小麦株高基因的分子标记
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摘要
小麦抗旱性遗传改良是小麦生产上的一个重要研究课题。小麦抗旱性涉
及到形态、生理、生化及遗传等多方面的因素，是一个复杂的数量性状。而
株高是与抗旱性及籽粒产量密切相关的重要性状。
本研究选用生产上推广的高秆（100cm以上）抗旱品种晋麦5号（不含
矮秆基因）与半矮秆（70cm左右）水敏感品种鲁麦14（含矮秆基因Rht）为
亲本进行杂交，对其杂交F_1代进行花药离体培养，得到的再生植株经秋水仙
素处理获得的加倍单倍体（DH）群体。以该群体为作图群体，借助于苗期赤霉
酸敏感性鉴定、成株期株高分析，用SSR（简单序列重复）技术对矮秆基因
进行分子标记作图。
结果表明，鲁麦14携带一个矮秆基因，在苗期和成株期均表现为显性
遗传。共用98对引物进行扩增和筛选，其中43对在亲本间扩增出多态性片
断，9对在DH群体的高矮DNA池间显示出多态性。分子作图表明该矮秆基
因Rht位于小麦染色体4BS上。获得3个与该基因连锁的SSR标记，其中
WMS368离Rht基因位置最近，图距为15.3cM，另两个SSR标记为WMS495
（图距为18.1cM）和WMS513（图距为26.9cM），它们在分子连锁图谱上的
顺序为WMS368-Rht-WMS513-WMS495。该基因的定位及其分子标记的获得
丰富了该连锁群的遗传图谱，为进一步深入研究该基因对小麦抗旱性的籽粒
产量影响和小麦辅助标记育种提供了有用信息。
Wheat genetic improvement on drought resistance is an imporant project in
wheat producion. Wheat drought resistance is a complicated quantitative trait
which involves morphological, physiological biological and hereditory elements,
etc. It is found that wheat height is an important trait closely relevant to drought
resistance and seeds production. In this study, molecular mapping of the GA
insensitive gene Rht3 is carried using the DH population of Jinmai 5 and Lumai
14 (caning Rht gene). The results are as follows:
Lumai 14 carried a gene Rht3. It behaved as a dominant trait at seedlings and
adult stages. Molecular mapping indicated that this gene was located on the
chromosome 4B. Using 98 primers to amplify and screen the population, of them
43 primers have been found to be polymorphismic between the parents. However,
9 primers can amplified the polymorphism between the DNA pool of plant height.
Our results showed that Bulked Segregant Analysis and rnicrosatellite primer
screening of DH population identified three chromosome 4B microsatellite
markers, WMS368, WMS495 and WMS5 13, only WMS368 closely linked to
Rh13. The linkage distance between WMS368 and Rht3 was 15.3cM, WMS5 13,
WMS495 and Rht3 was 18.1cM, 26.9cM respectively.
The location of Rht3 gene and the acquisition of the molecular markers
enriched the genetic map of this linked group. They also provided valuable
information for our further research on the effects of this gene on wheat drought
resistance and seeds production, and for marker-assistant wheat breeding.
