瘦肉型猪选育群的遗传分析
摘要
应用GBS软件(动物模型BLUP)和MTDFREML法对湖南正虹和益阳两个种猪场猪群的繁殖性状和生长发育性状进行了遗传分析。数据来源于两个种猪场的大约克、长白和杜洛克1998-2003年间4061头繁殖母猪的记录(总产仔数、产活仔数、初生窝重、21日龄窝重)和2000-2003年间共1171头后备猪性能测定记录(四月龄体重、测定日龄、背膘厚、体长、胸围、体高、腿臀围及六月龄体重、测定日龄、背膘厚、体长、胸围、体高、腿臀围)，其中公猪485头，母猪686头。估计了主要性状的育种值和遗传参数，同时分析了主要性状的遗传趋势。
通过估计育种值发现，益阳种猪场大约克、杜洛克的达100Kg日龄育种值分别下降了0.45和0.44天，背膘厚育种值分别下降了0.21和0.07毫米，父系指数分别提高了3.99和3.06，正虹种猪场大约克、长白、杜洛克的达100Kg日龄育种值分别下降了1.21、1.17和0.87天，背膘厚育种值分别下降了0.25、0.45和0.36毫米，父系指数分别提高了7.86、4.76和4.81。
繁殖性状估计遗传力的变化范围为0.07～0.21。大约克、长白和杜洛克三个品种的繁殖性状总产仔数(TNB)、产活仔数(NBA)、初生窝重(LBW)的遗传力分别为0.12、0.13、0.15，0.10、0.13、0.07，0.15、0.21、0.12。生长发育性状估计遗传力的变化范围为0.11～0.58，120日龄个体重(W_(120))、120日龄背膘厚(BF_(120))、120日龄体长(BL_(120))、120日龄胸围(CG_(120))、120日龄腿臀围(HG_(120))、120日龄体高(BH_(120))的遗传力估计值，大约克分别为0.41、0.55、0.34、0.30、0.39、0.28，杜洛克分别为0.35、0.29、0.32、0.45、0.34、0.18；180日龄个体重(W_(180))、180日龄体长(BL_(180))、180日龄胸围(CG_(180))、180日龄腿臀围(HG_(180))、180日龄体高(BH_(180))和180日龄背膘厚(BF_(180))的遗传力估计值，大约克分别为0.49、0.53、0.38、0.37、0.11、0.36，长白分别为0.52、0.55、0.36、0.31、0.20、0.30，杜洛克分别为0.58、0.46、0.43、0.43、0.27、0.34。
繁殖性状和生长发育性状的遗传相关：三个品种繁殖性状NBA／TNB、NBA／LBW、TNB／LBW的遗传相关估计值分别为0.78、0.56、0.43，0.79、0.68、0.56，0.82、0.75、0.62。估计的相关中，NBA／TNB呈较强正相关，NBA／LBW、TNB／LBw呈中等正相关。生长发育性状中W_(120)／BF_(120)、W_(180)／BF_(180)的遗传相关估计值为负的强相关，体尺性状间的遗传相关为较低的正相关。
各性状的遗传趋势：繁殖性状TNB、NBA、LBW的遗传趋势大约克、长白、杜洛克分别为0.0038、0.0583、0.0677，0.0024、0.0359、0.0235，0.0032、0.0421、0.0187，遗传趋势是有利的。生长发育性状的遗传趋势与表型趋势一致，也是有利的。
本研究结果表明，这两个猪场通过多年的不断选育，各性状基本朝着有利的方向发展，猪群质量在不断提高，这是符合当前育种目标要求的。
Genetic analyses had been carried out on reproduction and production traits both Zhenghong and Yiyang farm of Hunan. Multi-traits animal model BLUP and MTDFREML were used to estimate variance and covariance of estimate breeding value(EBV), additive effect, maternal effect, maternal permanent environmental effect, common litter effect, and residual effect on 4061 records of reproduction traits from 1998 to 2003, and 1171 records of production traits from 2000 to 2003 .The traits were analyzed including total number of piglets born(TNB), number of piglets born alive (NBA), litter weight of born(LBW), age of reach 100Kg(AGE100),weight of 120 day (W,2o),back fat thickness of 120 day(BF120),body length of 120 day(BL120), chest girth of 120 day(CG120), hip girth of 120 day(HG120), body height of 120 day(BH120), weight of 180 day(W180), body length of 180 day(BL180) chest girth of 180 day(CG180) hip girth of 180 day(HG180), body height of 180 day(WH180) and back fat thickness of 180 day(BF180).
