滩羊3号染色体遗传连锁图谱构建及羊毛性状QTL探测
|
摘要
本研究利用滩羊与小尾寒羊杂交的8个父系半同胞参考家系共576个个体,分析了绵羊基因组3号染色体上13个微卫星标记的等位基因频率、多态信息含量和群体杂合度,构建了3号染色体遗传连锁框架图谱,采用差异群体设计对影响滩羊10个表型性状的数量性状位点进行了检测研究,表型性状包括:初生重、二毛期活重、初生毛长、初生弯曲数、肩部弯曲数、股部弯曲数、毛股花穗类型、毛股紧实度、花案清晰度、弯曲比例。
所选的微卫星标记在参考群体中平均检测到11个(5~20个)等位基因,平均多态信息含量为0.5778 ( 0.2042~0.8320 ),群体杂合度平均值为0.5951(0.2094~0.8459)。13个标记中有9个呈现高度多态性,2个呈现中度多态性,2个呈现低度多态性,为该群体分子水平上的研究提供了可靠的依据。
构建的滩羊第3号染色体遗传连锁框架图总长度为466.7cM,标记平均间距为38.9cM,其中标记ILSTS049与CSRD264间距最大,为65.6cM,而BM2830和MCMA13间距最小,为4.9cM。图谱中13个标记的顺序与美国USDA-MARC和英国罗斯林研究所公布的SM4.7最新绵羊连锁图相同,但大部分标记间距大于后两者。
对参考群体中161个后代的10个重要性状的表型资料进行了分析。相关分析的结果表明初生重与初生毛长、初生弯曲数、二毛期活重、股部弯曲数间呈极显著正相关;初生毛长与初生弯曲数、花案清晰度间呈极显著正相关,与二毛期活重、肩部弯曲数、股部弯曲数呈显著正相关;初生弯曲数与花案清晰度、肩部弯曲数、股部弯曲数、弯曲比例呈极显著正相关,与毛股紧实度呈显著正相关;花案清晰度与弯曲比例呈极显著正相关;花穗类型与肩部弯曲数呈极显著正相关,与股部弯曲数呈显著正相关;毛股紧实度与肩部弯曲数、股部弯曲数、弯曲比例呈极显著正相关;肩部弯曲数与股部弯曲数、弯曲比例呈极显著正相关;股部弯曲数与弯曲比例呈显著正相关。方差分析结果表明家系(即公羊效应)对花案清晰度和毛股紧实度的影响不显著、对花穗类型影响显著之外对其余7个性状影响均极显著。最小二乘分析结果表明10个性状中除花案清晰度、毛股紧实度和花穗类型在各家系间差异不显著之外,其它各个性状在家系间存在不同程度的差异。
10个表型性状的QTL检测分析结果表明:QTL Express软件检测到了两个QTL区域,花案清晰度的QTL和花穗类型的QTL,但二者均未达到显著水平,未检测到明显的有关其他性状的QTL。
A referencing flock comprising eight paternal half-sib families totaling 576 individuals from population produced Tan sheep crossing with Small Tailed Han sheep was utilized to build a framework linkage map of the 3rd chromosome. Thirteen microsatellites were selected and genotyped; then allele frequency, polymorphism information content and heterozygosity of these markers were computed within the experimental group. Phenotypic data of ten traits, namely birth weight, weight at the time of kivskin, length of wool at birth, numbers of plait crimps of wool at shoulder and thigh, definition of plait crimps of wool and so on.
The average number of alleles, average PIC and average heterozygosis of micro satellites were 11(from 5 to 20), 0.5778(from 0.2042 to 0.8320) and 0.5951(0.2094~ 0.8459) respectively. And nine markers were high polymorphic, two were middle and two were low, those could substantially guarantee the reliability of the result of the research.
The sex average linkage map of the 3rd chromosome of Tan sheep was constructed by Crimap(2.4). The complete length of the map was 466.7cM, which had an average marker interval of 38.9cM as well as the maximum and minimum interval of 65.6cM and 4.9cM. Besides that, the sequence of loci was coincident with the USDA-MARC and SM4.7(ARKdb) ovine map, however most marker intervals were larger than the latter two.
Phenotypes of ten traits were handled statistically by SAS6.12 in the 161 progenies. The“corr”consequence revealed the significant correlation between several traits, weight and wool length, wool length and numbers of plait crimps of wool, numbers of plait crimps of wool and ratio of flexural on wool, and so on. Meanwhile, the“GLM”result demonstrated sire factor has robust influence on birth weight, length of wool at birth, numbers of plait crimps of wool at shoulder and thigh, ratio of flexural on wool.
The QTL mapping was accomplished through interval mapping. A software named QTL Express was applied to analyze simultaneously. A putative QTL affecting definition of plait crimps of wool and another putative QTL affecting type of flower of wool were detected, but both of them have no significant.