及到形态、生理、生化及遗传等多方面的因素，是一个复杂的数量性状。而
株高是与抗旱性及籽粒产量密切相关的重要性状。
本研究选用生产上推广的高秆（100cm以上）抗旱品种晋麦5号（不含
矮秆基因）与半矮秆（70cm左右）水敏感品种鲁麦14（含矮秆基因Rht）为
亲本进行杂交，对其杂交F_1代进行花药离体培养，得到的再生植株经秋水仙
素处理获得的加倍单倍体（DH）群体。以该群体为作图群体，借助于苗期赤霉
酸敏感性鉴定、成株期株高分析，用SSR（简单序列重复）技术对矮秆基因
进行分子标记作图。
结果表明，鲁麦14携带一个矮秆基因，在苗期和成株期均表现为显性
遗传。共用98对引物进行扩增和筛选，其中43对在亲本间扩增出多态性片
断，9对在DH群体的高矮DNA池间显示出多态性。分子作图表明该矮秆基
因Rht位于小麦染色体4BS上。获得3个与该基因连锁的SSR标记，其中
WMS368离Rht基因位置最近，图距为15.3cM，另两个SSR标记为WMS495
（图距为18.1cM）和WMS513（图距为26.9cM），它们在分子连锁图谱上的
顺序为WMS368-Rht-WMS513-WMS495。该基因的定位及其分子标记的获得
丰富了该连锁群的遗传图谱，为进一步深入研究该基因对小麦抗旱性的籽粒
产量影响和小麦辅助标记育种提供了有用信息。
Wheat genetic improvement on drought resistance is an imporant project in
wheat producion. Wheat drought resistance is a complicated quantitative trait
which involves morphological, physiological biological and hereditory elements,
etc. It is found that wheat height is an important trait closely relevant to drought
resistance and seeds production. In this study, molecular mapping of the GA
insensitive gene Rht3 is carried using the DH population of Jinmai 5 and Lumai
14 (caning Rht gene). The results are as follows:
Lumai 14 carried a gene Rht3. It behaved as a dominant trait at seedlings and
adult stages. Molecular mapping indicated that this gene was located on the
chromosome 4B. Using 98 primers to amplify and screen the population, of them
43 primers have been found to be polymorphismic between the parents. However,
9 primers can amplified the polymorphism between the DNA pool of plant height.
Our results showed that Bulked Segregant Analysis and rnicrosatellite primer
screening of DH population identified three chromosome 4B microsatellite
markers, WMS368, WMS495 and WMS5 13, only WMS368 closely linked to
Rh13. The linkage distance between WMS368 and Rht3 was 15.3cM, WMS5 13,
WMS495 and Rht3 was 18.1cM, 26.9cM respectively.
The location of Rht3 gene and the acquisition of the molecular markers
enriched the genetic map of this linked group. They also provided valuable
information for our further research on the effects of this gene on wheat drought
resistance and seeds production, and for marker-assistant wheat breeding.
引文
1． 傅大雄，阮仁武等．小麦显性矮秆基因Rht10“成长”的发现．21世纪小麦遗传育种国际学术会议讨论会文集．2001，288-293
2． 贾继增．分子标记种质资源鉴定和分子标记育种．中国农业科学．1996，29(4)：1-10
3． 贾继增．小麦的矮秆基因及矮源．作物品种资源．1987，(3)：22-24
4． 贾继增，丁寿康等．小麦新品种宛原50-2矮秆基因的染色体定位．作物学报．1994，5：297-301
5． 贾继增，丁寿康等．中国小麦的主要矮秆基因及矮源的研究．中国农业科学．1992，25(1)：1-5
6． 景蕊莲，昌小平．SSR标记在小麦种质资源研究中的作用 作物品种资源．1999，2：17-20
7． 景蕊莲．作物抗旱相关性状的QTL定位研究进展(已投稿)．
8． 何中虎，黄钢等译．提高小麦产量潜力．中国科学技术出版社，北京，1999
9． 李生海，温辉芹等．冬小麦抗(耐)旱种质基因库的建拓研究．小麦显性矮秆基因ght10“成长”的发现．21世纪小麦遗传育种国际学术会议讨论会文集．2001，354-358
10． 刘秉华，杨丽．矮败小麦及利用前景．科学通报．1991，306-308
11． 刘秉华，杨丽．矮败小麦及其在矮化育种中应用．中国农业科学．1994，27(5)：17-21
12． 刘秉华，杨丽等．矮败小麦遗传研究．作物学报．1994，20(3)：306-309
13． 陆朝福，朱立煌．植物育种中分子标记辅助选择。生物工程进展．1995，15(4)：11-17
14． 马渐新，周荣华，董玉琛，贾继增．小麦抗条锈病基因定位及分子标记研究进展．生物技术同报．1999，15(1)：1-6
15． 孙美荣，李岩华等．水、旱交叉选育抗旱高产稳产小麦新品种的研究．小麦显性矮秆基因Rht10“成长”的发现。21世纪小麦遗传育种国际学术会议讨论会文集．2001，439-442