The results showed that EBV of AGE100 of Yorkshire and Duroc in Yiyang farm decreased 0.45 day and 0.44 day, And their EBV of BF decreased 0.21mm and 0.07mm, respectively. At the same time, patrilineal index increased 3.99 and 3.06, respectively. Furthermore, EBV of AGE100 and BF of Yorkshire, Landrace and Duroc in Zhenghong farm also reduced 1.21, 1.17, 0.87day and 0.25, 0.45, 0.36mm, patrilineal index has increased 7.86, 4.76 and 4.81, respectively.
The estimated heritability of reproductive traits ranged 0.07-0.21. Heritability of TNB,NBA and LBW in Yorkshire, Landrace and Duroc were 0. 12, 0. 13, 0. 15, 0.10, 0.13, 0. 07, 0.15, 0. 21, 0. 12, respectively. However, the estimated heritability of production traits ranged 0.11-0.58. Heritability of W120, BF120,BL120, CG120, HG120, BH120 in Yorkshire were 0.41,0.55,0.34,0.30,0.39,0.28 and in Duroc were 0.35, 0.29, 0.32, 0.45, 0.34,0.18, respectively. Heritability of W180, BF180, BL180, CG180,HG180, BH180 in Yorkshire were 0.49,0.53,0.38,0.37,0.11,0.36, in Landrace were 0.52, 0.55, 0.36, 0.31,0.20 0.30 and in Duroc were 0.58, 0.46,0.43,0.43,0.27,0.34,respectively.
Genetic correlation of NBA/TNB, NBA/LBW, TNB/LBW for Yorkshire, Landrace and Duroc was 0. 78, 0. 56, 0. 43, 0. 79, 0. 68, 0. 56, 0. 82, 0. 75, 0. 62, respectively. And genetic correlation of W120/BF120, W180/BF180 was negative, genetic correlation of the other traits were positive.
The genetic trends of reproduction traits and production traits were favorable. The genetic trends of TNB, NBA, LBW for Yorkshire, Landrace and Duroc were 0.0038, 0.0583, 0.0677, 0.0024, 0.0359, 0.0235, 0.0032, 0.0421, 0.0187, respectively. And production traits, the genetic trends were coincide with the performance trends, they were all favorable.
All the results showed that the developing tendency of all traits was favorable, and the herds quality of the two farms have been improved by breeding over the past years.
通过估计育种值发现，益阳种猪场大约克、杜洛克的达100Kg日龄育种值分别下降了0.45和0.44天，背膘厚育种值分别下降了0.21和0.07毫米，父系指数分别提高了3.99和3.06，正虹种猪场大约克、长白、杜洛克的达100Kg日龄育种值分别下降了1.21、1.17和0.87天，背膘厚育种值分别下降了0.25、0.45和0.36毫米，父系指数分别提高了7.86、4.76和4.81。
繁殖性状估计遗传力的变化范围为0.07～0.21。大约克、长白和杜洛克三个品种的繁殖性状总产仔数(TNB)、产活仔数(NBA)、初生窝重(LBW)的遗传力分别为0.12、0.13、0.15，0.10、0.13、0.07，0.15、0.21、0.12。生长发育性状估计遗传力的变化范围为0.11～0.58，120日龄个体重(W_(120))、120日龄背膘厚(BF_(120))、120日龄体长(BL_(120))、120日龄胸围(CG_(120))、120日龄腿臀围(HG_(120))、120日龄体高(BH_(120))的遗传力估计值，大约克分别为0.41、0.55、0.34、0.30、0.39、0.28，杜洛克分别为0.35、0.29、0.32、0.45、0.34、0.18；180日龄个体重(W_(180))、180日龄体长(BL_(180))、180日龄胸围(CG_(180))、180日龄腿臀围(HG_(180))、180日龄体高(BH_(180))和180日龄背膘厚(BF_(180))的遗传力估计值，大约克分别为0.49、0.53、0.38、0.37、0.11、0.36，长白分别为0.52、0.55、0.36、0.31、0.20、0.30，杜洛克分别为0.58、0.46、0.43、0.43、0.27、0.34。
繁殖性状和生长发育性状的遗传相关：三个品种繁殖性状NBA／TNB、NBA／LBW、TNB／LBW的遗传相关估计值分别为0.78、0.56、0.43，0.79、0.68、0.56，0.82、0.75、0.62。估计的相关中，NBA／TNB呈较强正相关，NBA／LBW、TNB／LBw呈中等正相关。生长发育性状中W_(120)／BF_(120)、W_(180)／BF_(180)的遗传相关估计值为负的强相关，体尺性状间的遗传相关为较低的正相关。
各性状的遗传趋势：繁殖性状TNB、NBA、LBW的遗传趋势大约克、长白、杜洛克分别为0.0038、0.0583、0.0677，0.0024、0.0359、0.0235，0.0032、0.0421、0.0187，遗传趋势是有利的。生长发育性状的遗传趋势与表型趋势一致，也是有利的。
