中国乌骨鸡遗传多样性及家鸡起源研究
|
摘要
本文共测定了19个乌骨鸡品种(品系)及8个非乌骨鸡品种(品系)的243个家鸡和38个原鸡个体(属于Gallus.gallus.spadiceus和Gallus.gallus.jabouillee亚种)的线粒体D-环高变区(mtDNA HVS-Ⅰ)539bp的部分序列，另外还测定了5个红原鸡个体线粒体细胞色素b的289 bp序列、19个珍珠鸡个体、18个骡鸡个体的线粒体D环高变区序列、30个家鸡个体线粒体D环非高变区的690bp序列以及34个家鸡及原鸡个体的MC1R基因编码区的945bp。对这些序列片段及其它已经发表的相应序列片段进行了邻接法、网络图法等分析，得出如下主要结论：
1、8个丝羽乌骨鸡群体的66个个体的线粒体D环539bp中共有23个变异位点，14种单倍型，分析表明丝羽乌骨鸡来源于多个母系，是在不同的地方长期人工选育的结果。
2、124个常羽乌骨鸡的线粒体D环高变区序列中共有39个变异位点，可分为34种单倍型，从中可以看出常羽乌骨鸡的母系来源类型很多，至少有六个不同的原鸡母系来源。
3、丝羽乌骨鸡为常羽乌骨鸡的突变型，常羽乌骨鸡有比丝羽乌骨鸡更多的单倍型类型，而且在丝羽乌骨鸡群体中出现的几种主体单倍型类型也是常羽乌骨鸡群体的主体单倍型。
4、在中国的两个红原鸡亚种的线粒体D环高变区序列与在泰国等地分布的红原鸡亚种的此段序列差异不大，可归为一类，而与爪哇岛上分布的红原鸡亚种的此段序列差异较大。我们的结果支持红原鸡可以分为陆地型亚种和海岛型亚种。
5、314个家鸡个体和62个原鸡个体的线粒体高变区序列共有66个变异位点，79种单倍型，可划分为9大类群。家鸡和原鸡共同拥有的单倍型有7个，由此可推论所有家鸡都有共同的祖先，可能都来源于陆地型红原鸡，并经受了多次独立驯化。
6、在本实验中，线粒体D环非高变区序列对家鸡起源分析的结果同于高变区序列分析，所以，选择高变区来研究物种的遗传多样性和起源进化是经济、有效的。
7、在家鸡和原鸡黑色素受体1(MC1R)基因编码区的945个碱基中，34个个体共有10个变异位点，其中第28位、69位、159位、637位为同义突变；翻译成氨基酸后，有变异的6个氨基酸的位置分别是10、71、92、213、215、307，其中71、213、215、307有氨基酸杂合的情况。在这些突变的氨基酸位点中，突变与羽毛颜色间除了丝羽乌鸡外均有一一对应关系，这可能提示乌鸡的丝羽
口
及颜色受多个基因的调控，MC IR基因可能是其中的一个主效但不是唯一的质
量性状基因。
8、黑羽常羽乌骨鸡和珍珠鸡的线粒体D环高变区序列存在长度变异，乌骨鸡的此
段序列长度为539hp，珍珠鸡的序列是498hp，而骡鸡个体的此段序列或同于乌
骨鸡、或同于珍珠鸡。因此，本结果支持骡鸡是乌骨鸡和珍珠鸡的杂交后代。
In the present paper, we determined the mitochondrial DNA sequence 539bp of the high variable I D-loop regions(mtDNA HVS-I) for 243 domestic fowls (19 were Wugu (or black-bone chicken) and 8 non-wugu breeds/races) and 38 red jungle fowls (Gallus. Gallus. spadiceus and Gallus. Gallus. jabouillei). We also sequenced 289bp mitochondrial DNA of Cyt-b for 5 red jungle fowls; D-loop region (mtDNA HVS-I)for 19 individuals Guinea fowl and 18 individuals mule chickens (Guinea fowl $ x Wugu chickens^ ) respectively; mtDNA HVS- II of D-loop region for 34 individuals of domestic fowls and red jungle fowls in 690bp and the MC1R code gene in 945 bp. Analyzing this data together with the same sequence part available in genbank by neighbor joining and network etc, we can draw some conclusion as follow:
1. There were 23 mutation sites in mtDNA HVS-I of 66 individuals among 8 populations; it could be divided into 14 haplotypes. The result suggested that Silkies had several maternal ancestors. It was the result of long artificial selection in different place.
2. There were 39 mutation sites in mtDNA HVS-I of 124 individuals; it could be divided into 34 haplotypes. The normal feather Wugu chickens also had a lot of maternal ancestors; the result showed that at least six populations of red jungle fowl contributed to the gene pool of the domestic fowls.
3. Silkies were the mutation type of nomal feather Wugu chicken for it had more haplotypes and shared the major haplotypes with silkies.
4. The sub-species of G. g. spadcieus and G. g. jabouillee in China were at the same clade with the G. g. gallus in and around the Thailand area, but were in difference clade with G. g. bankiva in Java island, so our result support the opinion that red jungle fowl could be classified as mainland type and island type.
5. There were 66 mutation sites in mtDNA HVS-I of 314 domestic fowls and 62 red jungle fowls, it could be divided into 79 haplotypes, these haplotypes belong to nine clades. So, All the domestic fowl shared the same maternal ancestor of red jungle fowl and were domesticated several times independently in different places.
6. In our studying, the mtDNA HVS-I and the mtDAN HVS- II showed the same result, so, it was suitable , high efficiency and economical to choose mtDNA HVS-I as a marker studying for the genetic divergence and evolution of species.
7. There were 10 mute sites in the MC1R gene in 945 bp code region of the domestic fowls and red jungle fowls in all 34 individuals, in these mute sites, the 28th, 69th, 159th, 637th point mute sites were synonymous mutes, while the 10th 71th,92th, 2135h, 215th, 307th sites were amino acid mutation sites, among them , 71th, 213th, 215th, 307th were hybrid sites. We couldn't see the line-relationship of the mute sites and the feather color types; maybe MC1R was just one major but not only silky feather color functional gene, the feather color maybe controlled by several genes.
8. There were length differences when using the same primer to amplify the mtDNA HVS-1 of black normal feather Wugu chickens and Guinea fowl. The PCR production sequence length of them were 539bp and 498bp respectively, the same region sequence of mule chicken were the same as Guinea fowl or black normal feather Wugu chickens. So, it suggested that the mule chickens were the progeny of Guinea fowl and the black normal feather Wugu chicken.
1、8个丝羽乌骨鸡群体的66个个体的线粒体D环539bp中共有23个变异位点，14种单倍型，分析表明丝羽乌骨鸡来源于多个母系，是在不同的地方长期人工选育的结果。
2、124个常羽乌骨鸡的线粒体D环高变区序列中共有39个变异位点，可分为34种单倍型，从中可以看出常羽乌骨鸡的母系来源类型很多，至少有六个不同的原鸡母系来源。
3、丝羽乌骨鸡为常羽乌骨鸡的突变型，常羽乌骨鸡有比丝羽乌骨鸡更多的单倍型类型，而且在丝羽乌骨鸡群体中出现的几种主体单倍型类型也是常羽乌骨鸡群体的主体单倍型。
4、在中国的两个红原鸡亚种的线粒体D环高变区序列与在泰国等地分布的红原鸡亚种的此段序列差异不大，可归为一类，而与爪哇岛上分布的红原鸡亚种的此段序列差异较大。我们的结果支持红原鸡可以分为陆地型亚种和海岛型亚种。
5、314个家鸡个体和62个原鸡个体的线粒体高变区序列共有66个变异位点，79种单倍型，可划分为9大类群。家鸡和原鸡共同拥有的单倍型有7个，由此可推论所有家鸡都有共同的祖先，可能都来源于陆地型红原鸡，并经受了多次独立驯化。
6、在本实验中，线粒体D环非高变区序列对家鸡起源分析的结果同于高变区序列分析，所以，选择高变区来研究物种的遗传多样性和起源进化是经济、有效的。
7、在家鸡和原鸡黑色素受体1(MC1R)基因编码区的945个碱基中，34个个体共有10个变异位点，其中第28位、69位、159位、637位为同义突变；翻译成氨基酸后，有变异的6个氨基酸的位置分别是10、71、92、213、215、307，其中71、213、215、307有氨基酸杂合的情况。在这些突变的氨基酸位点中，突变与羽毛颜色间除了丝羽乌鸡外均有一一对应关系，这可能提示乌鸡的丝羽
口
及颜色受多个基因的调控，MC IR基因可能是其中的一个主效但不是唯一的质
量性状基因。
8、黑羽常羽乌骨鸡和珍珠鸡的线粒体D环高变区序列存在长度变异，乌骨鸡的此
段序列长度为539hp，珍珠鸡的序列是498hp，而骡鸡个体的此段序列或同于乌
骨鸡、或同于珍珠鸡。因此，本结果支持骡鸡是乌骨鸡和珍珠鸡的杂交后代。
In the present paper, we determined the mitochondrial DNA sequence 539bp of the high variable I D-loop regions(mtDNA HVS-I) for 243 domestic fowls (19 were Wugu (or black-bone chicken) and 8 non-wugu breeds/races) and 38 red jungle fowls (Gallus. Gallus. spadiceus and Gallus. Gallus. jabouillei). We also sequenced 289bp mitochondrial DNA of Cyt-b for 5 red jungle fowls; D-loop region (mtDNA HVS-I)for 19 individuals Guinea fowl and 18 individuals mule chickens (Guinea fowl $ x Wugu chickens^ ) respectively; mtDNA HVS- II of D-loop region for 34 individuals of domestic fowls and red jungle fowls in 690bp and the MC1R code gene in 945 bp. Analyzing this data together with the same sequence part available in genbank by neighbor joining and network etc, we can draw some conclusion as follow:
1. There were 23 mutation sites in mtDNA HVS-I of 66 individuals among 8 populations; it could be divided into 14 haplotypes. The result suggested that Silkies had several maternal ancestors. It was the result of long artificial selection in different place.