16． 王玉成，薛秀庄，唐国顺，王秋颖．“矮变一号”小麦株高的单体分析．作物学报．1982，8：193-198
17． 魏惠军，杜胜利，马德华．分子标记在黄瓜遗传育种研究中的应用．生物技术通报．1999，第二期
18．卫云宗，乔蕊清，谢福来，刘玲玲等．旱地小麦F1代主要性状杂种优势的遗传分析，麦类作物．1998，18(3)：8-10
19．吴兆苏编著．小麦育种学．农业出版社．1990，82-99
20．熊云仲，王石平，留克德，戴先凯，Saghai M．A，等．微卫星DNA和AFLP标记在水稻分子标记连锁图上的分布．植物学报．1998，40(7)：605-614
21．徐云碧，朱立煌著．分子数量遗传学．中国农业出版社．1994，86-113
22．徐吉臣，朱立煌．遗传图谱中的分子标记．生物工程进展．1992，12(5)：1-3
23．杨品官，Saghai Maroof M．A，张启发，秦昌华，罗忠训．水稻－多拷贝微卫星DNA多态性分析．遗传．1998，20(2)：27-30
24．张灿军，高海涛，王书子等．冬小麦水旱品种间杂交株高构成因素的遗传分析．小麦显性矮秆基因Rbt10“成长”的发现．21世纪小麦遗传育种国际学术会议讨论会文集．2001，435-438
25．张德水，陈受宜．DNA分子标记、基因组作图及其在植物遗传育种上的应用．生物技术通报．1998，5：15-22
26．张正斌，山仑．作物抗旱生理性状遗传研究进展．科学通报．1998，43(17)：1812-1817
27. Akkays MS, Bhagwat AA, Cregan PB. 1992, Length polymorphisms of simple sequence repeat DNA in Soybean. Genetic. 132:1131-1139
28. Becker and Heun. 1995, Mapping of digested and condigested random amplified microsatellite polymorphisms in barely. Genome, 38:991-998
29. Borner A., Roder M., Korzun V. 1995, Comparative molecular mapping of GA insensitive Rht loci on chromosomes 4B and 4D of common wheat (Triticum aestirum L.). Theor. Appl. Genet. 95:1133-1137
30. Cadalen T., Sourdile P., Charmet G., Tixier M. H., Gay G., Boeuf C., Bernard S., Leroy P., Bernard M. 1998, Molecular markers linked to genes affecting plant height in wheat using a doubled-haploid population. Theor. Appl. Genet. 96:933-940
31. Chap S., Sharp PJ, Worland AJ, Warham EJ et al. 1989, RFLP-based genetic maps of wheat homoelogous group 7 chromosomes. Theor. Appl. Genet. 78: 495-504
32. Daravsi A. et al. 1993, Selective DNA polling for determination of linkage between a maker locus and a quantative trait loci. TAG. 85:353-359
33. Eastwood R. F., Lagudah E. S., Apples R. 1994, Adirect search of DNA sequences
tightly linked to cereal cyst nematode resistance genes in Triticum tauschii. Genome 37: 311-319．
34． Ehdaie B. & Waines J.G. 1994, Growth and transpiration efficiency of near-isogenic lines for height in a spring wheat. Crop Sci. 34:6, 1443-1451
35． Fang D. Q., Roose M. L. 1997, Identification of closely related citrus cultivars with inter-simple sequence repeat markers. Theor. Appl. Genet. 66: 249-256
36． ForsterBP, Russed JR, Ellis RP, Haudley LL et al. 1997, Locating genotypes and genes for abiotic stress tolerance in barely: a strategy using maps markers and the wild soecies. New Phytol. 137: 141-147
37． Frova C., Krajeuski P., Viha M., Sari-Gorla M. 1999, Genetic analysis of drought resistance in maize by molecular markers. I. Yield Components. Theor. Appl. Genet. 99: 280-288
38． Henvy T., Nguyen, Chaudra Babu R., and Blum A. 1997, Breeding for Drought Resietance in Rice: Physiology and Molecular Genetics Coniderations. Crop Sci. 37: 1426-1434
39． Hesham A. S. , Agrama & Moumir E., Moussa. 1996, Mapping QTLs in breeding for drought tolerance in Maize ( Zea Mays L. ). Euphytica 91: 89-97
40． Hoz et al. 1996, Simple Sequence Repeats for gerplasm analysis and mapping in Maize. Genome. 39: 112-117