所选的微卫星标记在参考群体中平均检测到11个(5~20个)等位基因,平均多态信息含量为0.5778 ( 0.2042~0.8320 ),群体杂合度平均值为0.5951(0.2094~0.8459)。13个标记中有9个呈现高度多态性,2个呈现中度多态性,2个呈现低度多态性,为该群体分子水平上的研究提供了可靠的依据。
构建的滩羊第3号染色体遗传连锁框架图总长度为466.7cM,标记平均间距为38.9cM,其中标记ILSTS049与CSRD264间距最大,为65.6cM,而BM2830和MCMA13间距最小,为4.9cM。图谱中13个标记的顺序与美国USDA-MARC和英国罗斯林研究所公布的SM4.7最新绵羊连锁图相同,但大部分标记间距大于后两者。
对参考群体中161个后代的10个重要性状的表型资料进行了分析。相关分析的结果表明初生重与初生毛长、初生弯曲数、二毛期活重、股部弯曲数间呈极显著正相关;初生毛长与初生弯曲数、花案清晰度间呈极显著正相关,与二毛期活重、肩部弯曲数、股部弯曲数呈显著正相关;初生弯曲数与花案清晰度、肩部弯曲数、股部弯曲数、弯曲比例呈极显著正相关,与毛股紧实度呈显著正相关;花案清晰度与弯曲比例呈极显著正相关;花穗类型与肩部弯曲数呈极显著正相关,与股部弯曲数呈显著正相关;毛股紧实度与肩部弯曲数、股部弯曲数、弯曲比例呈极显著正相关;肩部弯曲数与股部弯曲数、弯曲比例呈极显著正相关;股部弯曲数与弯曲比例呈显著正相关。方差分析结果表明家系(即公羊效应)对花案清晰度和毛股紧实度的影响不显著、对花穗类型影响显著之外对其余7个性状影响均极显著。最小二乘分析结果表明10个性状中除花案清晰度、毛股紧实度和花穗类型在各家系间差异不显著之外,其它各个性状在家系间存在不同程度的差异。
10个表型性状的QTL检测分析结果表明:QTL Express软件检测到了两个QTL区域,花案清晰度的QTL和花穗类型的QTL,但二者均未达到显著水平,未检测到明显的有关其他性状的QTL。
A referencing flock comprising eight paternal half-sib families totaling 576 individuals from population produced Tan sheep crossing with Small Tailed Han sheep was utilized to build a framework linkage map of the 3rd chromosome. Thirteen microsatellites were selected and genotyped; then allele frequency, polymorphism information content and heterozygosity of these markers were computed within the experimental group. Phenotypic data of ten traits, namely birth weight, weight at the time of kivskin, length of wool at birth, numbers of plait crimps of wool at shoulder and thigh, definition of plait crimps of wool and so on.
The average number of alleles, average PIC and average heterozygosis of micro satellites were 11(from 5 to 20), 0.5778(from 0.2042 to 0.8320) and 0.5951(0.2094~ 0.8459) respectively. And nine markers were high polymorphic, two were middle and two were low, those could substantially guarantee the reliability of the result of the research.
The sex average linkage map of the 3rd chromosome of Tan sheep was constructed by Crimap(2.4). The complete length of the map was 466.7cM, which had an average marker interval of 38.9cM as well as the maximum and minimum interval of 65.6cM and 4.9cM. Besides that, the sequence of loci was coincident with the USDA-MARC and SM4.7(ARKdb) ovine map, however most marker intervals were larger than the latter two.
Phenotypes of ten traits were handled statistically by SAS6.12 in the 161 progenies. The“corr”consequence revealed the significant correlation between several traits, weight and wool length, wool length and numbers of plait crimps of wool, numbers of plait crimps of wool and ratio of flexural on wool, and so on. Meanwhile, the“GLM”result demonstrated sire factor has robust influence on birth weight, length of wool at birth, numbers of plait crimps of wool at shoulder and thigh, ratio of flexural on wool.
The QTL mapping was accomplished through interval mapping. A software named QTL Express was applied to analyze simultaneously. A putative QTL affecting definition of plait crimps of wool and another putative QTL affecting type of flower of wool were detected, but both of them have no significant.
引文
[1] Nilsson-Ehle, H. Kreuzunguntersuchungen an Hafer und Wizen. Lunds Universit. Arsskr. N. F. 1909,5(2): 1-122.
[2]余雳.猪2、4号Chr.连锁图构建和肉质性状QTL的鉴定[D].武汉:华中农业大学,2001,5.
[3]狄冉,马月辉.微卫星标记及其在绵、山羊中的研究进展[J].家畜生态学报.2006,27(1):96-100
[4]周明亮.凉山半细毛羊第3号染色体遗传连锁图谱构建及体质性状QTL定位的研究[D].雅安,四川农业大学, 2005.5.
