[J].农业生物技术学报,2009,17(1)：148～152
[97]Sacco F, Suilrez E Y, Naranjo T. Mapping of the leaf rust resistance gene Lr3 on chromosome 6B of Sinvalocho MA wheat[J].Genome,1998,41(5):686～690
[98]Gupta S K, Charpe A, Koul S, et al.Development and validation of molecular markers linked to an Aegilops umbellulata-derived leaf rust resistance gene, Lr9, for marker assisted selection in bread wheat[J].Genome,2005,48(5):823～830
[99]Schachermayr G,Feuillet C,Keller B.Molecular markers for the detection of the wheat leaf rust resistance gene Lr10 in diverse genetic backgrounds[J].Mol Breeding,1997,9(3):65～74
[100]Feuillet C,Schachermayr G,Keller B,et al.Molecular cloning of a new receptor-like kinase gene encoded at the Lr10 disease resistance locus of wheat[J].The Plant Journal,1995,11(1):45～52
[101]Nils S,Catherine F,Thomas W,et al.Subgenome chromosome walking in wheat:A 450-kb physical contig in Triticum monococcum L.spans the Lr10 resistance locus in hexaploid wheat(Triticum astivum L.)[J].PNAS,2000,97:13436-13441
[102]Seyfarth R,Feuillet C,Schachermayr G,et al.Molecular mapping of the adult-plant leaf rust resistance gene Lr13 in wheat(Triticum astivum L.) [J].Genet Breed,2000,54:193～198
[103]Prins R J,Groene wald Z G,Marais F,et al.AFLP and STS tagging of Lr19, a gene conferring resistance to leaf rust in wheat[J].Theor Appl Genet,2001,103:618-624
[104]李星,杨文香,李亚宁,等.小麦抗叶锈病基因Lr19的SSR标记[J].中国农业科学,2005,38(6)：1156-1159
[105]刘雅辉,闫红飞,杨文香,等.小麦抗叶锈病基因Lr19的SRAP标记[J].华北农学报,2007,2007,22(4)：193～196
[106]闫红飞,杨文香,陈云芳,等.偃麦草属E染色体组特异SCAR标记对Lr19的特异性和稳定性研究[J].植物病理学报,2009,39(1)：76～81
[107]Neu C H,Stein N,Keller B.Genetic mapping of the Lr20-pml resistance locus reveals suppressed recombination on chromosome arm 7AL in hexaploid wheat[J].Genome,2002,45:737～744
[108]Huang L,Gill B S.An RGA-like marker detects all known Lr21 leaf rust resistance gene family members in Aegilops tauschii and wheat[J].Theor Appl Genet,2001,103:1007～1013
[109]Schachermayr G M,Messmer M M, Feuillet C,et al.Identification of molecular markers linked to the Agropyron elongatum-derived leaf resistance gene Lr24 in wheat[J].Theor Appl Genet,1995,90:982～990
[110]Dedryver F,Jubier M F,Thouvenin J,et al.Molecular markers linked to the rust resistance genes Lr24 in different wheat cultivars[J].Genome,1996,39(5):830～835
[111]张娜,陈玉婷,李亚宁,等.小麦抗叶锈病基因Lr24的一个新STS标记[J].作物学报,2008,34(2)：212～216
[112]Procunier J D,Townley-Smith T,Fox S,et al.PCR-based RAPD/DGGE markers linked to leaf rust resistance genes Lr29 and Lr25 in wheat(Triticum aestivum L.)[J].Genet Breeding,1995,49:87～ 92
[113]Chai J F,Zhou R H,Jia J Z,et al.Development and application of a new codominant PCR marker for detecting 1BL/1RS wheat-rye chromosome translocations[J].Plant Breed,2006,125:302～304
[114]Nelson J C,Singh R P,Autrique E,et al.Mapping genes conferring and suppressing leaf rust resistance in wheat[J].Crop Science,1997,37:1928～1935