41． James C. Nelson, Allen E. Van Deynze, Enrique Autrique, Mark E. Sorrells, YunHai Lu, SylvieNegre, Michel Bernard, and Philippe Leroy. 1995, Molecular Mapping of wheat. Homoeologous Group 3． Genome. 38: 525-533
42． Kato K., Miura H., Sawade S., 1999, QTL mapping of genes controlling ear emergence time and plant height on chromosome 5A of wheat. Theor. Appl. Genet. 98: 472-477
43． Korzun V. et al. 1997 , Abstract of International Triticeae Mapping Initiative, Clermont-Ferrand, Framce June, 25-27
44． Korzun V. Melz G., Bormer A. 1996, RFLP mapping of the Dwarfing ( Ddw1 ) and Hairy peduncle (HP) genes on chromosome 5 of rye ( Secale cereal L ). Theor. Appl. Genet 92: 8, 1073-1077
45． Korzun V., RoderM. S., Ganal M. W., Worland A. J., Law C. N., 1998, Genetic analysis of the dwarfing gene (Rht8) in wheat. Part 1． Molecular mapping
of Rht8 on the short arm of chromosome 2Dof bread wheat ( Triticum aestivum L ). Theor. Appl. Genet 96: 1104-1109
46． Kristin A., Schneider, Mary E. Brothers, and Tames D. Kelly. 1997, Marker-Assisted Selection to Improve Drought Resietance in Common Bean. Crop. Sci. 37: 51-60
47． Law C. N. 1967, The location of genetic factors controlling a number of quantitative characters in wheat. Genetics 56: 445-461
48． Law C. N., Snape J. W & Worland A. J. 1981, Interspecific chromosome manipulation. Philosophical Transactions of the Royal Society of London (B) 292: 509-518
49． Lee SJ and Devos KM. 1996, Characterzation of loci containingmicrosatellite sequences among Canadian wheat cultivars. Genome. 39: 123-130
50． LevisionG., Gutman G. A. 1987, Slipped-strand mispairing: a major mechanism for DNA sequence evolution, Mol. Biol. Evol. 4: 203-221
51． Martin G. B., Williams J. G. K. , Tankley S. D. 1991, Rapid identification of markers linked to a Pseudomonas resistance gene in tomato by using random primers and near-isognic lines. Proc. Natl. Sci. USA 88: 2336-2340
52． Michelmore RW, Paran I, Kesseli RV. 1991, Identification of markers linked to disease resistance genes by bulked segregant analysis: a rapid method to detect markers in specific genomic regions by using segregating population. Proc Natl Acad Sci USA 88: 9828-9832
53． Moore G., Devos K. M., Wang Z., Gale M. D. 1995, Cereal genome evolution: Grasses, line up and from a circle. Current Biology 5: 737-740
54． Morgan J M., Tan M K., 1996, Chromosomal location of a wheat osmoregulation gene using RFLP analysis. Aust. J. Plant Physiol. 23: 803-806
55． MorganteM, Olivieri. 1993, PCR-amplifiedmicrosatellite as markers in plant genetics. Plant J. 3: 175-182
56． Nagaoka T, Ogihara Y. 1997, Applicability of inter-simple sequence repeat polymorphhisn in wheat for use as DNA markers in comparison to RFLP and RAPD markers. Theor. Appl. Genet. 94: 597-602
57． Patnaparkwe MB, Tekeoglu M, Muehlbauer H. 1998 Inter-simple-sequence-repeat (ISSR) polymorphism are useful for finding markers associated with disease resistance gene clusters. Theor. Appl. Genet.