2. There were 39 mutation sites in mtDNA HVS-I of 124 individuals; it could be divided into 34 haplotypes. The normal feather Wugu chickens also had a lot of maternal ancestors; the result showed that at least six populations of red jungle fowl contributed to the gene pool of the domestic fowls.
3. Silkies were the mutation type of nomal feather Wugu chicken for it had more haplotypes and shared the major haplotypes with silkies.
4. The sub-species of G. g. spadcieus and G. g. jabouillee in China were at the same clade with the G. g. gallus in and around the Thailand area, but were in difference clade with G. g. bankiva in Java island, so our result support the opinion that red jungle fowl could be classified as mainland type and island type.
5. There were 66 mutation sites in mtDNA HVS-I of 314 domestic fowls and 62 red jungle fowls, it could be divided into 79 haplotypes, these haplotypes belong to nine clades. So, All the domestic fowl shared the same maternal ancestor of red jungle fowl and were domesticated several times independently in different places.
6. In our studying, the mtDNA HVS-I and the mtDAN HVS- II showed the same result, so, it was suitable , high efficiency and economical to choose mtDNA HVS-I as a marker studying for the genetic divergence and evolution of species.
7. There were 10 mute sites in the MC1R gene in 945 bp code region of the domestic fowls and red jungle fowls in all 34 individuals, in these mute sites, the 28th, 69th, 159th, 637th point mute sites were synonymous mutes, while the 10th 71th,92th, 2135h, 215th, 307th sites were amino acid mutation sites, among them , 71th, 213th, 215th, 307th were hybrid sites. We couldn't see the line-relationship of the mute sites and the feather color types; maybe MC1R was just one major but not only silky feather color functional gene, the feather color maybe controlled by several genes.
8. There were length differences when using the same primer to amplify the mtDNA HVS-1 of black normal feather Wugu chickens and Guinea fowl. The PCR production sequence length of them were 539bp and 498bp respectively, the same region sequence of mule chicken were the same as Guinea fowl or black normal feather Wugu chickens. So, it suggested that the mule chickens were the progeny of Guinea fowl and the black normal feather Wugu chicken.
引文
1．陈灵芝 主编 1993，中国的生物多样性——现状及其保护对策99-113科学出版社 北京
2．施立明，1990 遗传多样性及其保护层生命科学信息 2：158-164
3．史宪伟 1997 中国主要家鹅品种遗传多样性及其起源分化研究 博士学位论文
4．黄勇富 1996 中国部分地方猪种遗传多样性及起源分化研究 博士学位论文
5．吕雪梅等 1996 以RAPD标记分析广东四个鸡品种的亲缘关系
6．洪夏等 1997 河田鸡线粒体DNA的物理图谱。第八次全国家禽学术讨论会论文集。东北林大出版社 49-51
7．孙汉等 1997 江西地方鸡种六种血清酶多态性的研究 第八次全国家禽学术讨论会论文集东北林大出版社 82-85
8．吴常信 1999 对“畜禽遗传资源保护和利用”立项研究的建议 第九次全国家禽学术讨论会论文集 1-3
9．顾志良 2000 我国地方鸡种遗传多样性研究进展 中国家禽 7
10．陈幼春 1995 中国家畜多样性保护的意义 生物多样性 3(3)：143-146。
11．邱祥聘，陈锷，陈育新等 1988 中国家禽品种志 上海科技出版社 上海 3-16。
12．刘丑生，魏绍征 1995 DNA指纹的研究及其在畜禽遗传育种中的应用 甘肃畜牧兽医 (5)：27-29。
13．丁波 1994 DNA指纹技术及其在畜禽遗传育种中的应用 青海畜牧兽医杂志 (4)：42-44。
14．王金玉，龚允陈等 1995 鸡 DNA 指纹遗传研究进展及其在育种上的应用 中国家禽 (5)：29-31。
15．魏彩虹 1997 动物mtDNA遗传研究进展 甘肃畜牧兽医 (1)：23-25。
16．李建凡 1995 家禽分子生物学技术研究进展 国外畜牧科技 (5)：53-57。
17．孟安明 1993 DNA指纹技术在农业上的应用 生物工程进展 13(3)。
18．张沅 1991 生物技术在动物育种中应用的现状与前景 遗传 13(4)：39-42。
19．孟安明，齐顺章，周顺伍，宫桂芬 1993 家禽遗传图谱初报 遗传 15(2)：9-13。
20．曾福玪，李宁，王峰等 1995．鹌鹑品种(系)的随机扩增多态性DNA指纹分析 农业生物技术学报 3：61-64。
21．杨丽萍，1999 RAPD技术在家兔遗传分析中的应用。硕士学位论文。中国农科院。
22．王晓梅，宋文芹等，1998 用RAPD技术检测野生鲫鱼和四个金鱼代表品种的基因组DNA多态性。遗传20(5)：7-11。
23．李祥龙 1997 中国主要山羊品种遗传多样性及起源分化研究 四川农业大学博士论文。
24．张细权，吕学梅，杨玉华等，1998 用微卫星多态性和RAPD分析广东地方鸡种的群体遗传变异。遗传学报 25(2)：112—119。
25．周怀军，陈宽维等 1997 RAPD技术对地方鸡种群体遗传结构的分析 中国家禽 5(7)。
26．张学余，黄凡美等 1998 乌骨鸡种群遗传结构的RAPD分析 中国家禽 5(1)。
27．刘浚凡等 1998 矮小型黑羽乌骨鸡选育 中国家禽 1998 1(8)。
28．扬志勤 1997 乌骨鸡的品种与开发 (王红宁，朱庆 主编：养禽与禽病防治新进展) 四川科技出版社。42-45。
29．龚炎长，彭中镇等，1988。畜禽微卫星的应用。34(6)：51-52。
30．顾志良，王志跃，张军。1997。微卫星DNA标记及其在鸡的基因图谱构建中的应用。中国家禽。(6)：27-29。
31．刘浚凡等，1998。矮小型黑羽乌骨鸡选育 中国家禽 1(8)。
32．邱祥聘等，1999。丝羽乌骨鸡新品系的选育工作初报 第九次全国家禽学术讨论会论文集 7-10。
33．罗正雄等，1999。盐津乌骨鸡肉用性能的研究 第九次全国家禽学术讨论会论文集 86-91。
34．杜金平等，1999。金水乌骨鸡选育综合报告第九次全国家禽学术讨论会论文集 91-94。
35．金深逊等，1999。乌蒙黄羽乌骨鸡肉用性能研究 第九次全国家禽学术讨论会论文集 96-98。
36．金冬雁，金奇，侯云德，1996．核酸和蛋白质的化学合成与序列分析，北京：科学
37．出版社，49-70
38．张亚平．1996．从DNA序列到物种树，动物学研究，17(3)：247-252
39．张亚平，施立明．1992．动物线粒体DNA多态性的研究概况．动物学研究，13(3)：289—298
40．刘树俊，程光潮．1992．DNA指纹技术的应用及局限性．生物工程进展，12(5)：13-17
41．王斌，翁曼丽．1996．AFLP的原理及其应用．杂交水稻，(5)：27—30
42．包世增，家禽育种学，1994，29-43。北京：农业出版社
43．常洪，家畜遗传资源学纲要，1995，44-45。北京：中国农业出版社。
44．张细权，李加琪，杨关福，动物遗传标记，1997，7-8，中国农业大学出版社。
45．徐幸莲等，黑色素的研究进展，中国畜产与食品，1997，4(3)135-136。
46．江志洁等，黑色素形成机理的新概念及复合美白剂的应用，日用化学品科学，1998(4)：138-140。
47．徐幸莲等，乌骨鸡黑色素对延缓果蝇衰老的作用，南京农业大学学报，1999，22(2)105-108。
48．贾青等，泰和鸡黑色素的发生初探，河北农业大学学报．1998，21(3)．72-74。
49．吴常信主编，21世纪中国家禽业的研究与发展---第九次全国家禽学术讨论会论文集，1999．11，上海。
50．耿拓宇等，泰和乌鸡出雏后黑色素的分布与沉积，中国家禽，2000，22(7)：10-12。
51．朱子玉等，丝毛乌骨鸡和北京油鸡超氧化物歧化酶活性比较研究，中国家禽，2000(10) 6-7。
52. Abed, Y., Davinj-Regli,A.,Bollet,C.,De Micco,P.1995. Efficient discrimination of Mycobacherium tubemlosis strains by 16s spacer region based random amplified polymorphic DNA analysis. Clin Microbiol.33:1448-1420
53. Adachi S, Morii E, Kim D, Ogihara H, Jippo T, Ito A, Lee Y-M, Y Kitamura.Involvement of mi-transcription factor in expression of α-melanocyte-stimulating hormone receptor in cultured mast cells of mice. J. Immunology, 2000.164:855～860
54. Akahashi, K., and M. Nei. 2000. Efficiencies of fast algorithms of phylogenetic inference under the criteria of maximum parsimony, minimum evolution, and maximum likelihood when a large number of sequences are used. Molecular Biology and Evolution 17:1251-1258.