[115]Naik S,Gill K S,Prakasa Rao V S,et al.Identification of a STS marker linked to the Aegilops speltoides-derived leaf rust resistance gene Lr28 in wheat[J].Theor Appl Genet,1998,97:535～540
[116]Yogesh V,Parveen C,Rippy S,et al.Tagging of Aegilops speltoides-derived leaf rust resistance gene Lr28 with a microsatellite marker in wheat[J].Plant Biochemistry Biotechnology, 2004,13:47～49
[117]Tar M, Punnhauser L, Csosz L, et al. Identification of molecular markers for an efficient leaf rust resistance gene Lr29 in wheat[J]. Acta Biologica Szegediensis,2002,46:133～134
[118]Autrique E,Singh R P, Tanksley S D,et al.Molecular markers for four leaf rust resistance genes introgressed into wheat from wild relatives[J].Genome,1995,38:75～83
[119]E S Lagudah, H. McFadden, R P Singh,et al. Molecular genetic characterization of the Lr34/Yr18 slow rusting resistance gene region in wheat[J].Theor Appl Genet,2006,114(1):21～ 30
[120]Evans S. Lagudah, Simon G. Krattinger, Sybil Herrera-Foessel, et al.Gene-specific markers for the wheat gene Lr34|Yr18|Pm38 which confers resistance to multiple fungal pathogens[J]. Theor Appl Genet,2009,119(5):889～898
[121]Seyfarth R, Feuillet C, Schachermayr Q,et al.Development of a molecular for the adult plant leaf rust resistance gene Lr35 in wheat[J]. Theor Appl Genet,1999,99:554～560
[122]Gold J,Harder D,Townley Smith F,et al.Development of a molecular marker for rust resistance genes Sr39 and Lr35 in wheat breeding lines[J].Electronic Joural of Biotechnology, 1999,2(1):45～52
[123]李丽娜.小麦抗叶锈基因Lr35的AFLP和SSR分子标记[D].保定,河北农业大学,2005
[124]Robert O, Abelard C, Dedryver F,et al.Identification of molecular markers for the wheat of the yellow rust resistance gene Yr17 in wheat[J].Mol Breed,1999,5:167～175
[125]Seah S,Bariana H,Jahier J,et al.The introgressed segment carry rust resistance genes Yrl7, Lr37 and Sr38 in wheat can be assayed by a cloned disease resistance gene-like sequence[J]. Theor Appl Genet,2001,102:600～605
[126]张立荣,徐大庆,杨文香,等.小麦抗叶锈基因Lr37 ISSR分子标记[J].农业生物技术学 报,2004,1：86～89
[127]杨文香Lr37,Lr44 AFLP分子标记及124个小麦品种(系)抗叶锈基因鉴定[D].保定,河北农业大学,2003
[128]闫红飞,杨文香,褚栋,等.小麦抗叶锈基因Lr38的一个新标记[J].中国农业科学,2008,41(11)：3604～3609
[129]Mebrate SA,Oerke EC, Dehne HW, et al.Mapping of the leaf rust resistance gene Lr38 on wheat chromosome arm 6DL using SSR markers[J].Euphytica,2008,162:457～466
[130]Raupp W J,Sukhwinder-Singh G L,Brown-Guedira B S,et al.Cytogenetic and molecular mapping of the rust resistance gene Lr39 in wheat[J].Theor Appl Genet,2001,102:347～352
[131]Huang L, Korzun V, Gill B S.Molecular markers linked to the rust resistance genes Lr39 and Lr40 of wheat introgressed from Aegilops tauschii[D].Plant & Animal Genome Ⅶ Conference, Town & Country Hotel, San Diego,CA:1999～2001,383
[132]Lottering J M,Botha A M, Klopper R F.AFLP and RAPD markers linked to leaf rust resistance gene Lr4l in wheat[J].Plant and Soil,2002,19:1～6