97: 515-519
58． Payburn AL, Gill BS. 1985, Use of biotin-labeled probes to map specific DNA sequences on wheat chromosomes. Heredity. 76: 78-81
59． Phinney B. O., Spray C. 1982, Chemical genetics and the gibberellin pathway in Zea Mays L. In plant Growth Substances. Aberystwyth, Wales, July 1982． pp101-110． Ed. By P. E. Wareing. Academic Press, New York
60． Plaschke J., Korzun V., Koebner R. M. D., Borner A. 1995, Mapping the GA3~insensitive dwarfing gene ct1 on chromosome 7 in rye. Plant Breeding 114: 113-116
61． Roder M. S., Korzun J., Wendehake K. et al, 1998, A microsatellite map of wheat. Genetics 149: 2007-2023
62． Roder MS, Plashke J, Konig SU, Borner A, Sorrells ME, Tanksley SD, Ganal MW. 1995, Abundance variability and chromosomal location of microsatellite in wheat. Mol. Gen. Genet. 246: 327-333
63． Schondelmaier J., Steinruken G., Jung C, 1996, Amplified fragment length polymorphism (AFLP) in soybean: species diversity , inheritance, and near-isogenic line analysis. Theor. Appl. Genet. 93: 3, 392-401
64． Skirver D, Mundy J. 1990, Gene expression in response to abscisic acid and osmotic stress. Plant Cell. 2: 503-512
65． Sugi H., and Murakami Y. 1968, Occurrence of a rice mutant deficient in gibberellin-like substances. Plant Cell Physiology. 9: 411-414
66． Tanksley S. D., Young N. D., Paterson A. H. et al. 1989, RFLP mapping in plant breeding: New tools for an old science. Bio/Technology 7: 257-264．
67． Tsumura Y., Ohba K., Strauss S. 1996, Diversity and inheritance of inter-simple sequence repeat polymorphisms in Douglas-fir ( Pseudotsuga menziesii ) and sugi ( Crytomeria japonica ), Theor. Appl. Genet. 92: 40-45
68． Wu KS, Tanksley SD. 1993, Abundance polymorphism and genetic mapping of microsatellite in rice. Mol. Gen. Genet. 241: 225-235
69． Zeng 7 B., Kao C H 1998, In: Abstract of 18th International Congress of Genetics. Beijing, China. P132
70． Zeven A. C., Knott D. R., Johnson R. 1983, Investigation of linkage drag in neat isogenic lines of wheat by testing for seedling reaction to races of stem rust, leaf rust and yellow rust. Euphytica 32: 319-327
2． 贾继增．分子标记种质资源鉴定和分子标记育种．中国农业科学．1996，29(4)：1-10
3． 贾继增．小麦的矮秆基因及矮源．作物品种资源．1987，(3)：22-24
4． 贾继增，丁寿康等．小麦新品种宛原50-2矮秆基因的染色体定位．作物学报．1994，5：297-301
5． 贾继增，丁寿康等．中国小麦的主要矮秆基因及矮源的研究．中国农业科学．1992，25(1)：1-5
6． 景蕊莲，昌小平．SSR标记在小麦种质资源研究中的作用 作物品种资源．1999，2：17-20
7． 景蕊莲．作物抗旱相关性状的QTL定位研究进展(已投稿)．
8． 何中虎，黄钢等译．提高小麦产量潜力．中国科学技术出版社，北京，1999
9． 李生海，温辉芹等．冬小麦抗(耐)旱种质基因库的建拓研究．小麦显性矮秆基因ght10“成长”的发现．21世纪小麦遗传育种国际学术会议讨论会文集．2001，354-358
10． 刘秉华，杨丽．矮败小麦及利用前景．科学通报．1991，306-308
11． 刘秉华，杨丽．矮败小麦及其在矮化育种中应用．中国农业科学．1994，27(5)：17-21
12． 刘秉华，杨丽等．矮败小麦遗传研究．作物学报．1994，20(3)：306-309
13． 陆朝福，朱立煌．植物育种中分子标记辅助选择。生物工程进展．1995，15(4)：11-17