[5]吴登俊,马丁·菲尔斯特.主要家畜种基因组连锁图现状和展望[J].国外畜牧科技.1999, 26(6):21-25.
[6]周明亮,吴登俊,曾云琦.绵羊遗传连锁图的研究现状[J].贵州农业科学.2005,33(3):96-97.
[7]涂友仁等.中国羊品种志.上海科学技术出版社[M].1989.4.:45-47.
[8] Rust C C.Hormonal Control of pelage cycles in the mink[J].J Mammal,1965,(46):549-565.
[9] Rose J.Induction of winter fur growth in mink with melatonin[J]. Anim Sci,1984,58(1):57.
[10] Rose J.The effects of photoperiod melatonin on serum prolaction levels of mink during the autumn moult.[J]. Pineal Research,1985,(2):13.
[11]林仲凡.毛皮动物埋植褪黑激素的研究进展[J].动物学杂志,1988.23(5):45-47.
[12]柳建昌,尹协镇,方天祺.褪黑素对中国绒山羊在非生绒期促绒生长与绒产量的影响[J].动物学杂志,1998,33(3):8-12.
[13]柳建昌.含褪黑素微型胶囊促使银狐毛皮提早成熟的效应[J].动物学杂志,1995,30(1):48-51.
[14]盛雪.褪黑素植入物在狐狸毛皮成熟方面的应用[J].山东畜牧兽医,1997,(2):51-52.
[15]谢宝胜,杨占兴,马武龙,等.褪黑素对门源牦牛春季促绒生长及绒产量的影响[J].中国草食动物,1999,1(6):12-13.
[16] Wynn PC, et al. Randomised controlled trial of intravenous antibiotic treatme.[J].Aust J Biol Sci,1988,41(2):177-187.
[17] McDonald B J,et al. [J]. Aust J Biol,Sci,1982,35(1):33-47.
[18] Maddocks S, et al. [J].Aust J Biol Sci,1985,38(4):405-410,
[19] Philpott MP,Sanders DA,Kealey T. Effects of insulin and insulin-like growth factors on cultured human hair follicles:IGF-I at physiologic concentiations is an important regulator of
hair follicle growth in vitro[J]. Invest Dermatol 1994,102:857-861.
[20] Bates E J, P.I. Hynd, N.M.Penn,et al.Serum-free culture of [J].Br J Dermatol,1997,137(4): 498-505.
[21] Bond J J,et al.[J].Biol Anim, 1994,30A:2,90-98.
[22] Bond J J,et al.[J].Arch Dermatol Res,1996,288(7):373-382.
[23]赵改名等.羊毛生长的内分泌调节研究进展[J].中国畜牧杂志,2003,39(3):44-46.
[24]狄江,车文功.羊毛弯曲清晰度与其它数量及质量性状的相关研究[J].草食家畜,1999,4(105): 18-20.
[25]赵国先,宋智娟,张晓云等.毛皮质量的营养调控[J].中国饲料,2005,13:5-9.
[26]赵俊丽.凉山半细毛羊第3号染色体遗传连锁图谱构建及体重性状QTL定位的研究[D].雅安,四川农业大学.2004.
[27]梁春年,姚军,杨博辉,等.高原超细型细毛羊经济性状的微卫星标记研究[J].畜牧兽医学报, 2005, 36(12):1265-1269.
[28]牟艳军,刘国龄,廖和荣,等.利用mRNA差异显示技术筛选绵羊羊毛性状相关基因[J].石河子大学学报(自然科学版),2006,2,24(1):95-97.
[29]张鹦俊.凉山半细毛羊2号染色体羊毛性状的QTL定位研究[D].雅安.四川农业大学硕士,2005.6.
[30] Bray A.R., O’Connell D., Saville D.J., Genes with major effects on wool traits detected in Finn cross sheep, Proc. NZ Soc. Anim. Prod. 62 (2002) 65–68.
[31] Parsons'tim,Piper LR,Cooper DW.Linkage relationships between keratin-associated protein(KRTAP) genes and growth hormone in sheep[J].Genomics.1994,20(3):500-502
[32] Henry H.M, Dodds K. G.,Wuliji T., et a1..A genome screen for QTL for wool traits in a Merino×Romney backcross flock[J].Wool Technology and Sheep Breeding,1998(46):213-217.
[33] Ponz R,et a1.Assessement of genetic variation explained by markers for wool traits in sheep via segment mapping approach,Mamm.Genome,2001(12):569-572.
[34] Rogers G.R,Hickford J.G.H,Biekerstaffe R.Polymorphism in two genes for B2 sulfur proteins of wool,Anim[J].Genet,1994(25):407-415.
[35] Allain D,et al.Adesign aiming at detecting QTL controlling wool traits and other traitsin the INRA401 sheep line,in:Proc.6th World Cong.Genet.App1.Livest [J].Prod.Armidale.1998, 24:51-54.