55. Akitoshi NOZAKI et al; The Surface color Measurement of Major tissues of Silky Fowls and White Leghorns, J.Vet. Med. Sci. 1998,60:489-493
56. Ashwell, M.S. Rexroad, C. E. Miller. J.R.H, etal. 1992. Animal genetics. 23:133-142.
57. Ayala F.J & J.W, Valentine 1991. Evolving: the theory and progress of organic evolution: menlo park: Benjamin-cummings.
58. B. Van Wijk. Wildlife management principles. What is biodiversity 103-104
59. Ballou, J.D; M.Gilpin, T.J, foise, eds. Population management for survival and recovery,analytical methods and strategies in small population conservation. Columbia University
Press, New York, 1995．
60． Barnes,Nick: conflicts over biodiversty. In environmental policy in an international context part SOU, 1996．
61． Boston B A, Cone R D.Characterization of melanocortin receptor subtype expression in murine adipose tissues and in the 3T3-L1 cell line. Endocrinology , 1996． 137:2043-2050
62． Brown W.M.,et al. 1979． Rapid evolution of animal mitochondrial DNA. Proc. Natl. Acad. Sci. USA. 76:1967-1971．
63． Brown,W.M. 1983． Evolution of animal mtDNA. In:Nei,M.,and Koehn,R.K. (eds.). Evolution of Gene and Proteins,sinauer,sunderland,62-68
64． Bultman S P, Michaud E J, Woychik R P. Molecular characterisation of the mouse agouti locus. Cell ,1992． 71:1-20
65． Chakraborty A, Slominski A, Ermak G, Hwang J, Pawelek J. Ultraviolet B and melanocyte-stimulating hormone (MSH) stimulate mRNA production for alpha MSH receptors and proopiomelanocortin-derived peptides in mouse melanoma cells and transformed keratinocytes. J Invest Dermatol, 1995． 105:655-659
66． Cheng H. Etal. 1994． Poultry science. 73: 539-546; 74: 1855-1874．
67． Chhajlani V, Muceniece R, Wiaberg J E S. Moleculat cloning of a novel human menocortin-1 receptor. Biochem Biophys Res Commun,1993． 195:866-873
68． Chhajlani V, Wikberg J E S. Molecular cloning and expression of the human melanocyte stimulating hormone receptor cDNA. FEBS Lett, 1992． 309: 417-420．
69． Chhajlani V, Xu X, Blauw J, Sudarshi S. Identification of ligand binding residues in extracellular loops of the melanocortin 1 receptor. Biochem Biophys Res Commun Feb, 1996． 15;219(2) :521-525
70． Chluba-de Tapia J, Bagutti C, Cotti R, Eberle A N. Induction of constitutive melanogenesis in amelanotic mouse melanoma cells by transfection of the human melanocortin-1 receptor gene. J Cell Sci, 1996． 109 (Pt 8) :2023-2030．
71． Cone R D, Lu D, Koppula S, Vage D I, Klungland H, Boston B, Chen W, Orth D N, Pouton C, Kesterson R A. The melanocortin receptors: agonists, antagonists, and the hormonal control of pigmentation. Rec. Prog Horm Res, 1996． 51: 287-318．
72． Crawford RD. Poultry Genetics and Breeding ,1989．
73． D.C Millar C.I &Libby W.J.1991 strategies for conserving clinal, ecotypic, and disjunct
population diversity in widespread species. In Falk D.A & K.E. Holsinger (eds) genetic and conservation of rare plants New York: Oxford University Press.
74． Darwin C. 1896 The variation of animals and plants under domestication. 2nd editon. Appleton and company. New York.
75． Dunnington E.A etal. 1991 Poultry science 70:463-467; 1992 71:1251-1258．
76． Felsenstein, J. 1978． Cases in which parsimony or compatibility methods will be positively misleading. Systematic Zoology 27:401-410．
77． Ferguson CA, et al; The regulation of tyrosinase gene transcription.Pigment Cell Res 1997 Jun;10(3) :127-38
78． Fitch, W. M. 1971． Towards defining the course of evolution: Minimum change for a specific tree topology. Systematic Zoology 20:406-416．
79． Fitch, W. M. and E. Margoliash. 1967． Construction of phylogenetic trees. Science 155:279284．
80． Folkertsma,R.T.,Jeroen,N.A.M.,Der Voort,R.V.,De Groot, K.E.1996． Gene pool similarities of potato cyst nematode populations assessed by AFLP analysis.Molecular Plant-Microbe Interactions,9(l):47-54
81． Gantz I, Miwa H, Konda Y, Shimoto Y, Tashiro T, Watson S J, DelValle J, Yamada T, 1993b. Molecular clonging, expression, and gene localization of a fourth melanocortin receptor. J Biol Chem, 268:15174-15179
82． Gantz I, Tashiro T, Barcroft C, Konda Y, Shimoto Y, Miwa H, Glover T, Munzert G, T Yamada. Localization of the genes encoding the melanocortin-2 (adrenocor-ticotropic hormone) and melanocortin-3 receptors to chromosomes 18p11． 2 and 20q13． 2-q13． 3 by fluorescence in situ hybridization. Genomics , 1993c. 18:166-167
83． Gantz I, Y Konda, T Tashiro, Y Shimoto, H Miwa, G Munzert, S J Watson, J DelValle and T Yamada. Molecular clonging of a novel melanocortin receptor. J. Biol. Chem, 1993a.268:8246-8250
84． Georges. M., Nielsen. S., Mackinnon, M. Etal . 1994． Proc. 5th world congress. Genet. Appl. Livest. Prod. 21:81-85．
85． Grant V. 1991, the evolutionary process: A critical study of evolutionary theory (2nd). New York: Columbia University Press.
86． Grimont,F.,Grimont,P.A.D.1986． Ribosomal ribinucleic acid gene restrict-ion patterns as potential taxonomic tools.Ann.Inst.Pasteur/Microbi-01, 1378:165-175
87． Gu W, Tu Z, Kleyn P W, Kissebah A, Duprat L, Lee J, Chin W, Maruti S, Deng N, Fiaher S L, Franco L S, Burn P, Yagaloff K A, Nathan J, Heymsfield S, Albu J, Pi-Sunyer F X, Allison D E. Identification and functional analysis of novel human melanocortin-4 receptor variants. Diabetes, 1999． 48:635-639
88． Hans.H et al, 1995 Development of a genetic map of the chicken with markers of high utility, poultry science. 74: 1855-1874．
89． Harding R M, Healy E, Ray A J, Ellis N S, Flanagan N, Todd C, Dixon C, Sajantila A, Jackson I J, Birch-Machin M A, Rees J L. Evidence for variable selective pressures at MC1 R. Am. J Hum Genet, 2000． 66:1351-1361
90． Harrison P.G., et al., 1980 . animal mitochodrial DNA as a genetic marker in population and evolutionary biology. Tree. 4:6-11．
91． HillelJ etal, 1989 DNA fingerprints of poultry. Animal Genetics. 20:25-35．
92． Hillel.J etal, 1990． DNA fingerprints applied to gene introgression in breeding progress. Genetics, 124:783-789．
93． Hustad C M, Perry W L, Siracusa L D, Rasberry C, Cobb L, Cattanach B M, Kovatch R, Copeland N G, Jenkins N A. Molecular genetic characterisation of six recesive viable alleles of the mouse agouti locus. Genetics , 1995． 140:255-265
94． Huszer D, Lynch C A, Fairchild-Huntress V, Dunmore J H, Fang Q, Berkemeier L R, Gu W, Kesterson R A, Boston B A, Cone R D, Smith F J, Campfield L A, Burn P,-Lee F. Targeted disruption of the melanocortin-4 receptor results in obesity in mice. Cell, 1997． 88:131-141．
95． Iwahana,H.,Yoshimoto,K.,Itakura,M.1992． Dection of point mutations by SSCP of PCR-amplified DNA after endonuclease digestion. Biotechniques, 12:64-66
96． Jackson I J. Coat color-more to color than meets the eye. Curr. Biol ,1993a. 3:518-521
97． Jackson I J. Color coded switches. Nature, 1993b. 362:587-588
98． Jackson I J. Homologous pigmentation mutations in human, mouse and other model organisms. Hum Mol Genet, 1997． 6(10) :1613-1624
99． Joerg H, Fries H R, Meijerink E, Stranzinger G F. Red coat color in Holstein cattle is associated with a deletion in the MSHR gene. Mamm. Genome, 1996． 7:317-318
100． Jordan S A, Jackson I J. Melanocortin receptors and antagonists regulate pigmentation and body weight. BioEssays, 1998． 20:603-606．
101． Jukes, T. H. and C. R. Cantor. 1969． Evolution of protein molecules. In H. N. Munro, ed.,
Mammalian Protein Metabolism, pp. 21-132, Academic Press, New York.