14． 马渐新，周荣华，董玉琛，贾继增．小麦抗条锈病基因定位及分子标记研究进展．生物技术同报．1999，15(1)：1-6
15． 孙美荣，李岩华等．水、旱交叉选育抗旱高产稳产小麦新品种的研究．小麦显性矮秆基因Rht10“成长”的发现。21世纪小麦遗传育种国际学术会议讨论会文集．2001，439-442
16． 王玉成，薛秀庄，唐国顺，王秋颖．“矮变一号”小麦株高的单体分析．作物学报．1982，8：193-198
17． 魏惠军，杜胜利，马德华．分子标记在黄瓜遗传育种研究中的应用．生物技术通报．1999，第二期
18．卫云宗，乔蕊清，谢福来，刘玲玲等．旱地小麦F1代主要性状杂种优势的遗传分析，麦类作物．1998，18(3)：8-10
19．吴兆苏编著．小麦育种学．农业出版社．1990，82-99
20．熊云仲，王石平，留克德，戴先凯，Saghai M．A，等．微卫星DNA和AFLP标记在水稻分子标记连锁图上的分布．植物学报．1998，40(7)：605-614
21．徐云碧，朱立煌著．分子数量遗传学．中国农业出版社．1994，86-113
22．徐吉臣，朱立煌．遗传图谱中的分子标记．生物工程进展．1992，12(5)：1-3
23．杨品官，Saghai Maroof M．A，张启发，秦昌华，罗忠训．水稻－多拷贝微卫星DNA多态性分析．遗传．1998，20(2)：27-30
24．张灿军，高海涛，王书子等．冬小麦水旱品种间杂交株高构成因素的遗传分析．小麦显性矮秆基因Rbt10“成长”的发现．21世纪小麦遗传育种国际学术会议讨论会文集．2001，435-438
25．张德水，陈受宜．DNA分子标记、基因组作图及其在植物遗传育种上的应用．生物技术通报．1998，5：15-22
26．张正斌，山仑．作物抗旱生理性状遗传研究进展．科学通报．1998，43(17)：1812-1817
27. Akkays MS, Bhagwat AA, Cregan PB. 1992, Length polymorphisms of simple sequence repeat DNA in Soybean. Genetic. 132:1131-1139
28. Becker and Heun. 1995, Mapping of digested and condigested random amplified microsatellite polymorphisms in barely. Genome, 38:991-998
29. Borner A., Roder M., Korzun V. 1995, Comparative molecular mapping of GA insensitive Rht loci on chromosomes 4B and 4D of common wheat (Triticum aestirum L.). Theor. Appl. Genet. 95:1133-1137
30. Cadalen T., Sourdile P., Charmet G., Tixier M. H., Gay G., Boeuf C., Bernard S., Leroy P., Bernard M. 1998, Molecular markers linked to genes affecting plant height in wheat using a doubled-haploid population. Theor. Appl. Genet. 96:933-940
31. Chap S., Sharp PJ, Worland AJ, Warham EJ et al. 1989, RFLP-based genetic maps of wheat homoelogous group 7 chromosomes. Theor. Appl. Genet. 78: 495-504
32. Daravsi A. et al. 1993, Selective DNA polling for determination of linkage between a maker locus and a quantative trait loci. TAG. 85:353-359
33. Eastwood R. F., Lagudah E. S., Apples R. 1994, Adirect search of DNA sequences
tightly linked to cereal cyst nematode resistance genes in Triticum tauschii. Genome 37: 311-319．
34． Ehdaie B. & Waines J.G. 1994, Growth and transpiration efficiency of near-isogenic lines for height in a spring wheat. Crop Sci. 34:6, 1443-1451
35． Fang D. Q., Roose M. L. 1997, Identification of closely related citrus cultivars with inter-simple sequence repeat markers. Theor. Appl. Genet. 66: 249-256
36． ForsterBP, Russed JR, Ellis RP, Haudley LL et al. 1997, Locating genotypes and genes for abiotic stress tolerance in barely: a strategy using maps markers and the wild soecies. New Phytol. 137: 141-147
37． Frova C., Krajeuski P., Viha M., Sari-Gorla M. 1999, Genetic analysis of drought resistance in maize by molecular markers. I. Yield Components. Theor. Appl. Genet. 99: 280-288