[36]薛科邦,左秉峰.滩羊主要经济性状的表型相关及回归分析[J].中国畜牧杂志,1992.28(1): 27-28.
[37]马振东.滩羊二毛羔羊主要性状遗传相关的分析[J].中国草食动物.2001,3(1):20-21.
[38]李积友,刘霞,武广善,等.甘肃滩羊血液蛋白多态性与部分生产性能相关关系的研究[J].甘肃农业大学学报,2001,36(3):322-328.
[39]张振汉,魏智清,于洪川,等.宁夏滩羊体大品系血清生长激素及生长介素的测定分析研究[C].中国草食家畜,2003,Z1:137-138.
[40]于洪川等.滩羊体大品系生长激素基因的RFLPs的检测[J].甘肃畜牧兽医,2002,32(2):6-8.
[41]于洪川,魏智清,张振汉,等.滩羊体大品系遗传标记的研究[J].中国草食家畜,2003, Z1: 44-48.
[42]于洪川等.宁夏滩羊体大品系OB基因的RFLP检测[J].中国草食家畜,2003,Z1:73-74.
[43]赵倩君.中国部分绵羊群体的起源、遗传多样性及保护研究[D].北京.中国农业科学院.2007.6.
[44]武艳萍.五个山羊品种微卫星DNA的遗传多样性研究[D].太谷.山西农业大学.2005.6
[45]郭军.中国主要绵羊类群进化关系分析[D].北京.中国农业科学院.2007.6.
[46]仲涛.中国部分绵羊品种的遗传多样性研究[D].北京.中国农业科学院.2007.6
[47]张明亚.凉山半细毛羊第1号染色体遗传连锁图谱构建及羊毛性状QTL定位研究[D].雅安,四川农业大学.2004.5.
[48]陈艺.凉山半细毛羊第2号染色体遗传连锁框架图的构建及体重性状QTL定位的研究[D].雅安,四川农业大学.2004.5.
[49]周明亮.凉山半细毛羊第3号染色体遗传连锁图谱构建及体重性状QTL定位的研究[D].雅安,四川农业大学.2005.5.
[50]赵俊丽.利用微卫星进行凉山半细毛羊羊毛性状的QTL定位研究[D].雅安,四川农业大学,2002.6.
[51]鲁绍雄,连林生.SAS统计分析系统在畜牧科学中的应用[M].昆明:云南科技出版社,2001.94-97.
[52]高之仁.数量遗传学[M].成都:四川大学出版社,1986.
[53] Ian William Purvis et al.,Major genes and QTL influencing wool production and quality: a review[J]. Genet. Sel. 2005 ,Evol. 37(1):S97–S107.
[54] Lande R.Isolation by distance in a quantitative trait[J].Genetics.1991,26:451-466.
[55]狄江,王琼,李勇,等.毛囊角蛋白基因及表达[J].草食家畜.2004,125(4):17-19.
[56]刘霞,李积友,武广善,等.甘肃滩羊转铁蛋白及白蛋白基因座多态性研究[J].甘肃农业大学学报.2001,36(4): 374-378.
[57]于洪川,龚伟宏,杜立新,等.滩羊生长激素基因PCR-RFLPs反应体系的优化[J].宁夏农学院学报, 2003,24(1): 29-33.
[58]魏智清,于洪川,张振汉,等.宁夏滩羊体大品系、裘皮品系及其杂交后代血红蛋白多态性比较[J].中国草食家畜.2003,Z1:57-58.
[59]陈巍,龚伟宏.宁夏滩羊血清蛋白组分含量的研究[J].中国草食家畜.2003,Z1:70-71.
[60]鲁生霞,常洪,角田建司等.利用结构基因座分析小尾寒羊、滩羊群体遗传分化水平[J].中国农业科学, 2005,38(9):1890-1897.
[61]于洪川,魏智清.双羔滩羊血红蛋白多态性的研究[J].宁夏大学学报(自然科学版),1999,20(1):90-91.
[62] Almasy L. and Blangero J. Multipoint quantitative-trait linkage analysis in general pedigrees[J]. Am,J.Hum.GENET.1998,62;1198-1211.
[63] Bovenhuis H, Spelman RJ.Selective genotyping to detect quantitative trait loci for multiple traits in outbred populations[J]. Dairy Sci. 2000 Jan; 83(1):173-180.
[64] Christopher J. Basten.Bruce S. Weir, Zhao-Bang Zeng. QTL cartographer Version 1.15 use manual,2001,6.30.
[65] Crawford, Dodds KG, Ede AJ, et.al.An Autosomal Genetic Linkage Map Of The Sheep Genome[J]. Genetics.1995,140(2):703-724.
[66] Allian D., Lantier L., Elsen J.M.,et.al. A design aiming at detecting QTL controlling wool traits and other traits in the INRA401 sheep line[C].proc.6th WCGALP.52-54.