102． Kijas J M H, Wales R, Tornsten A, Chardon P, Moller M, Andersson L. Melanocortin receptor 1 (MC1R) mutations and coat color in pigs. Genetics , 1998． 150:1177-1185．
103． Kimura, M. 1980． A simple method for estimating evolutionary rate of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution 16:111-120．
104． Klebig M L, Wilkinson J E, Geisler J G, Woychik R P. Ectopie expression of the agouti gene in transgenic mice causes obesity, features of type-II diabetes, and yellow fur. Proc Natl Acad Sci USA1995 . 92:4728-4732
105． Klungland H, Vage D I, Gomezraya L, Adalsteinsson S, Lien S. The role of melanocyte-stimulating hormone (MSH) receptor in bovine coat color determination. Mammalian Genome, 1995． 6:636-639．
106． Krude H, Biebermann H, Luck W, Horn R, Braban G, Gruters A. Several early-onset obesity, adrenal insufficiency and red hair pigmentation caused by POMC mutations in humans. Nature genetics, 1998． 19:155-157
107． Kucer G, Bortner D M, Rosenberg M P. Overexpression of an agouti cDNA in the skin of transgenic mice recapitulates dominant coat color of spontaneous mutants. Dev. Biol, 1996． 173:162-173
108． Kuhnlein U; Y.Zadworny; R.W. Dawe, etal. 1990 assessment of inbreeding by DNA fingerprinting: development of a calibration curve using defined strains of chichens. Genetics. 125:161-165．
109． Kuhnlein.U; Y.Dawe, D.Zadworny, & J.S Gawora, 1989 DNA fingerprinting: a tool for determining genetic distances between strains of poultry. Theor. Appl. Genet. 77:669-672．
110． Lewontin,R.C.1966． A molecular approach to the study of genetic hetrozygosity in natural populations. II .Amount of variation and degree of hetrozygisity in natural populations of Drosphila pseudoobscura. Genetics,54:595-609
111． Loftus, R. T. & hammond, K. 1994． Animal genetics. 25(suppl), 109．
112． Maddison, W. P. and D. R. Maddison. 1992． MacClade: Analysis of phylogeny and character evolution. Version 3． Sinauer Associates, Sunderland, Massachusetts.
113． Manne J, Argeson A C, Siracusa L D. Mechanisms for the pleiotropic effects of the agouti gene. Proc. Natl. Acad. Sci USA, 1995． 92:4721-4724
114． Marklund L, Moller M J, Sandberg K, Andersson L. A missense mutation in the gene for melanocyte-stimulating hormone receptor (MC1R) is associated with the chestnut coat color in horses. Mamm Genome Dec, 1996． 7(12) :895-899．
115． Moore,W.S.1995． Inferring phylogenies from mtDNA variation: Mitochondrial-gene trees versus nuclear-gene trees. Evolution, 49(4) :718-726
116． Moritz C. And D.M. Hillis. 1990 Molecular systematics: context and controversies molecular systematics. Sunderland: sinauer 1-11
117． Mountjoy K G, Robbins L S, Mortrud M T, Cone R D. The cloning of a family of genes that encode the melanocortin receptors. Science, 1992． 257:1248-1251．
118． Muroya S, Tanabe R, Nakajima I, Chikuni K; Molecular characteristics and site specific distribution of the pigment of the silky fowl. J Vet Med Sci 2000 Apr;62(4) :391-5
119． Nei, M., J. C. Stephens, and N. Saitou. 1985． Methods for computing the standard errors of branching points in an evolutionary tree and their application to molecular data from humans and apes. Molecular Biology and Evolution 2:66-85．
120． Nei, M., S. Kumar, K. Takahashi. 1998． The optimization principle in phylogenetic analysis tends to give incorrect topologies when the number of nucleotides or amino acids used is small. Proceedings of the National Academy of Sciences USA 95:12390-12397
121． Nei. M., 1987． Molecular evolutionary genetics. Colombia university press. New York. 122． Nozaki,a. etal;Energy dispersive X-ray fluorescence spectrometry of major tissures of silky fowl. J.Vet.Med.Sci. 1998,60:485-488．
123． Orita,M., Suzuki,T., Hayashi,K.1989． Rapid and sensitive detection of piont mutation and DNA polymorphisms using the polymerase chain reaction. Genomics, 5:874-879
124． Perry W L, Hustad C M, Swing D A, Jenkins N A, Copeland N G. A transgenic mouse assay for agouti protein activity. Genetics, 1995． 140:267-274
125． Prota G, Lamoreux M L, Muller J, Kobayashi T, Napolitano A, Vincensi M R, Sakai C, Hearing V J. Comparative analysis of melanins and melanosomes produced by various coat color mutants.Pigment Cell Res, 1995． 8:153-163
126． Qi,X.,Lindhout,P.1997． Development of AFLP markers in barely. Molecualr & General Genetics,254(3) :330-336
127． Ralph,D.,McCleuand,M.,Welsh,J.,Baranton,G.,Oerolat,P.1993． Leptospira species categorized by arbitrarily primed polymerase chain reaction (PCR) and by mapped
restriction polymorphisms in PCR-amplified rRNA genes.J. Bacteriol. 175:973-981
128． Rana B K, Hewett-Emmett D, Jin L, Chang B H-J, Sambuughin N, Lin M, Watkins S, Bamshad M, Jorde L B, Ramsay M, Jenkin T, L W-H. High polymorphism at the human melanocortin 1 receptor locus. Genetics, 1999． 151:1547-1557．
129． Robbins LS, Nadeau JH, Johnson KR, Kelly MA, Roselli-Rehfuss L, Baack E, Mountjoy KG, Cone RD. Pigmentation phenotypes of variant extension locus alleles result from point mutations that alter MSH receptor function. Cell ,1993． 72(6) :827-834．
130． Rolfs,A.,Schuller,I.,Finckh,U.,Weber-Rolfs,I.1992． Detection of single base changes using PCR,m: PCR: Clinical Diagnotics and Research,Speinger-Verlag, Berlin,Germany,pp. 149-167
131． Ron M., Band .M., Yanai, A. Etal. 1994． Animal. Genetics. 25: 259-264．
132． Rozas, J. and R. Rozas. 1999． DnaSP versions: an integrated program for molecular population genetic and molecular evolution analysis. Bioinformatics,!5:174-175．
133． Saitou, N. and M. Nei. 1987． The neighbor-joining method: A new method for reconstructing phylogenetic trees. Molecular Biology and Evolution 4:406-425．
134． Sankoff, D. and R. J. Cedergren. 1983． Simultaneous comparison of three or more sequences related by a tree. In D. Sankoff and J. B. Kruskal, eds., Tune Warps, String Edits, and Macromolecules: The Theory and Practice of Sequence Comparison, pp. 253-263． Addison-Wesley, Reading, Massachusetts.
135． Sarkar,G.,Yoon,H.S.,Sommer,S.1992b. Screening for mutations by RNA single-strand conformation polymorphism (rSSCP):comparison with DNA-SSCP.Nucl. Acids Res,20:871-878
136． Sarkar,G.,Yoon,H.S.Sommer,S.,1992a. Dideoxy fingerprinting(ddF: A rapid and efficient screen for the presence of mutations.)Genomics, 13:441-443
137． Sazanov A, Masabanda J, Ewald D, Takeuchi S, Tixier-Boichard M, Buitkamp J, Friews R. Evolutionarily conserved telomeric location of BBCl and MCIR on a microchromosome questions the identity of MCIR and a pigmentation locus on chromosome 1 in chicken. Chromosome Res Dec, 1998:6(8) :651-654
138． Sazanov A,et al. Evolutionarily conserved telomeric location of BBCl and MCIR on a microchromosome questions the identity of MCIR and a pigmentation locus on chromosome 1 in chicken.[J].Chromosome Res, 1998 ,6(8) :651-4．
139． Sazanov A,et al; Evolutionarily conserved telomeric location of BBCl and MCIR on a
microchromosome questions the identity of MC1R and a pigmentation locus on chromosome 1 in chicken.Chromosome Res 1998 Dec;6(8) :651-4
140． Shi P., Lu X M, Luo H R, Xiang-Yu J-G, Zhang Y P. Melanocortin-1 receptor gene variants in four Chinese ethnic populations. Cell Research, 2001． 11(1) :81-84．
141． Smith E., C.P.John, K.E.Nestor. 1996． Use of random amplified polymorphic DNA markers for genetic analysis of relatedness and diversity hi chickens and turkey. Poultry science. 75: 579-584．
142． Smyth JR Jr et al; Relationships between genes affecting melanin pigmentation and other traits in the fowl. Worlds Poult Sci J 1969 Jan-Mar;25(l):6-14
143． Smyth JR Jr. in poultry breeding and genetics(edited by Crawford R D), pp109-168,1990, elsevier, Amsterdam, Netherland.