本研究结果表明，这两个猪场通过多年的不断选育，各性状基本朝着有利的方向发展，猪群质量在不断提高，这是符合当前育种目标要求的。
Genetic analyses had been carried out on reproduction and production traits both Zhenghong and Yiyang farm of Hunan. Multi-traits animal model BLUP and MTDFREML were used to estimate variance and covariance of estimate breeding value(EBV), additive effect, maternal effect, maternal permanent environmental effect, common litter effect, and residual effect on 4061 records of reproduction traits from 1998 to 2003, and 1171 records of production traits from 2000 to 2003 .The traits were analyzed including total number of piglets born(TNB), number of piglets born alive (NBA), litter weight of born(LBW), age of reach 100Kg(AGE100),weight of 120 day (W,2o),back fat thickness of 120 day(BF120),body length of 120 day(BL120), chest girth of 120 day(CG120), hip girth of 120 day(HG120), body height of 120 day(BH120), weight of 180 day(W180), body length of 180 day(BL180) chest girth of 180 day(CG180) hip girth of 180 day(HG180), body height of 180 day(WH180) and back fat thickness of 180 day(BF180).
The results showed that EBV of AGE100 of Yorkshire and Duroc in Yiyang farm decreased 0.45 day and 0.44 day, And their EBV of BF decreased 0.21mm and 0.07mm, respectively. At the same time, patrilineal index increased 3.99 and 3.06, respectively. Furthermore, EBV of AGE100 and BF of Yorkshire, Landrace and Duroc in Zhenghong farm also reduced 1.21, 1.17, 0.87day and 0.25, 0.45, 0.36mm, patrilineal index has increased 7.86, 4.76 and 4.81, respectively.
The estimated heritability of reproductive traits ranged 0.07-0.21. Heritability of TNB,NBA and LBW in Yorkshire, Landrace and Duroc were 0. 12, 0. 13, 0. 15, 0.10, 0.13, 0. 07, 0.15, 0. 21, 0. 12, respectively. However, the estimated heritability of production traits ranged 0.11-0.58. Heritability of W120, BF120,BL120, CG120, HG120, BH120 in Yorkshire were 0.41,0.55,0.34,0.30,0.39,0.28 and in Duroc were 0.35, 0.29, 0.32, 0.45, 0.34,0.18, respectively. Heritability of W180, BF180, BL180, CG180,HG180, BH180 in Yorkshire were 0.49,0.53,0.38,0.37,0.11,0.36, in Landrace were 0.52, 0.55, 0.36, 0.31,0.20 0.30 and in Duroc were 0.58, 0.46,0.43,0.43,0.27,0.34,respectively.
Genetic correlation of NBA/TNB, NBA/LBW, TNB/LBW for Yorkshire, Landrace and Duroc was 0. 78, 0. 56, 0. 43, 0. 79, 0. 68, 0. 56, 0. 82, 0. 75, 0. 62, respectively. And genetic correlation of W120/BF120, W180/BF180 was negative, genetic correlation of the other traits were positive.
The genetic trends of reproduction traits and production traits were favorable. The genetic trends of TNB, NBA, LBW for Yorkshire, Landrace and Duroc were 0.0038, 0.0583, 0.0677, 0.0024, 0.0359, 0.0235, 0.0032, 0.0421, 0.0187, respectively. And production traits, the genetic trends were coincide with the performance trends, they were all favorable.
All the results showed that the developing tendency of all traits was favorable, and the herds quality of the two farms have been improved by breeding over the past years.