[67] Lee G, J. and Archibald A.L. Detection of quantitative trait loci in complex pedigrees: a two-sheep variance component approach[J].Genetics.2000;156;2081-2092.
[68] Guo Xiao-Ling, Looft Christian, Reinsch Norbert. A Porcine Linkage Map of Microsatellite Makers Using a Commercial Population as a Reference[J].Acta Genetica Sinica.2003,30(7): 653-656.
[69] Hediger R, Ansari HA, Stranzinger GF. Chromoosome banding and gene locations support extensive conservation of chromosome structure between cattle and sheep[J].Cytogenet Cell Genet. 1991,57:127-134.
[70] Kao C-H, Zeng Z-B, Teasdele R D. Multiple Interval mapping for quantitative trait loci[J]. Genetics. 1999,21:158-159.
[71] King AH. Jiang Z. Mapping Quantitave Trait loci Affecting Female Reproductive Traits on Porcine Chromosome 8[J]. Biol Reprod.2003,22[qpud ahead of print].
[72] Kiniecyzn, A. and Ausubel F.M. A procedure for mapping Arabidopsis mutations using co-dominant ecotype-specific PCR-base markers[J]. Plant.1993,4:403-410.
[73] Knott S.A.,Elsen J-M. and Haley C.S. Methods for multiple-maker mapping of quantitative trait loci in half-sib population[J].Theoretical and Applied Genetics.1996,93:71-80.
[74] Owen J.L., Gomez-Raya L. A genome scan for quantitative trait loci affecting milk production in Norwegian dairy cattle[J].Dairy Sci.2002,85(11):3124-3130.
[75] Parsons YM, and Cooper DW. Linkage of wool protein genes in the sheep genome[C].Proc Aust Assoc Anim Breed Genet.1995,11:469-472.
[76] Plowman JE, Bryson WG, Jordan TW. Application of proteomics for determining protein markers for wool quality traits[J].Electrophoresis.2000,21(9):1899-1906.
[77] Robert J. Mc laren, Geraldine R. et al. Linkage mapping of wool keratin and keratin-associated protein genes in sheep[J]. Mammalian Genome.1997,8:938-940.
[78] Robison BD. Composite interval mapping reveals a major locus influencing embryonic development rate in rainbow trout(Oncorhynchus mykiss)[J].Red.2001,92(1):16-22.
[79] Wada Y., Akita T. Quantitative trait loci(QTL) analysis in a Meishan×Gottingen cross population[J]. anim Genet. 2000,31(6):376-384.
[80] Tuiskula-Haavisto M, Honkatukia M. Mapping of quantitative trait loci affecting quality and production traits in egg layers[J]. Poult Sci.2002.81(7):919-927.
[81] Wilkie PJ. Paszek AA. Agenomic scan of porcine reproductive traits reveals possible quantitative trait loci(QTLs) for number of corpora lutea[J].Mamm Genome.1999,10(6): 573-578.
[82] Zeng Z-B. Precision mapping of quantitative trait loci[J]. Gennetics.1994,136:1457-1468.
[83]储明星译,Falconer D.s,Marckay T.F.C.eds.数量遗传学导论.北京,中国农业科技出版社,2000,281-295.
[84]何小红,徐辰武,蒯建敏,等.构建数量性状基因图谱的几种统计方法[J].遗传,2001,23(5):482-486.
[85]屈彦纯,邓昌彦.猪6号染色体部分遗传连锁图谱的构建与QTL定位[J].华中农业大学学报, 2002, 21(5):409-413.
[86]左波,熊远著,苏玉虹等.利用24个微卫星进行猪数量性状定位及其遗传效应分析[J].畜牧兽医学报. 2003,34(2):139-146.
[87] Jasen R C. Interval mapping of multiple quantitative trait loci[J]. Genetics.1993,135:205-211.
[88]徐云碧,陈英.QTL区间作图的统计理论和计算机软件及其应用[J].作物学报.1995, 21(1): 1-8.
[89]苏玉虹,熊远著,张勤等.大白×梅山猪资源家系生长性状QTL的检测[J].遗传学报.2002,29(7): 607-611.
[90]张付军,李宁.动物主基因(QTL)的定位策略[J].农业生物技术学报.1996,4(2):166-173.
[91]高爱保,吴登俊.利用微卫星标记进行凉山半细毛羊亲权鉴定研究[J].遗传.2005,27(1): 085-090.
[92]李玉奎,吴常信. RFLP在动物育种中应用的前景兼对数量性状基因定位的的讨论[J].中国畜牧杂志. 1991.27: 57-59
[2]余雳.猪2、4号Chr.连锁图构建和肉质性状QTL的鉴定[D].武汉:华中农业大学,2001,5.