38． Henvy T., Nguyen, Chaudra Babu R., and Blum A. 1997, Breeding for Drought Resietance in Rice: Physiology and Molecular Genetics Coniderations. Crop Sci. 37: 1426-1434
39． Hesham A. S. , Agrama & Moumir E., Moussa. 1996, Mapping QTLs in breeding for drought tolerance in Maize ( Zea Mays L. ). Euphytica 91: 89-97
40． Hoz et al. 1996, Simple Sequence Repeats for gerplasm analysis and mapping in Maize. Genome. 39: 112-117
41． James C. Nelson, Allen E. Van Deynze, Enrique Autrique, Mark E. Sorrells, YunHai Lu, SylvieNegre, Michel Bernard, and Philippe Leroy. 1995, Molecular Mapping of wheat. Homoeologous Group 3． Genome. 38: 525-533
42． Kato K., Miura H., Sawade S., 1999, QTL mapping of genes controlling ear emergence time and plant height on chromosome 5A of wheat. Theor. Appl. Genet. 98: 472-477
43． Korzun V. et al. 1997 , Abstract of International Triticeae Mapping Initiative, Clermont-Ferrand, Framce June, 25-27
44． Korzun V. Melz G., Bormer A. 1996, RFLP mapping of the Dwarfing ( Ddw1 ) and Hairy peduncle (HP) genes on chromosome 5 of rye ( Secale cereal L ). Theor. Appl. Genet 92: 8, 1073-1077
45． Korzun V., RoderM. S., Ganal M. W., Worland A. J., Law C. N., 1998, Genetic analysis of the dwarfing gene (Rht8) in wheat. Part 1． Molecular mapping
of Rht8 on the short arm of chromosome 2Dof bread wheat ( Triticum aestivum L ). Theor. Appl. Genet 96: 1104-1109
46． Kristin A., Schneider, Mary E. Brothers, and Tames D. Kelly. 1997, Marker-Assisted Selection to Improve Drought Resietance in Common Bean. Crop. Sci. 37: 51-60
47． Law C. N. 1967, The location of genetic factors controlling a number of quantitative characters in wheat. Genetics 56: 445-461
48． Law C. N., Snape J. W & Worland A. J. 1981, Interspecific chromosome manipulation. Philosophical Transactions of the Royal Society of London (B) 292: 509-518
49． Lee SJ and Devos KM. 1996, Characterzation of loci containingmicrosatellite sequences among Canadian wheat cultivars. Genome. 39: 123-130
50． LevisionG., Gutman G. A. 1987, Slipped-strand mispairing: a major mechanism for DNA sequence evolution, Mol. Biol. Evol. 4: 203-221
51． Martin G. B., Williams J. G. K. , Tankley S. D. 1991, Rapid identification of markers linked to a Pseudomonas resistance gene in tomato by using random primers and near-isognic lines. Proc. Natl. Sci. USA 88: 2336-2340
52． Michelmore RW, Paran I, Kesseli RV. 1991, Identification of markers linked to disease resistance genes by bulked segregant analysis: a rapid method to detect markers in specific genomic regions by using segregating population. Proc Natl Acad Sci USA 88: 9828-9832
53． Moore G., Devos K. M., Wang Z., Gale M. D. 1995, Cereal genome evolution: Grasses, line up and from a circle. Current Biology 5: 737-740