引文
[1]张沅，张勤．畜禽育种中的线性模型[M]．北京：北京农业大学出版社，1993．
[2]盛志廉，陈瑶生．数量遗传学[M]．北京：科学出版社，1999．
[3]彭中镇．猪的遗传改良[M]．北京：农业出版社，1990．
[4]施启顺，柳小春．养猪业中的杂种优势利用[M]．湖南：湖南科技出版社，1997．
[5]陈瑶生，中国动物遗传育种研究进展[C]，北京：中国农业科学技术出版社，2003
[6]GBS工作室．GBS种猪生产管理及育种分析系统[M]．北京：中国农业科技出版社，1999．
[7]储明星，吴常信，张建生．二花脸猪和大白猪产仔数加性-显性遗传模型以及基因效应的研究[J]．南京农业大学学报，2001，(1)：89-91．
[8]张保军，孙超，张丽娟等．猪育种趋势及其研究进展[J]．家畜生态，2002，23(4)：45-48．
[9]郑友民，王立贤．国外对太湖猪的研究进展[J]．国外畜牧科技，1998，(4)：22-26．
[10]储明星，吴常信，张建生等．母猪生产力性状的遗传分析[J]．遗传学报，2000，(11)：947-952．
[11]杨泽民，熊远著，喻传洲．影响猪遗传评估效果的主要因素研究[J]．华中农业大学学报，2001，(6)：598-602．
[12]郑友民，王立贤，张沅．用MTDFREML方法估计大白猪遗传参数[J]．养猪，1999，(1)：32-33．
[13]张伟峰，张沅．应用动物模型REML方法估计遗传参数的研究[J]．养猪，1999，(3)：31-3225．
[14]陈海燕，朱海平，傅衍．影响外来猪种早期生长的固定效应[J]．浙江大学学报，2003，29(4)：429-432．
[15]郑友民，张沅．大白猪繁殖和生产性状母体遗传效应估计[J]．畜牧兽医学报，1999，(5)：394-398．
[16]储明星，吴常信，张建生等．猪窝产仔数的遗传分析[J]．遗传学报，2001，(4)：317-321．
[17]张胜利，杨秀文，靳明武等．遗传参数不同估计方法的比较[J]．遗传，1995，(6)：21-23．
[18]张勤，张沅．方差组分估计方法MIVQUE和REML的模拟比较[J]．遗传学报，1995，(6)：424-430．
[19]施启顺，柳小春，李元英．宁乡猪数量性状遗传规律的研究Ⅰ繁殖性状[J]．遗传，1986，8(1)23-26．
[20]施启顺，柳小春，李元英．宁乡猪数量性状遗传规律的研究Ⅱ肥育与胴体性状[J]．遗传，1986，8(1)27-30．
[21]陈斌，柳小春，施启顺等．长白猪数量性状表型和遗传参数估测[A]，见：施启顺主编．中国畜禽遗传育种进展(第七次全国畜禽遗传育种学术讨论会论文集)[C]．长沙：湖南科学技术出版社，1993．19-24．
[22]施启顺、柳小春、李元英．宁乡猪数量性状遗传规律研究Ⅲ．生长发育性状[J]．遗传，1986，8(4)：24-26．
[23]张胜利、靳明武、陈廷槐．遗传参数不同估计方法的比较[J]．见：施启顺主编．中国畜禽遗传育种进展(第七次全国畜禽遗传育种学术讨论会论文集)[C]．长沙：湖南科学技术出版社，1993．56-59．
[24]常英新，张沅，张勤．估计猪早期生长性状和繁殖性状的动物模型比较[J]．黑龙江畜牧兽医，2001，(7)：13-15．
[25]谢保胜，徐宁迎．应用动物模型REML法估计金华猪生长性状遗传参数[J]．黑龙江畜牧兽医，2002，10，1-3．
[26]金访中，周珠丽．长白猪生长发育性状的遗传参数估测[J]．浙江畜牧兽医，2001，3．
[27]刘建华，曹果清等．山西瘦肉型猪SD-Ⅱ系生长发育性状遗传参数估测[J]．山西农业大学学报，2001，(3)：212-214．
[28]金访中，周珠丽，钱成刚．长白猪S_Ⅳ系生长发育性状的遗传参数估计[J]．畜牧与兽医，2002，(2)：18-20．
[29]郑友民，张沅．大白猪繁殖和生产性状母体遗传效应估计[J]．畜牧兽医学报，1999，(5)：394-398．
[30]付茂忠，何治平，林小伟．瘦肉系专门化父系杜洛克遗传参数估测[J]．四川畜牧兽医，1997(1)：8-10．
[31]王楚端，马卫东，陈清明等．利用多性状混合模型估计猪繁殖性状的遗传参数[J]．中国农业大学学报，2001，6(3)：96-100．
[32]郑友民，王立贤，张沅．大白猪生产和繁殖性状遗传趋势的分析[J]．中国农业科学，1999，32(3)：85-89．
[33]徐来仁，王维洪，仇志军等．约克夏猪繁殖性状直接的和母体的加性遗传效应分析[J]．畜牧与兽医，1999，31(1)：16-17．
[34]连林生，鲁绍雄．撒坝猪繁殖性状的表型和遗传参数分析[J]．畜牧与兽医，2001，33(2)：6-8．
[35]曹洪战，吴常信，师守堃．利用MTDFREML估计大约克双肌臀种猪活体背膘厚
遗传力[J]．畜牧与兽医，2003，35(1)：6-7．
[36] Henderson.C.R. Estimation of variance and cavariance components Biometrics[J]. 1953, 9:226-252.