[3]狄冉,马月辉.微卫星标记及其在绵、山羊中的研究进展[J].家畜生态学报.2006,27(1):96-100
[4]周明亮.凉山半细毛羊第3号染色体遗传连锁图谱构建及体质性状QTL定位的研究[D].雅安,四川农业大学, 2005.5.
[5]吴登俊,马丁·菲尔斯特.主要家畜种基因组连锁图现状和展望[J].国外畜牧科技.1999, 26(6):21-25.
[6]周明亮,吴登俊,曾云琦.绵羊遗传连锁图的研究现状[J].贵州农业科学.2005,33(3):96-97.
[7]涂友仁等.中国羊品种志.上海科学技术出版社[M].1989.4.:45-47.
[8] Rust C C.Hormonal Control of pelage cycles in the mink[J].J Mammal,1965,(46):549-565.
[9] Rose J.Induction of winter fur growth in mink with melatonin[J]. Anim Sci,1984,58(1):57.
[10] Rose J.The effects of photoperiod melatonin on serum prolaction levels of mink during the autumn moult.[J]. Pineal Research,1985,(2):13.
[11]林仲凡.毛皮动物埋植褪黑激素的研究进展[J].动物学杂志,1988.23(5):45-47.
[12]柳建昌,尹协镇,方天祺.褪黑素对中国绒山羊在非生绒期促绒生长与绒产量的影响[J].动物学杂志,1998,33(3):8-12.
[13]柳建昌.含褪黑素微型胶囊促使银狐毛皮提早成熟的效应[J].动物学杂志,1995,30(1):48-51.
[14]盛雪.褪黑素植入物在狐狸毛皮成熟方面的应用[J].山东畜牧兽医,1997,(2):51-52.
[15]谢宝胜,杨占兴,马武龙,等.褪黑素对门源牦牛春季促绒生长及绒产量的影响[J].中国草食动物,1999,1(6):12-13.
[16] Wynn PC, et al. Randomised controlled trial of intravenous antibiotic treatme.[J].Aust J Biol Sci,1988,41(2):177-187.
[17] McDonald B J,et al. [J]. Aust J Biol,Sci,1982,35(1):33-47.
[18] Maddocks S, et al. [J].Aust J Biol Sci,1985,38(4):405-410,
[19] Philpott MP,Sanders DA,Kealey T. Effects of insulin and insulin-like growth factors on cultured human hair follicles:IGF-I at physiologic concentiations is an important regulator of
hair follicle growth in vitro[J]. Invest Dermatol 1994,102:857-861.
[20] Bates E J, P.I. Hynd, N.M.Penn,et al.Serum-free culture of [J].Br J Dermatol,1997,137(4): 498-505.
[21] Bond J J,et al.[J].Biol Anim, 1994,30A:2,90-98.
[22] Bond J J,et al.[J].Arch Dermatol Res,1996,288(7):373-382.
[23]赵改名等.羊毛生长的内分泌调节研究进展[J].中国畜牧杂志,2003,39(3):44-46.
[24]狄江,车文功.羊毛弯曲清晰度与其它数量及质量性状的相关研究[J].草食家畜,1999,4(105): 18-20.
[25]赵国先,宋智娟,张晓云等.毛皮质量的营养调控[J].中国饲料,2005,13:5-9.
[26]赵俊丽.凉山半细毛羊第3号染色体遗传连锁图谱构建及体重性状QTL定位的研究[D].雅安,四川农业大学.2004.
[27]梁春年,姚军,杨博辉,等.高原超细型细毛羊经济性状的微卫星标记研究[J].畜牧兽医学报, 2005, 36(12):1265-1269.
[28]牟艳军,刘国龄,廖和荣,等.利用mRNA差异显示技术筛选绵羊羊毛性状相关基因[J].石河子大学学报(自然科学版),2006,2,24(1):95-97.
[29]张鹦俊.凉山半细毛羊2号染色体羊毛性状的QTL定位研究[D].雅安.四川农业大学硕士,2005.6.
[30] Bray A.R., O’Connell D., Saville D.J., Genes with major effects on wool traits detected in Finn cross sheep, Proc. NZ Soc. Anim. Prod. 62 (2002) 65–68.
[31] Parsons'tim,Piper LR,Cooper DW.Linkage relationships between keratin-associated protein(KRTAP) genes and growth hormone in sheep[J].Genomics.1994,20(3):500-502
[32] Henry H.M, Dodds K. G.,Wuliji T., et a1..A genome screen for QTL for wool traits in a Merino×Romney backcross flock[J].Wool Technology and Sheep Breeding,1998(46):213-217.
[33] Ponz R,et a1.Assessement of genetic variation explained by markers for wool traits in sheep via segment mapping approach,Mamm.Genome,2001(12):569-572.
[34] Rogers G.R,Hickford J.G.H,Biekerstaffe R.Polymorphism in two genes for B2 sulfur proteins of wool,Anim[J].Genet,1994(25):407-415.