144． Soller M, et al 1986 restriction fragment length polymorphisms in poultry breeding. Poultry Science. 65: 1476-1488．
145． Sudhir Kumar, Koichiro Tamura, Ingrid B. Jakobsen, and Masatoshi Nei (2001) MEGA2: Molecular Evolutionary Genetics Analysis software, Bioinformatics (submitted).
146． Suzuki I, Cone R C, Im S, Nordlund J, Abdel-Malek Z A. Binding of melanitropic hormones to the melanocortin receptor MC1R on human melano cytes stimulates proliferation and melanohenesis. Endocrinology, 1996． 137: 1627-1633
147． Swofford, D. L. 1998． PAUP: Phylogenetic Analysis Using Parsimony (and Other Methods) Sunderland, MA: Sinauer Associates.
148． Tajima, F. 1983． Evolutionary relationship of DNA sequences in finite populations. Genetics 105:437-460．
149． Tajima, F. 1993． Simple methods for testing molecular clock hypothesis. Genetics 135:599-607．
150． Tajima, F. and M. Nei. 1984． Estimation of evolutionary distance between nucleotide sequences. Molecular Biology and Evolution 1:269-285．
151． Takeuchi S, et al; A possible involvement of melanocortin 1-receptor in regulating feather color pigmentation in the chicken. Biochim Biophys Acta 1996 Aug 14;1308(2) :164-8
152． Takeuchi S, Suruki H, Yabuuchi M, Takahashi S. A possible involvement of melanocortin 1-receptor in regulating feather color pigmantation in the chicken. Biochim. Biophys. Acta Gene Struct Exp, 1996． 1308:164-168
153． Takeuchi S,et al; Melanocortin receptor genes hi the chicken-tissue distributions.Gen
Comp Endocrinol 1998 Nov;112(2) :220-31
154． Tamura, K. 1992． Estimation of the number of nucleotide substitutions when there are strong transition-transversion and G + C-content biases. Molecular Biology and Evolution 9:678-687．
155． Tamura, K. and M. Nei. 1993． Estimation of the number of nucleotide substitutions in the control region of mitochondrial DNA in humans and chimpanzees. Molecular Biology and Evolution 10:512-526．
156． Tan C P, McKee K K, Weinberg D H, MacNeil T, Palyha O C, Feighner S D, Hreniuk D L,Van Der Ploeg L H, MacNeil D J, Howard A D. Molecular analysis of a new splice variant of the human melanocortin-1 receptor. FEBS Lett, 1999． 451:137-141
157． Tanaka, T. and M. Nei. 1989． Positive Darwinian selection observed at the variable-region genes of immnoglobulins. Molecular Biology and Evolution 6:447-459．
158． Thomas,C.M.,Vos,P.,Zabeau,M.,Jones,D.A.,Norcott,K.A.,Chadwick,B.P.,Jone s,.D.G.1995． Identification of amplified fragment length polymorphism (AFLP)markers tightly linked to the tomato cf-9 gene for resistance to Cladosporium fulvum. Plant J.,8(5) :785-794
159． Vage D I, Lu D, Klungland H, Lien S, Adalsteinsson S, Cone R D. A non-epistatic interaction of agouti and extension in the fox, Vulpes vulpes. Nat. Genet, 1997． 15:311-315
160． Valverde P, Healy E, Jackson I, Rees J L, Thordy A J. Variants of the melanocyte stimulating hormone receptor gene are associated with melanoma. Hum Mol Genet, 1995． 11:328-330．
161． Vos,R,Hogers,R.M,Bleeker,M.,Reijians,M.,Lee,T.V.de,Hornes,M.,Frijte-rs,A.,Pot,J.,Peleman,J.,Kuiper,M.,Azbeau,M.1995． AFLP:a new technique for DNA fingerprinting.Nucleic Acids Research, 23(21) :4407-4414
162． Vrieling H, Duhl D M, Millar S E, Millar K A and Barsh G S. Differences in dorsal and ventral pigmentation result from regional expression of the mouse agouti gene. Proc. Natl. Acad. Sci USA, 1994． 91: 5667-5671
163． wilson, E.O.(1988) .Biodiversity, national academy press, Washington
164． Xia, X. 2000． Data analysis in molecular biology and evolution. Kluwer Academic Publishers. Boston/Dordrecht/London.
165． Yao Y-G, Lu X-M, Luo H-R, Li W-H, Zhang Y-P. Gene admixture in the Silk Road
region of China-evidence from mtDNA and melanocortin 1 receptor polymorphism. Genes & Genetics Systems., 2000． 75:173-178．
166． Yoshimoto. 1992． Role of P53 mutations in endocrine tumorigenesis: Mutation detection by polymerase chain reaction-single strand conformation polymorphism.Cancer Res.,52:5061-5064
167． Zhang X, C.R.Mcdniel, J.J.Giambrone, 1995． Random amplified polymorphic DNA comparisons among broiler lines selected for incidence of tibial dyschondroplasia. Poultry science. 74:1253-1258．
168． Zhu J, K.E.Nestor, I.Moritsu, 1996 relationship between band sharing levels of DNA fingerprints and inbreeding coefficients and estimation of true inbreeding in turkey lines. Poultry Science 75:25-28．
2．施立明，1990 遗传多样性及其保护层生命科学信息 2：158-164
3．史宪伟 1997 中国主要家鹅品种遗传多样性及其起源分化研究 博士学位论文
4．黄勇富 1996 中国部分地方猪种遗传多样性及起源分化研究 博士学位论文
5．吕雪梅等 1996 以RAPD标记分析广东四个鸡品种的亲缘关系
6．洪夏等 1997 河田鸡线粒体DNA的物理图谱。第八次全国家禽学术讨论会论文集。东北林大出版社 49-51
7．孙汉等 1997 江西地方鸡种六种血清酶多态性的研究 第八次全国家禽学术讨论会论文集东北林大出版社 82-85
8．吴常信 1999 对“畜禽遗传资源保护和利用”立项研究的建议 第九次全国家禽学术讨论会论文集 1-3
9．顾志良 2000 我国地方鸡种遗传多样性研究进展 中国家禽 7
10．陈幼春 1995 中国家畜多样性保护的意义 生物多样性 3(3)：143-146。
11．邱祥聘，陈锷，陈育新等 1988 中国家禽品种志 上海科技出版社 上海 3-16。
12．刘丑生，魏绍征 1995 DNA指纹的研究及其在畜禽遗传育种中的应用 甘肃畜牧兽医 (5)：27-29。
13．丁波 1994 DNA指纹技术及其在畜禽遗传育种中的应用 青海畜牧兽医杂志 (4)：42-44。
14．王金玉，龚允陈等 1995 鸡 DNA 指纹遗传研究进展及其在育种上的应用 中国家禽 (5)：29-31。
15．魏彩虹 1997 动物mtDNA遗传研究进展 甘肃畜牧兽医 (1)：23-25。
16．李建凡 1995 家禽分子生物学技术研究进展 国外畜牧科技 (5)：53-57。
17．孟安明 1993 DNA指纹技术在农业上的应用 生物工程进展 13(3)。
18．张沅 1991 生物技术在动物育种中应用的现状与前景 遗传 13(4)：39-42。
19．孟安明，齐顺章，周顺伍，宫桂芬 1993 家禽遗传图谱初报 遗传 15(2)：9-13。
20．曾福玪，李宁，王峰等 1995．鹌鹑品种(系)的随机扩增多态性DNA指纹分析 农业生物技术学报 3：61-64。
21．杨丽萍，1999 RAPD技术在家兔遗传分析中的应用。硕士学位论文。中国农科院。
22．王晓梅，宋文芹等，1998 用RAPD技术检测野生鲫鱼和四个金鱼代表品种的基因组DNA多态性。遗传20(5)：7-11。
23．李祥龙 1997 中国主要山羊品种遗传多样性及起源分化研究 四川农业大学博士论文。
24．张细权，吕学梅，杨玉华等，1998 用微卫星多态性和RAPD分析广东地方鸡种的群体遗传变异。遗传学报 25(2)：112—119。
25．周怀军，陈宽维等 1997 RAPD技术对地方鸡种群体遗传结构的分析 中国家禽 5(7)。
26．张学余，黄凡美等 1998 乌骨鸡种群遗传结构的RAPD分析 中国家禽 5(1)。
27．刘浚凡等 1998 矮小型黑羽乌骨鸡选育 中国家禽 1998 1(8)。
28．扬志勤 1997 乌骨鸡的品种与开发 (王红宁，朱庆 主编：养禽与禽病防治新进展) 四川科技出版社。42-45。
29．龚炎长，彭中镇等，1988。畜禽微卫星的应用。34(6)：51-52。
30．顾志良，王志跃，张军。1997。微卫星DNA标记及其在鸡的基因图谱构建中的应用。中国家禽。(6)：27-29。
31．刘浚凡等，1998。矮小型黑羽乌骨鸡选育 中国家禽 1(8)。
32．邱祥聘等，1999。丝羽乌骨鸡新品系的选育工作初报 第九次全国家禽学术讨论会论文集 7-10。
33．罗正雄等，1999。盐津乌骨鸡肉用性能的研究 第九次全国家禽学术讨论会论文集 86-91。
34．杜金平等，1999。金水乌骨鸡选育综合报告第九次全国家禽学术讨论会论文集 91-94。
35．金深逊等，1999。乌蒙黄羽乌骨鸡肉用性能研究 第九次全国家禽学术讨论会论文集 96-98。
36．金冬雁，金奇，侯云德，1996．核酸和蛋白质的化学合成与序列分析，北京：科学
37．出版社，49-70
38．张亚平．1996．从DNA序列到物种树，动物学研究，17(3)：247-252