[133]N. ZHANG,W. YANG,H. YAN,et al.Molecular markers for leaf rust resistance gene Lr45 in wheat based on AFLP analysis[J].Agri Sci in China,2006,5(12):938～943
[134]William M,Singh R P,Huerta Espino J,et al.Molecular marker mapping of leaf rust resistance gene Lr46 and its association with stripe rust resistance gene Yr29 in wheat[J]. Phytopatholgy,2003,93(2):153～159
[135]Helguera M,Khan I A,Dubcovsky J.Development of PCR markers for the wheat leaf rust resistance gene Lr47[J].Theor Appl Genet,2000,100(1):137～143
[136]Bansal U K, Hayden M J,Venkata B P, et al.Genetic mapping of adult plant leaf rust resistance genes Lr48 and Lr49 in common wheat[J].Theor Appl Genet,2008,117:307～312
[137]Brown Guedira G L,Singh S, Fritz A K.Performance and mapping of leaf rust resistance transferred to wheat from Triticum armeniacum[J].Phytopathol,2003,93(7):784～789
[138]Helguem M,Vanzetti L,Soria M,et al.PCR markers for Triticum Speltoides leaf rust resistance gene Lr51 and their use to develop isogenic hard red spring wheat lines[J].Crop Sci,2005, 45:728-734
[139]Colin H,Julian T,Brent M.Locating the broad spectrum wheat leaf rust resistance gene Lr52(LrW)to chromosome 5B by a new cytogenetic method [J].Theor Appl Genet,2005, 110:1453-1457
[140]Hiebert CW,Thomas JB,McCallum B D,et al.Genetic mapping of the wheat leaf rust resistance gene Lr60 (LrW2)[J].Crop Sci,2008,48:1020～1026
[141]Herrera-Foessel SA,Singh RP,Huerta-Espino J,et al.Molecular mapping of a leaf rust resistance gene on the short arm of chromosome 6B of durum wheat[J].Plant Dis,2008,92:1650～1654
[142]刘志勇,王小玲,倪中福,等.小麦抗叶锈基因Lr9,Lr24的分子标记辅助选择研究[J].农业生物技术学报,2000,8(1)：13～16
[143]陈云芳,刘莉,杨文香,等.33个小麦品种(系)抗叶锈基因Lr19分子检测[J].分子植物育种,2008,6(5)：1015～1018
[144]Lukasz S,Lidia GJerzy C,et al.Leaf rust resistance genes of wheat-identification in cultivars and resistance sources[J].J Appl Genet,2003,44(2):139～149
[145]L. G. Tyryshkin,E.I.Gul'tyaeva,N. V.Alpat'eva,et al. Identification of effective leaf rust resistance genes in wheat (Triticum aestivum) using STS markers[J].Russian Journal of Genetics,2006,42(6):662～666
[146]张利军,李在峰,Mortcn Lillcmo,等.CIMMYT小麦品种Saar的叶锈成株抗性QTL分析[J].中国农业科学,2009,42(2)：388～397
[147]Li Huang.High-resolution mapping and molecular cloning of the leaf rust resistance gene Lr21 in wheat[D]. Ph.D Thesis,2002
[148]王海燕TcLr35小麦抗病相关基因的克隆及分析[D].保定,河北农业大学,2006
[149]Simon G K,Evans S L,Wolfgang S,et al.A Putative ABC Transporter Confers Durable Resistance to Multiple Fungal Pathogens in Wheat[J]. Science,2009,323(5919):1360～1363
[150]Roebner R M D,Gennaro A, Ceoloni C.A candidate for Lr19,an exotic gene conditioning leaf rust resistance in wheat[J].Funct Integr Genomics,2009,9(3):325～334
[151]McIntosh RA,Wellings CR,Park RF. Wheat rusts:an atlas of resistance genes (ed. A. Cloud-Guest & K. Jeans)[M]. Dordrecht:Kluwer,1995:1～200.