[35] Allain D,et al.Adesign aiming at detecting QTL controlling wool traits and other traitsin the INRA401 sheep line,in:Proc.6th World Cong.Genet.App1.Livest [J].Prod.Armidale.1998, 24:51-54.
[36]薛科邦,左秉峰.滩羊主要经济性状的表型相关及回归分析[J].中国畜牧杂志,1992.28(1): 27-28.
[37]马振东.滩羊二毛羔羊主要性状遗传相关的分析[J].中国草食动物.2001,3(1):20-21.
[38]李积友,刘霞,武广善,等.甘肃滩羊血液蛋白多态性与部分生产性能相关关系的研究[J].甘肃农业大学学报,2001,36(3):322-328.
[39]张振汉,魏智清,于洪川,等.宁夏滩羊体大品系血清生长激素及生长介素的测定分析研究[C].中国草食家畜,2003,Z1:137-138.
[40]于洪川等.滩羊体大品系生长激素基因的RFLPs的检测[J].甘肃畜牧兽医,2002,32(2):6-8.
[41]于洪川,魏智清,张振汉,等.滩羊体大品系遗传标记的研究[J].中国草食家畜,2003, Z1: 44-48.
[42]于洪川等.宁夏滩羊体大品系OB基因的RFLP检测[J].中国草食家畜,2003,Z1:73-74.
[43]赵倩君.中国部分绵羊群体的起源、遗传多样性及保护研究[D].北京.中国农业科学院.2007.6.
[44]武艳萍.五个山羊品种微卫星DNA的遗传多样性研究[D].太谷.山西农业大学.2005.6
[45]郭军.中国主要绵羊类群进化关系分析[D].北京.中国农业科学院.2007.6.
[46]仲涛.中国部分绵羊品种的遗传多样性研究[D].北京.中国农业科学院.2007.6
[47]张明亚.凉山半细毛羊第1号染色体遗传连锁图谱构建及羊毛性状QTL定位研究[D].雅安,四川农业大学.2004.5.
[48]陈艺.凉山半细毛羊第2号染色体遗传连锁框架图的构建及体重性状QTL定位的研究[D].雅安,四川农业大学.2004.5.
[49]周明亮.凉山半细毛羊第3号染色体遗传连锁图谱构建及体重性状QTL定位的研究[D].雅安,四川农业大学.2005.5.
[50]赵俊丽.利用微卫星进行凉山半细毛羊羊毛性状的QTL定位研究[D].雅安,四川农业大学,2002.6.
[51]鲁绍雄,连林生.SAS统计分析系统在畜牧科学中的应用[M].昆明:云南科技出版社,2001.94-97.
[52]高之仁.数量遗传学[M].成都:四川大学出版社,1986.
[53] Ian William Purvis et al.,Major genes and QTL influencing wool production and quality: a review[J]. Genet. Sel. 2005 ,Evol. 37(1):S97–S107.
[54] Lande R.Isolation by distance in a quantitative trait[J].Genetics.1991,26:451-466.
[55]狄江,王琼,李勇,等.毛囊角蛋白基因及表达[J].草食家畜.2004,125(4):17-19.
[56]刘霞,李积友,武广善,等.甘肃滩羊转铁蛋白及白蛋白基因座多态性研究[J].甘肃农业大学学报.2001,36(4): 374-378.
[57]于洪川,龚伟宏,杜立新,等.滩羊生长激素基因PCR-RFLPs反应体系的优化[J].宁夏农学院学报, 2003,24(1): 29-33.
[58]魏智清,于洪川,张振汉,等.宁夏滩羊体大品系、裘皮品系及其杂交后代血红蛋白多态性比较[J].中国草食家畜.2003,Z1:57-58.
[59]陈巍,龚伟宏.宁夏滩羊血清蛋白组分含量的研究[J].中国草食家畜.2003,Z1:70-71.
[60]鲁生霞,常洪,角田建司等.利用结构基因座分析小尾寒羊、滩羊群体遗传分化水平[J].中国农业科学, 2005,38(9):1890-1897.
[61]于洪川,魏智清.双羔滩羊血红蛋白多态性的研究[J].宁夏大学学报(自然科学版),1999,20(1):90-91.
[62] Almasy L. and Blangero J. Multipoint quantitative-trait linkage analysis in general pedigrees[J]. Am,J.Hum.GENET.1998,62;1198-1211.
[63] Bovenhuis H, Spelman RJ.Selective genotyping to detect quantitative trait loci for multiple traits in outbred populations[J]. Dairy Sci. 2000 Jan; 83(1):173-180.
[64] Christopher J. Basten.Bruce S. Weir, Zhao-Bang Zeng. QTL cartographer Version 1.15 use manual,2001,6.30.
[65] Crawford, Dodds KG, Ede AJ, et.al.An Autosomal Genetic Linkage Map Of The Sheep Genome[J]. Genetics.1995,140(2):703-724.