[37] Henderson.C.R. Appliations of liner models in animal breeding[J].Univ of Guelph, Ontario. 1984, Canada.
[38] Rao.C.R. Estimation of heteroscedastic variances in linear models[J]. J.Am.Stat Assoc, 1970,65:161-172.
[39] Hartley.H.O, J.N.K.Rao. Maximum-likelihood estimation for the mixed analysis of variance model [J].Biomtrika,1967,54:93-108.
[40] Wang.C.S J,J.Rutledgye, D.Gianola. Marshal inferences about variance components in a mixed linear model using Gibbs sampling[J].Genet Sel Evol, 1993,25:41-64.
[41] Jensen.J, C.S.wang, D.A.Sorenson, et al. Beyesian inference on variance and covariance components for traits influenced by maternal and direct genetic effects using the Gibbs sampler[J]. Acta. Agric. Stand, 1994,44:193-201.
[42] Van.Tassell, C.P.G.Casella, E.J.Pollak. Effects of selection on estimates of variance componects using Gibbs samply and restricted maximum likelihood[J]. J. Dairy Sci, 1995,78:678-692.
[43] Reverter.A.B, L.Golden, R.M.Bourdon, et al. Method R. variance components procedure Application on the simple breeding value model[J]. J.Anim.Sci, 1994,72:2247-2253.
[44] See,J.W.Mabry, J.K.Bertrand. Restricted Maximum likelihood Estimation of variance components from Field Data for Number of pigs Born Alive[J]. J.Anim Sci, 1993,71:2905-2909.
[45] S.Culbertson,J.W.Mabry, I.Miszatal,et.al. estimation of Dominance in purebred Yorkshire swine[J].J.Anim Sci, 1998,76:448-45.
[46] Druet,I.Misztal,M.Duangjinda,et.al. Estimation of genetic covariances with Mothed R[J].J.Anim Sci ,2001,79:605-615.
[47] Crump R E,Haley C S,Thompson R et..al,Individual animal model estimates of genetic parameters for reproduction traits of landrace pigs performance test in a commercial nucleus herd[J]. Animal sci, 1997,65:285-290.
[48] Adamec,V.and Johnson,R.K. Genetic analysis kf rebreeding intervals,litter traits,and production traits in sows kf the National Czech nucleus[J]. 1997,48(1):13-22.
[49] Alfonso,L,Noguera,J.L,Babot,D.Estany,j.Estimation of genetic parameters for
litter size at different parities in pigs[J].Livst.Prod.Sci. 1997, 47(2): 149-156.
[50] Roche R,kermedy B W.Estimation of genetic parameters for litter size in canadian Yorkshine and landrace Swine with each parity of farrowing freated as a different trait[J]. J Anim sci. 1995,73:2959-2970.
[51] Mercer J,T, et.al.Genetic parameter estimates for reproduction traits in purebred landrace pigs[J].proc 4th world congress Genet Apply in Ani. Prod.xv:489-492.
[52] T Ishida, T kuroki H Harada,et..al.Estimation of Additine and Dominance Genetic Variances in Line Breeding Swine[J].Anim Sci.2001,vol 14(1): 1-6.
[53] Bidanel,J.P, Ducos A, Labroue F, et.al.Genetic parameters of backfat thickness,age at 100 kg and meat quality traits in pietrain pigs[J].Ann Zootech.43:141-149.