54． Morgan J M., Tan M K., 1996, Chromosomal location of a wheat osmoregulation gene using RFLP analysis. Aust. J. Plant Physiol. 23: 803-806
55． MorganteM, Olivieri. 1993, PCR-amplifiedmicrosatellite as markers in plant genetics. Plant J. 3: 175-182
56． Nagaoka T, Ogihara Y. 1997, Applicability of inter-simple sequence repeat polymorphhisn in wheat for use as DNA markers in comparison to RFLP and RAPD markers. Theor. Appl. Genet. 94: 597-602
57． Patnaparkwe MB, Tekeoglu M, Muehlbauer H. 1998 Inter-simple-sequence-repeat (ISSR) polymorphism are useful for finding markers associated with disease resistance gene clusters. Theor. Appl. Genet.
97: 515-519
58． Payburn AL, Gill BS. 1985, Use of biotin-labeled probes to map specific DNA sequences on wheat chromosomes. Heredity. 76: 78-81
59． Phinney B. O., Spray C. 1982, Chemical genetics and the gibberellin pathway in Zea Mays L. In plant Growth Substances. Aberystwyth, Wales, July 1982． pp101-110． Ed. By P. E. Wareing. Academic Press, New York
60． Plaschke J., Korzun V., Koebner R. M. D., Borner A. 1995, Mapping the GA3~insensitive dwarfing gene ct1 on chromosome 7 in rye. Plant Breeding 114: 113-116
61． Roder M. S., Korzun J., Wendehake K. et al, 1998, A microsatellite map of wheat. Genetics 149: 2007-2023
62． Roder MS, Plashke J, Konig SU, Borner A, Sorrells ME, Tanksley SD, Ganal MW. 1995, Abundance variability and chromosomal location of microsatellite in wheat. Mol. Gen. Genet. 246: 327-333
63． Schondelmaier J., Steinruken G., Jung C, 1996, Amplified fragment length polymorphism (AFLP) in soybean: species diversity , inheritance, and near-isogenic line analysis. Theor. Appl. Genet. 93: 3, 392-401
64． Skirver D, Mundy J. 1990, Gene expression in response to abscisic acid and osmotic stress. Plant Cell. 2: 503-512
65． Sugi H., and Murakami Y. 1968, Occurrence of a rice mutant deficient in gibberellin-like substances. Plant Cell Physiology. 9: 411-414
66． Tanksley S. D., Young N. D., Paterson A. H. et al. 1989, RFLP mapping in plant breeding: New tools for an old science. Bio/Technology 7: 257-264．
67． Tsumura Y., Ohba K., Strauss S. 1996, Diversity and inheritance of inter-simple sequence repeat polymorphisms in Douglas-fir ( Pseudotsuga menziesii ) and sugi ( Crytomeria japonica ), Theor. Appl. Genet. 92: 40-45
68． Wu KS, Tanksley SD. 1993, Abundance polymorphism and genetic mapping of microsatellite in rice. Mol. Gen. Genet. 241: 225-235
69． Zeng 7 B., Kao C H 1998, In: Abstract of 18th International Congress of Genetics. Beijing, China. P132
70． Zeven A. C., Knott D. R., Johnson R. 1983, Investigation of linkage drag in neat isogenic lines of wheat by testing for seedling reaction to races of stem rust, leaf rust and yellow rust. Euphytica 32: 319-327