39．张亚平，施立明．1992．动物线粒体DNA多态性的研究概况．动物学研究，13(3)：289—298
40．刘树俊，程光潮．1992．DNA指纹技术的应用及局限性．生物工程进展，12(5)：13-17
41．王斌，翁曼丽．1996．AFLP的原理及其应用．杂交水稻，(5)：27—30
42．包世增，家禽育种学，1994，29-43。北京：农业出版社
43．常洪，家畜遗传资源学纲要，1995，44-45。北京：中国农业出版社。
44．张细权，李加琪，杨关福，动物遗传标记，1997，7-8，中国农业大学出版社。
45．徐幸莲等，黑色素的研究进展，中国畜产与食品，1997，4(3)135-136。
46．江志洁等，黑色素形成机理的新概念及复合美白剂的应用，日用化学品科学，1998(4)：138-140。
47．徐幸莲等，乌骨鸡黑色素对延缓果蝇衰老的作用，南京农业大学学报，1999，22(2)105-108。
48．贾青等，泰和鸡黑色素的发生初探，河北农业大学学报．1998，21(3)．72-74。
49．吴常信主编，21世纪中国家禽业的研究与发展---第九次全国家禽学术讨论会论文集，1999．11，上海。
50．耿拓宇等，泰和乌鸡出雏后黑色素的分布与沉积，中国家禽，2000，22(7)：10-12。
51．朱子玉等，丝毛乌骨鸡和北京油鸡超氧化物歧化酶活性比较研究，中国家禽，2000(10) 6-7。
52. Abed, Y., Davinj-Regli,A.,Bollet,C.,De Micco,P.1995. Efficient discrimination of Mycobacherium tubemlosis strains by 16s spacer region based random amplified polymorphic DNA analysis. Clin Microbiol.33:1448-1420
53. Adachi S, Morii E, Kim D, Ogihara H, Jippo T, Ito A, Lee Y-M, Y Kitamura.Involvement of mi-transcription factor in expression of α-melanocyte-stimulating hormone receptor in cultured mast cells of mice. J. Immunology, 2000.164:855～860
54. Akahashi, K., and M. Nei. 2000. Efficiencies of fast algorithms of phylogenetic inference under the criteria of maximum parsimony, minimum evolution, and maximum likelihood when a large number of sequences are used. Molecular Biology and Evolution 17:1251-1258.
55. Akitoshi NOZAKI et al; The Surface color Measurement of Major tissues of Silky Fowls and White Leghorns, J.Vet. Med. Sci. 1998,60:489-493
56. Ashwell, M.S. Rexroad, C. E. Miller. J.R.H, etal. 1992. Animal genetics. 23:133-142.
57. Ayala F.J & J.W, Valentine 1991. Evolving: the theory and progress of organic evolution: menlo park: Benjamin-cummings.
58. B. Van Wijk. Wildlife management principles. What is biodiversity 103-104
59. Ballou, J.D; M.Gilpin, T.J, foise, eds. Population management for survival and recovery,analytical methods and strategies in small population conservation. Columbia University
Press, New York, 1995．
60． Barnes,Nick: conflicts over biodiversty. In environmental policy in an international context part SOU, 1996．
61． Boston B A, Cone R D.Characterization of melanocortin receptor subtype expression in murine adipose tissues and in the 3T3-L1 cell line. Endocrinology , 1996． 137:2043-2050
62． Brown W.M.,et al. 1979． Rapid evolution of animal mitochondrial DNA. Proc. Natl. Acad. Sci. USA. 76:1967-1971．
63． Brown,W.M. 1983． Evolution of animal mtDNA. In:Nei,M.,and Koehn,R.K. (eds.). Evolution of Gene and Proteins,sinauer,sunderland,62-68
64． Bultman S P, Michaud E J, Woychik R P. Molecular characterisation of the mouse agouti locus. Cell ,1992． 71:1-20
65． Chakraborty A, Slominski A, Ermak G, Hwang J, Pawelek J. Ultraviolet B and melanocyte-stimulating hormone (MSH) stimulate mRNA production for alpha MSH receptors and proopiomelanocortin-derived peptides in mouse melanoma cells and transformed keratinocytes. J Invest Dermatol, 1995． 105:655-659
66． Cheng H. Etal. 1994． Poultry science. 73: 539-546; 74: 1855-1874．
67． Chhajlani V, Muceniece R, Wiaberg J E S. Moleculat cloning of a novel human menocortin-1 receptor. Biochem Biophys Res Commun,1993． 195:866-873
68． Chhajlani V, Wikberg J E S. Molecular cloning and expression of the human melanocyte stimulating hormone receptor cDNA. FEBS Lett, 1992． 309: 417-420．
69． Chhajlani V, Xu X, Blauw J, Sudarshi S. Identification of ligand binding residues in extracellular loops of the melanocortin 1 receptor. Biochem Biophys Res Commun Feb, 1996． 15;219(2) :521-525
70． Chluba-de Tapia J, Bagutti C, Cotti R, Eberle A N. Induction of constitutive melanogenesis in amelanotic mouse melanoma cells by transfection of the human melanocortin-1 receptor gene. J Cell Sci, 1996． 109 (Pt 8) :2023-2030．
71． Cone R D, Lu D, Koppula S, Vage D I, Klungland H, Boston B, Chen W, Orth D N, Pouton C, Kesterson R A. The melanocortin receptors: agonists, antagonists, and the hormonal control of pigmentation. Rec. Prog Horm Res, 1996． 51: 287-318．
72． Crawford RD. Poultry Genetics and Breeding ,1989．
73． D.C Millar C.I &Libby W.J.1991 strategies for conserving clinal, ecotypic, and disjunct
population diversity in widespread species. In Falk D.A & K.E. Holsinger (eds) genetic and conservation of rare plants New York: Oxford University Press.
74． Darwin C. 1896 The variation of animals and plants under domestication. 2nd editon. Appleton and company. New York.
75． Dunnington E.A etal. 1991 Poultry science 70:463-467; 1992 71:1251-1258．
76． Felsenstein, J. 1978． Cases in which parsimony or compatibility methods will be positively misleading. Systematic Zoology 27:401-410．
77． Ferguson CA, et al; The regulation of tyrosinase gene transcription.Pigment Cell Res 1997 Jun;10(3) :127-38
78． Fitch, W. M. 1971． Towards defining the course of evolution: Minimum change for a specific tree topology. Systematic Zoology 20:406-416．
79． Fitch, W. M. and E. Margoliash. 1967． Construction of phylogenetic trees. Science 155:279284．
80． Folkertsma,R.T.,Jeroen,N.A.M.,Der Voort,R.V.,De Groot, K.E.1996． Gene pool similarities of potato cyst nematode populations assessed by AFLP analysis.Molecular Plant-Microbe Interactions,9(l):47-54
81． Gantz I, Miwa H, Konda Y, Shimoto Y, Tashiro T, Watson S J, DelValle J, Yamada T, 1993b. Molecular clonging, expression, and gene localization of a fourth melanocortin receptor. J Biol Chem, 268:15174-15179
82． Gantz I, Tashiro T, Barcroft C, Konda Y, Shimoto Y, Miwa H, Glover T, Munzert G, T Yamada. Localization of the genes encoding the melanocortin-2 (adrenocor-ticotropic hormone) and melanocortin-3 receptors to chromosomes 18p11． 2 and 20q13． 2-q13． 3 by fluorescence in situ hybridization. Genomics , 1993c. 18:166-167
83． Gantz I, Y Konda, T Tashiro, Y Shimoto, H Miwa, G Munzert, S J Watson, J DelValle and T Yamada. Molecular clonging of a novel melanocortin receptor. J. Biol. Chem, 1993a.268:8246-8250
84． Georges. M., Nielsen. S., Mackinnon, M. Etal . 1994． Proc. 5th world congress. Genet. Appl. Livest. Prod. 21:81-85．
85． Grant V. 1991, the evolutionary process: A critical study of evolutionary theory (2nd). New York: Columbia University Press.