[54] Bidanel,J.P, Dueos A,Gueblez R,et..al.Genetie parameters of back_fat thickness,age at 100 kg and ultimate pH in on-farm tested French Landrace and Large White pigs[J]. Livestock Sei. 1994,40:291-301.
[55] Cameron N D,Methodologies for estimation of genotype with environment interaction[J].Livest Sci. 1993,35(3-4):237-249.
[56] Cho Y C,Kim G W, Park H Y. Estimation of genetic parameters for major economic traits in swine[J].Korean Journal of Anita. Sci. 1998,40(5):447-454.
[57] Ducos A,Bidanel J P.Genetic coorelations between production and reproduction traits measured in the farm,on the Large White and French Landrace pigs breeds[J].Breed Genet. 1996,113(6):493-504.
[58] Estany J,Sorensen D.Estimation of genetic parameters for litter size in Danish Landrace and Yorkshire pigs[J].Anim Sci. 1995,60(2):315-324.
[59] Falconer D S,Mackey T F C. Introduction to quantitative genetics edition.Longman Harlon. 1996,Essex.
[60] Ferraz J B S Johnson R K. Animal model estimation of genetic parameters and response to selection for litter size and weight,growth and backfat in closed seedstock productions of Large White and Landrace swine[J].J of Anim Sci. 1993,71(4):850-858.
[61] Garreau H,Tribowt T Bidanel J P. New estimation of genetic parameters of production traits in Large White and French Landrace pigs[J], pig News and Information. 1999,20(2): 113.
[62] Hagger C. Litter permanent environmental,ramflock,and genetic effects on early weight gain of lambs[J].J of Anim Sci. 1998,76:452-457.
[63] Hofer A. Variance compenent estimation in animal breeding:a review[J].J Anim Breed genet. 1998,115:247-265.
[64] Keele J W, Long T E, Johnson R K. Comparison of methods of. estimating variance components in pig[J].J Anita Sci. 1991,69(4): 1428-1434.
[65] Kerr J C, Cameron N D. Genetic and phenotypic relationships between performance test and reproduction traits in Large White pigs[J]. Anim Sci. 1996,62(3):531-540.
[66] Meyer K, Smith S P, Restricted Maximum Likelihood estimation for animal models using derivatives of the likelihood[J].Genet sel Evol. 1997,28:23-49.
[67] Meyer K. Estimating variance and covariance for multivariate animal models by restricted maximum likelihood[J].Genet Sel Evol. 1991,23(1):67-83.
[68] Meyer K. Biased and sampling covarianee of estimates of variance components due to maternal effects[J].Genet. Sel. Evol. 1992,24(6):487-509.
[69] Meyer K.variance components due to direct and maternal effects for growth traits of Australian beef cattle[J].Livest.Prod.Sci. 1992,31:197-204.
[70] Misztel I.,Perez-Enciso,M. Sparse matrix inversion for REML estimates of variance components by expectation-maximization[J].J.Dairy.Sci.1993,76(5): 1479-1483.
[71] Mrode,R.A. Kennedy, B.W. Genetic variation in measures of food efficiency in pigs and their genetic relation ships with growth rate and backfat[J].Anim.Prod. 1993,56(2):225-232.
[72] Roehe,R,Kennedy, B.W. Estimation of Genetic parameters for litter size in Canadian Yorkshire and Landrace swine with each parity of farrowing treated as a different trait[J].J, of Anim. Sci. 1995,73 (10):2959-2970.
[73] Roehe,R.and Kennedy, B.W.The influence of maternal effects on accuracy of evaluation of litter size in swine[J].J.Anim.Sci. 1993,71 (10):2353.
[74] See,M.T.Mabry, J.W. and Bertrand,J.K. REML estimates of variance components from field data for number of pigs born alive[J]. J.Anim.Sci. 1993, 71 (11): 2905-2909.
[75] Skorupski,M.K.,Garrick,D.J.,Blair, H.J. Estimation of Genetic parameters for production and reproduction trains in three breeds of pigs[J].New zealand Journal of Agricultural Reseaeh.1996,39(3):387-395.
[76] Southwood, O.I. and Kennedy, B.W. Estimation of direct and maternal genetic variance for litter size in Canadian Yorkshire and Landrace swine using an animal model[J]. J.Anim.Sci. 1990,68:1841-1847.
[77] Southwood,O.I.and Kennedy, B.W. Genetic and environmental trends for litter size in swine[J].J. Anim.Sci. 1991,69:3117-3182.