[66] Allian D., Lantier L., Elsen J.M.,et.al. A design aiming at detecting QTL controlling wool traits and other traits in the INRA401 sheep line[C].proc.6th WCGALP.52-54.
[67] Lee G, J. and Archibald A.L. Detection of quantitative trait loci in complex pedigrees: a two-sheep variance component approach[J].Genetics.2000;156;2081-2092.
[68] Guo Xiao-Ling, Looft Christian, Reinsch Norbert. A Porcine Linkage Map of Microsatellite Makers Using a Commercial Population as a Reference[J].Acta Genetica Sinica.2003,30(7): 653-656.
[69] Hediger R, Ansari HA, Stranzinger GF. Chromoosome banding and gene locations support extensive conservation of chromosome structure between cattle and sheep[J].Cytogenet Cell Genet. 1991,57:127-134.
[70] Kao C-H, Zeng Z-B, Teasdele R D. Multiple Interval mapping for quantitative trait loci[J]. Genetics. 1999,21:158-159.
[71] King AH. Jiang Z. Mapping Quantitave Trait loci Affecting Female Reproductive Traits on Porcine Chromosome 8[J]. Biol Reprod.2003,22[qpud ahead of print].
[72] Kiniecyzn, A. and Ausubel F.M. A procedure for mapping Arabidopsis mutations using co-dominant ecotype-specific PCR-base markers[J]. Plant.1993,4:403-410.
[73] Knott S.A.,Elsen J-M. and Haley C.S. Methods for multiple-maker mapping of quantitative trait loci in half-sib population[J].Theoretical and Applied Genetics.1996,93:71-80.
[74] Owen J.L., Gomez-Raya L. A genome scan for quantitative trait loci affecting milk production in Norwegian dairy cattle[J].Dairy Sci.2002,85(11):3124-3130.
[75] Parsons YM, and Cooper DW. Linkage of wool protein genes in the sheep genome[C].Proc Aust Assoc Anim Breed Genet.1995,11:469-472.
[76] Plowman JE, Bryson WG, Jordan TW. Application of proteomics for determining protein markers for wool quality traits[J].Electrophoresis.2000,21(9):1899-1906.
[77] Robert J. Mc laren, Geraldine R. et al. Linkage mapping of wool keratin and keratin-associated protein genes in sheep[J]. Mammalian Genome.1997,8:938-940.
[78] Robison BD. Composite interval mapping reveals a major locus influencing embryonic development rate in rainbow trout(Oncorhynchus mykiss)[J].Red.2001,92(1):16-22.
[79] Wada Y., Akita T. Quantitative trait loci(QTL) analysis in a Meishan×Gottingen cross population[J]. anim Genet. 2000,31(6):376-384.
[80] Tuiskula-Haavisto M, Honkatukia M. Mapping of quantitative trait loci affecting quality and production traits in egg layers[J]. Poult Sci.2002.81(7):919-927.
[81] Wilkie PJ. Paszek AA. Agenomic scan of porcine reproductive traits reveals possible quantitative trait loci(QTLs) for number of corpora lutea[J].Mamm Genome.1999,10(6): 573-578.
[82] Zeng Z-B. Precision mapping of quantitative trait loci[J]. Gennetics.1994,136:1457-1468.
[83]储明星译,Falconer D.s,Marckay T.F.C.eds.数量遗传学导论.北京,中国农业科技出版社,2000,281-295.
[84]何小红,徐辰武,蒯建敏,等.构建数量性状基因图谱的几种统计方法[J].遗传,2001,23(5):482-486.
[85]屈彦纯,邓昌彦.猪6号染色体部分遗传连锁图谱的构建与QTL定位[J].华中农业大学学报, 2002, 21(5):409-413.
[86]左波,熊远著,苏玉虹等.利用24个微卫星进行猪数量性状定位及其遗传效应分析[J].畜牧兽医学报. 2003,34(2):139-146.
[87] Jasen R C. Interval mapping of multiple quantitative trait loci[J]. Genetics.1993,135:205-211.
[88]徐云碧,陈英.QTL区间作图的统计理论和计算机软件及其应用[J].作物学报.1995, 21(1): 1-8.
[89]苏玉虹,熊远著,张勤等.大白×梅山猪资源家系生长性状QTL的检测[J].遗传学报.2002,29(7): 607-611.
[90]张付军,李宁.动物主基因(QTL)的定位策略[J].农业生物技术学报.1996,4(2):166-173.
[91]高爱保,吴登俊.利用微卫星标记进行凉山半细毛羊亲权鉴定研究[J].遗传.2005,27(1): 085-090.
[92]李玉奎,吴常信. RFLP在动物育种中应用的前景兼对数量性状基因定位的的讨论[J].中国畜牧杂志. 1991.27: 57-59