86． Grimont,F.,Grimont,P.A.D.1986． Ribosomal ribinucleic acid gene restrict-ion patterns as potential taxonomic tools.Ann.Inst.Pasteur/Microbi-01, 1378:165-175
87． Gu W, Tu Z, Kleyn P W, Kissebah A, Duprat L, Lee J, Chin W, Maruti S, Deng N, Fiaher S L, Franco L S, Burn P, Yagaloff K A, Nathan J, Heymsfield S, Albu J, Pi-Sunyer F X, Allison D E. Identification and functional analysis of novel human melanocortin-4 receptor variants. Diabetes, 1999． 48:635-639
88． Hans.H et al, 1995 Development of a genetic map of the chicken with markers of high utility, poultry science. 74: 1855-1874．
89． Harding R M, Healy E, Ray A J, Ellis N S, Flanagan N, Todd C, Dixon C, Sajantila A, Jackson I J, Birch-Machin M A, Rees J L. Evidence for variable selective pressures at MC1 R. Am. J Hum Genet, 2000． 66:1351-1361
90． Harrison P.G., et al., 1980 . animal mitochodrial DNA as a genetic marker in population and evolutionary biology. Tree. 4:6-11．
91． HillelJ etal, 1989 DNA fingerprints of poultry. Animal Genetics. 20:25-35．
92． Hillel.J etal, 1990． DNA fingerprints applied to gene introgression in breeding progress. Genetics, 124:783-789．
93． Hustad C M, Perry W L, Siracusa L D, Rasberry C, Cobb L, Cattanach B M, Kovatch R, Copeland N G, Jenkins N A. Molecular genetic characterisation of six recesive viable alleles of the mouse agouti locus. Genetics , 1995． 140:255-265
94． Huszer D, Lynch C A, Fairchild-Huntress V, Dunmore J H, Fang Q, Berkemeier L R, Gu W, Kesterson R A, Boston B A, Cone R D, Smith F J, Campfield L A, Burn P,-Lee F. Targeted disruption of the melanocortin-4 receptor results in obesity in mice. Cell, 1997． 88:131-141．
95． Iwahana,H.,Yoshimoto,K.,Itakura,M.1992． Dection of point mutations by SSCP of PCR-amplified DNA after endonuclease digestion. Biotechniques, 12:64-66
96． Jackson I J. Coat color-more to color than meets the eye. Curr. Biol ,1993a. 3:518-521
97． Jackson I J. Color coded switches. Nature, 1993b. 362:587-588
98． Jackson I J. Homologous pigmentation mutations in human, mouse and other model organisms. Hum Mol Genet, 1997． 6(10) :1613-1624
99． Joerg H, Fries H R, Meijerink E, Stranzinger G F. Red coat color in Holstein cattle is associated with a deletion in the MSHR gene. Mamm. Genome, 1996． 7:317-318
100． Jordan S A, Jackson I J. Melanocortin receptors and antagonists regulate pigmentation and body weight. BioEssays, 1998． 20:603-606．
101． Jukes, T. H. and C. R. Cantor. 1969． Evolution of protein molecules. In H. N. Munro, ed.,
Mammalian Protein Metabolism, pp. 21-132, Academic Press, New York.
102． Kijas J M H, Wales R, Tornsten A, Chardon P, Moller M, Andersson L. Melanocortin receptor 1 (MC1R) mutations and coat color in pigs. Genetics , 1998． 150:1177-1185．
103． Kimura, M. 1980． A simple method for estimating evolutionary rate of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution 16:111-120．
104． Klebig M L, Wilkinson J E, Geisler J G, Woychik R P. Ectopie expression of the agouti gene in transgenic mice causes obesity, features of type-II diabetes, and yellow fur. Proc Natl Acad Sci USA1995 . 92:4728-4732
105． Klungland H, Vage D I, Gomezraya L, Adalsteinsson S, Lien S. The role of melanocyte-stimulating hormone (MSH) receptor in bovine coat color determination. Mammalian Genome, 1995． 6:636-639．
106． Krude H, Biebermann H, Luck W, Horn R, Braban G, Gruters A. Several early-onset obesity, adrenal insufficiency and red hair pigmentation caused by POMC mutations in humans. Nature genetics, 1998． 19:155-157
107． Kucer G, Bortner D M, Rosenberg M P. Overexpression of an agouti cDNA in the skin of transgenic mice recapitulates dominant coat color of spontaneous mutants. Dev. Biol, 1996． 173:162-173
108． Kuhnlein U; Y.Zadworny; R.W. Dawe, etal. 1990 assessment of inbreeding by DNA fingerprinting: development of a calibration curve using defined strains of chichens. Genetics. 125:161-165．
109． Kuhnlein.U; Y.Dawe, D.Zadworny, & J.S Gawora, 1989 DNA fingerprinting: a tool for determining genetic distances between strains of poultry. Theor. Appl. Genet. 77:669-672．
110． Lewontin,R.C.1966． A molecular approach to the study of genetic hetrozygosity in natural populations. II .Amount of variation and degree of hetrozygisity in natural populations of Drosphila pseudoobscura. Genetics,54:595-609
111． Loftus, R. T. & hammond, K. 1994． Animal genetics. 25(suppl), 109．
112． Maddison, W. P. and D. R. Maddison. 1992． MacClade: Analysis of phylogeny and character evolution. Version 3． Sinauer Associates, Sunderland, Massachusetts.
113． Manne J, Argeson A C, Siracusa L D. Mechanisms for the pleiotropic effects of the agouti gene. Proc. Natl. Acad. Sci USA, 1995． 92:4721-4724
114． Marklund L, Moller M J, Sandberg K, Andersson L. A missense mutation in the gene for melanocyte-stimulating hormone receptor (MC1R) is associated with the chestnut coat color in horses. Mamm Genome Dec, 1996． 7(12) :895-899．
115． Moore,W.S.1995． Inferring phylogenies from mtDNA variation: Mitochondrial-gene trees versus nuclear-gene trees. Evolution, 49(4) :718-726
116． Moritz C. And D.M. Hillis. 1990 Molecular systematics: context and controversies molecular systematics. Sunderland: sinauer 1-11
117． Mountjoy K G, Robbins L S, Mortrud M T, Cone R D. The cloning of a family of genes that encode the melanocortin receptors. Science, 1992． 257:1248-1251．
118． Muroya S, Tanabe R, Nakajima I, Chikuni K; Molecular characteristics and site specific distribution of the pigment of the silky fowl. J Vet Med Sci 2000 Apr;62(4) :391-5
119． Nei, M., J. C. Stephens, and N. Saitou. 1985． Methods for computing the standard errors of branching points in an evolutionary tree and their application to molecular data from humans and apes. Molecular Biology and Evolution 2:66-85．
120． Nei, M., S. Kumar, K. Takahashi. 1998． The optimization principle in phylogenetic analysis tends to give incorrect topologies when the number of nucleotides or amino acids used is small. Proceedings of the National Academy of Sciences USA 95:12390-12397
121． Nei. M., 1987． Molecular evolutionary genetics. Colombia university press. New York. 122． Nozaki,a. etal;Energy dispersive X-ray fluorescence spectrometry of major tissures of silky fowl. J.Vet.Med.Sci. 1998,60:485-488．
123． Orita,M., Suzuki,T., Hayashi,K.1989． Rapid and sensitive detection of piont mutation and DNA polymorphisms using the polymerase chain reaction. Genomics, 5:874-879
124． Perry W L, Hustad C M, Swing D A, Jenkins N A, Copeland N G. A transgenic mouse assay for agouti protein activity. Genetics, 1995． 140:267-274
125． Prota G, Lamoreux M L, Muller J, Kobayashi T, Napolitano A, Vincensi M R, Sakai C, Hearing V J. Comparative analysis of melanins and melanosomes produced by various coat color mutants.Pigment Cell Res, 1995． 8:153-163
126． Qi,X.,Lindhout,P.1997． Development of AFLP markers in barely. Molecualr & General Genetics,254(3) :330-336
127． Ralph,D.,McCleuand,M.,Welsh,J.,Baranton,G.,Oerolat,P.1993． Leptospira species categorized by arbitrarily primed polymerase chain reaction (PCR) and by mapped
restriction polymorphisms in PCR-amplified rRNA genes.J. Bacteriol. 175:973-981
128． Rana B K, Hewett-Emmett D, Jin L, Chang B H-J, Sambuughin N, Lin M, Watkins S, Bamshad M, Jorde L B, Ramsay M, Jenkin T, L W-H. High polymorphism at the human melanocortin 1 receptor locus. Genetics, 1999． 151:1547-1557．
129． Robbins LS, Nadeau JH, Johnson KR, Kelly MA, Roselli-Rehfuss L, Baack E, Mountjoy KG, Cone RD. Pigmentation phenotypes of variant extension locus alleles result from point mutations that alter MSH receptor function. Cell ,1993． 72(6) :827-834．
130． Rolfs,A.,Schuller,I.,Finckh,U.,Weber-Rolfs,I.1992． Detection of single base changes using PCR,m: PCR: Clinical Diagnotics and Research,Speinger-Verlag, Berlin,Germany,pp. 149-167
131． Ron M., Band .M., Yanai, A. Etal. 1994． Animal. Genetics. 25: 259-264．
132． Rozas, J. and R. Rozas. 1999． DnaSP versions: an integrated program for molecular population genetic and molecular evolution analysis. Bioinformatics,!5:174-175．
133． Saitou, N. and M. Nei. 1987． The neighbor-joining method: A new method for reconstructing phylogenetic trees. Molecular Biology and Evolution 4:406-425．
134． Sankoff, D. and R. J. Cedergren. 1983． Simultaneous comparison of three or more sequences related by a tree. In D. Sankoff and J. B. Kruskal, eds., Tune Warps, String Edits, and Macromolecules: The Theory and Practice of Sequence Comparison, pp. 253-263． Addison-Wesley, Reading, Massachusetts.