[152]Ling HQ, Zhu YX, Keller B.High-resolution mapping of the leaf rust disease resistance gene Lrl in wheat and characterization of BAC clones from the Lrl locus[J].Theor Appl Genet,2003,106: 875-882
[153]Ling HQ,Qiu JW,Singh RP,et al.Identification and genetic characterization of an Aegilops tauschii ortholog of the wheat leaf rust disease resistance gene Lr1[J].Theor Appl Genet,2004, 109:1133～1138
[154]Cloutier S, McCallum BD, Loutre C, et al.Leaf rust resistance gene Lrl, isolated from bread wheat(Triticum aestivum L.) is a member of the large psr567 gene family[J].Plant Mol Biol,2007, 65:93～106
[155]刘仙俊.小麦抗叶锈病基因Lr1的进化分析[D].雅安,四川农业大学,2008
[156]山仑.我国西北地区植物水分研究与旱地农业增产[J].植物学通讯,1983(5)：7～11
[157]吴永成,周顺利,王志敏.小麦与抗旱性有关的根系遗传改良研究进展[J].麦类作物学报,2004,24(3)：101～104
[158]Wang W,Vmocur B, Altman A.Plant responses to drought, salinity and extreme temperatures: towards genetics engineering for stress tolerance[J].Planta,2003,21(8):1～14
[159]武仙山,昌小平,景蕊莲.小麦灌浆期抗旱性鉴定指标的综合评价[J].麦类作物学报,2008,28(4)：626～632
[160]王贺正,李艳,马均,等.水稻苗期抗旱性指标的筛选[J].作物学报,2007,33(9)：1523～1529
[161]王春平,张伟,张改生,等.三种分析方法对小麦新品种的综合评价[J].麦类作物学报,2006,26(5)：45～48
[162]刘春芳,罗湘宁,李冬.灰色关联度分析法在亚麻品种评价中的应用[J].中国油料,1997,19(1)：23～26
[163]阎洁治,白春华.灰色关联度分析在大豆引种规律研究中的应用[J].农业系统科学与综合研究,1997,13(2)：88～90
[164]庞欣,李春俭,张福锁.部分根系供磷对小麦幼苗生长及同化物分配的影响[J].作物学报,2000,26(6)：719～724
[165]杨子光,张灿军,冀天会,等.小麦抗旱性鉴定方法及评价指标研究Ⅳ-萌发期抗旱指标的比较研究[J].中国农学通报,2007,23(12)：173～176
[166]曹宏,张希彪,陈靠山.“天达2116”浸种对小麦种子萌发与幼苗抗旱性的影响[J].中国农学通报,2001,17(6)：24～27
[167]张尧庭等.多元统计分析引论[M].北京,科学出版社,1993
[168]张玉梅,林琪,姜雯,等.渗透胁迫条件下不同抗旱性小麦品种萌发期生理生化指标的变化[J].麦类作物学报,2006,26(6)：125～130.
[169]王谨,刘桂茹,杨学举.PEG胁迫下小麦再生植株根系特性与抗旱性的关系[J].麦类作物学报,2006,26(3)：117～119
[170]黄清华,景蕊莲,吴新元,等.人工模拟水分胁迫条件下小麦幼苗抗旱性鉴定指标的分析[J].麦类作物学报,2007,27(4)：661～666
[171]周广生,周竹青,朱旭彤,等.用隶属函数法评价小麦的耐湿性[J].麦类作物学报,2001,21(4)：34～37
[172]李志博,尚勋武.灰色关联度分析法在兰州拉面粉质量评价中的应用[J].麦类作物学报,2007,27(1)：59～62
[173]成雪峰,柴守玺,张凤云.河西绿洲灌区不同灌溉模式下春小麦主要农艺性状与产量和WUE的灰色关联分析[J].麦类作物学报,2007,27(4)：699～704
[174]张灿军,姚宇卿,王育红,等.早稻抗旱性鉴定方法与指标研究-Ⅰ鉴定方法与评价指标[J].干旱地区农业研究,2005,23(3)：33～36
[175]郜俊红,廖先静,李晓青,等.水分胁迫对不同小麦品种种子萌发的影响[J].安徽农业科学,2008,36(8)：3106～3107
[176]Higuchi R,Dollinger Q Walsh P S,et al.Simultaneous amplification and detection of specific DNA sequences[J].Nature Biotechnology,1992,10:413～417
[177]庄巧生.中国小麦品种改良及系谱分析[M].北京,中国农业出版社,2003
[178]金善宝.中国小麦品种及其系谱[M].北京,中国农业出版社,1983
[179]冯晶,淦爱华,徐世昌,等.中国小麦条锈菌鉴别寄主南大2419抗条锈性遗传分析[J].中国农业大学学报,2009,14(3)：42～48
[180]张知仪,张阳,李俊,等.春化基因VRN-1在骨干亲本南大2419衍生系中的分布[J].西南农业学报,2009,22(3)：568～571
[181]司清林,刘新伦,刘智奎,等.阿夫及其衍生小麦品种(系)的SSR分析[J].作物学报,2009,35(4)：615～619
[182]李红琴,相吉山,郭青云,等.小麦骨干亲本阿夫及其衍生品种(系)的高分子量麦谷蛋白亚基演变分析[J].植物遗传资源学报,2009,10(1)：37～41
[183]张利军.中国小麦品种(系)抗叶锈性鉴定以及CIMMYT小麦Saar的成株抗叶锈QTL分析[D].保定,河北农业大学.2008