135． Sarkar,G.,Yoon,H.S.,Sommer,S.1992b. Screening for mutations by RNA single-strand conformation polymorphism (rSSCP):comparison with DNA-SSCP.Nucl. Acids Res,20:871-878
136． Sarkar,G.,Yoon,H.S.Sommer,S.,1992a. Dideoxy fingerprinting(ddF: A rapid and efficient screen for the presence of mutations.)Genomics, 13:441-443
137． Sazanov A, Masabanda J, Ewald D, Takeuchi S, Tixier-Boichard M, Buitkamp J, Friews R. Evolutionarily conserved telomeric location of BBCl and MCIR on a microchromosome questions the identity of MCIR and a pigmentation locus on chromosome 1 in chicken. Chromosome Res Dec, 1998:6(8) :651-654
138． Sazanov A,et al. Evolutionarily conserved telomeric location of BBCl and MCIR on a microchromosome questions the identity of MCIR and a pigmentation locus on chromosome 1 in chicken.[J].Chromosome Res, 1998 ,6(8) :651-4．
139． Sazanov A,et al; Evolutionarily conserved telomeric location of BBCl and MCIR on a
microchromosome questions the identity of MC1R and a pigmentation locus on chromosome 1 in chicken.Chromosome Res 1998 Dec;6(8) :651-4
140． Shi P., Lu X M, Luo H R, Xiang-Yu J-G, Zhang Y P. Melanocortin-1 receptor gene variants in four Chinese ethnic populations. Cell Research, 2001． 11(1) :81-84．
141． Smith E., C.P.John, K.E.Nestor. 1996． Use of random amplified polymorphic DNA markers for genetic analysis of relatedness and diversity hi chickens and turkey. Poultry science. 75: 579-584．
142． Smyth JR Jr et al; Relationships between genes affecting melanin pigmentation and other traits in the fowl. Worlds Poult Sci J 1969 Jan-Mar;25(l):6-14
143． Smyth JR Jr. in poultry breeding and genetics(edited by Crawford R D), pp109-168,1990, elsevier, Amsterdam, Netherland.
144． Soller M, et al 1986 restriction fragment length polymorphisms in poultry breeding. Poultry Science. 65: 1476-1488．
145． Sudhir Kumar, Koichiro Tamura, Ingrid B. Jakobsen, and Masatoshi Nei (2001) MEGA2: Molecular Evolutionary Genetics Analysis software, Bioinformatics (submitted).
146． Suzuki I, Cone R C, Im S, Nordlund J, Abdel-Malek Z A. Binding of melanitropic hormones to the melanocortin receptor MC1R on human melano cytes stimulates proliferation and melanohenesis. Endocrinology, 1996． 137: 1627-1633
147． Swofford, D. L. 1998． PAUP: Phylogenetic Analysis Using Parsimony (and Other Methods) Sunderland, MA: Sinauer Associates.
148． Tajima, F. 1983． Evolutionary relationship of DNA sequences in finite populations. Genetics 105:437-460．
149． Tajima, F. 1993． Simple methods for testing molecular clock hypothesis. Genetics 135:599-607．
150． Tajima, F. and M. Nei. 1984． Estimation of evolutionary distance between nucleotide sequences. Molecular Biology and Evolution 1:269-285．
151． Takeuchi S, et al; A possible involvement of melanocortin 1-receptor in regulating feather color pigmentation in the chicken. Biochim Biophys Acta 1996 Aug 14;1308(2) :164-8
152． Takeuchi S, Suruki H, Yabuuchi M, Takahashi S. A possible involvement of melanocortin 1-receptor in regulating feather color pigmantation in the chicken. Biochim. Biophys. Acta Gene Struct Exp, 1996． 1308:164-168
153． Takeuchi S,et al; Melanocortin receptor genes hi the chicken-tissue distributions.Gen
Comp Endocrinol 1998 Nov;112(2) :220-31
154． Tamura, K. 1992． Estimation of the number of nucleotide substitutions when there are strong transition-transversion and G + C-content biases. Molecular Biology and Evolution 9:678-687．
155． Tamura, K. and M. Nei. 1993． Estimation of the number of nucleotide substitutions in the control region of mitochondrial DNA in humans and chimpanzees. Molecular Biology and Evolution 10:512-526．
156． Tan C P, McKee K K, Weinberg D H, MacNeil T, Palyha O C, Feighner S D, Hreniuk D L,Van Der Ploeg L H, MacNeil D J, Howard A D. Molecular analysis of a new splice variant of the human melanocortin-1 receptor. FEBS Lett, 1999． 451:137-141
157． Tanaka, T. and M. Nei. 1989． Positive Darwinian selection observed at the variable-region genes of immnoglobulins. Molecular Biology and Evolution 6:447-459．
158． Thomas,C.M.,Vos,P.,Zabeau,M.,Jones,D.A.,Norcott,K.A.,Chadwick,B.P.,Jone s,.D.G.1995． Identification of amplified fragment length polymorphism (AFLP)markers tightly linked to the tomato cf-9 gene for resistance to Cladosporium fulvum. Plant J.,8(5) :785-794
159． Vage D I, Lu D, Klungland H, Lien S, Adalsteinsson S, Cone R D. A non-epistatic interaction of agouti and extension in the fox, Vulpes vulpes. Nat. Genet, 1997． 15:311-315
160． Valverde P, Healy E, Jackson I, Rees J L, Thordy A J. Variants of the melanocyte stimulating hormone receptor gene are associated with melanoma. Hum Mol Genet, 1995． 11:328-330．
161． Vos,R,Hogers,R.M,Bleeker,M.,Reijians,M.,Lee,T.V.de,Hornes,M.,Frijte-rs,A.,Pot,J.,Peleman,J.,Kuiper,M.,Azbeau,M.1995． AFLP:a new technique for DNA fingerprinting.Nucleic Acids Research, 23(21) :4407-4414
162． Vrieling H, Duhl D M, Millar S E, Millar K A and Barsh G S. Differences in dorsal and ventral pigmentation result from regional expression of the mouse agouti gene. Proc. Natl. Acad. Sci USA, 1994． 91: 5667-5671
163． wilson, E.O.(1988) .Biodiversity, national academy press, Washington
164． Xia, X. 2000． Data analysis in molecular biology and evolution. Kluwer Academic Publishers. Boston/Dordrecht/London.
165． Yao Y-G, Lu X-M, Luo H-R, Li W-H, Zhang Y-P. Gene admixture in the Silk Road
region of China-evidence from mtDNA and melanocortin 1 receptor polymorphism. Genes & Genetics Systems., 2000． 75:173-178．
166． Yoshimoto. 1992． Role of P53 mutations in endocrine tumorigenesis: Mutation detection by polymerase chain reaction-single strand conformation polymorphism.Cancer Res.,52:5061-5064
167． Zhang X, C.R.Mcdniel, J.J.Giambrone, 1995． Random amplified polymorphic DNA comparisons among broiler lines selected for incidence of tibial dyschondroplasia. Poultry science. 74:1253-1258．
168． Zhu J, K.E.Nestor, I.Moritsu, 1996 relationship between band sharing levels of DNA fingerprints and inbreeding coefficients and estimation of true inbreeding in turkey lines. Poultry Science 75:25-28．