[2]盛志廉，陈瑶生．数量遗传学[M]．北京：科学出版社，1999．
[3]彭中镇．猪的遗传改良[M]．北京：农业出版社，1990．
[4]施启顺，柳小春．养猪业中的杂种优势利用[M]．湖南：湖南科技出版社，1997．
[5]陈瑶生，中国动物遗传育种研究进展[C]，北京：中国农业科学技术出版社，2003
[6]GBS工作室．GBS种猪生产管理及育种分析系统[M]．北京：中国农业科技出版社，1999．
[7]储明星，吴常信，张建生．二花脸猪和大白猪产仔数加性-显性遗传模型以及基因效应的研究[J]．南京农业大学学报，2001，(1)：89-91．
[8]张保军，孙超，张丽娟等．猪育种趋势及其研究进展[J]．家畜生态，2002，23(4)：45-48．
[9]郑友民，王立贤．国外对太湖猪的研究进展[J]．国外畜牧科技，1998，(4)：22-26．
[10]储明星，吴常信，张建生等．母猪生产力性状的遗传分析[J]．遗传学报，2000，(11)：947-952．
[11]杨泽民，熊远著，喻传洲．影响猪遗传评估效果的主要因素研究[J]．华中农业大学学报，2001，(6)：598-602．
[12]郑友民，王立贤，张沅．用MTDFREML方法估计大白猪遗传参数[J]．养猪，1999，(1)：32-33．
[13]张伟峰，张沅．应用动物模型REML方法估计遗传参数的研究[J]．养猪，1999，(3)：31-3225．
[14]陈海燕，朱海平，傅衍．影响外来猪种早期生长的固定效应[J]．浙江大学学报，2003，29(4)：429-432．
[15]郑友民，张沅．大白猪繁殖和生产性状母体遗传效应估计[J]．畜牧兽医学报，1999，(5)：394-398．
[16]储明星，吴常信，张建生等．猪窝产仔数的遗传分析[J]．遗传学报，2001，(4)：317-321．
[17]张胜利，杨秀文，靳明武等．遗传参数不同估计方法的比较[J]．遗传，1995，(6)：21-23．
[18]张勤，张沅．方差组分估计方法MIVQUE和REML的模拟比较[J]．遗传学报，1995，(6)：424-430．
[19]施启顺，柳小春，李元英．宁乡猪数量性状遗传规律的研究Ⅰ繁殖性状[J]．遗传，1986，8(1)23-26．
[20]施启顺，柳小春，李元英．宁乡猪数量性状遗传规律的研究Ⅱ肥育与胴体性状[J]．遗传，1986，8(1)27-30．
[21]陈斌，柳小春，施启顺等．长白猪数量性状表型和遗传参数估测[A]，见：施启顺主编．中国畜禽遗传育种进展(第七次全国畜禽遗传育种学术讨论会论文集)[C]．长沙：湖南科学技术出版社，1993．19-24．
[22]施启顺、柳小春、李元英．宁乡猪数量性状遗传规律研究Ⅲ．生长发育性状[J]．遗传，1986，8(4)：24-26．
[23]张胜利、靳明武、陈廷槐．遗传参数不同估计方法的比较[J]．见：施启顺主编．中国畜禽遗传育种进展(第七次全国畜禽遗传育种学术讨论会论文集)[C]．长沙：湖南科学技术出版社，1993．56-59．
[24]常英新，张沅，张勤．估计猪早期生长性状和繁殖性状的动物模型比较[J]．黑龙江畜牧兽医，2001，(7)：13-15．
[25]谢保胜，徐宁迎．应用动物模型REML法估计金华猪生长性状遗传参数[J]．黑龙江畜牧兽医，2002，10，1-3．
[26]金访中，周珠丽．长白猪生长发育性状的遗传参数估测[J]．浙江畜牧兽医，2001，3．
[27]刘建华，曹果清等．山西瘦肉型猪SD-Ⅱ系生长发育性状遗传参数估测[J]．山西农业大学学报，2001，(3)：212-214．
[28]金访中，周珠丽，钱成刚．长白猪S_Ⅳ系生长发育性状的遗传参数估计[J]．畜牧与兽医，2002，(2)：18-20．
[29]郑友民，张沅．大白猪繁殖和生产性状母体遗传效应估计[J]．畜牧兽医学报，1999，(5)：394-398．
[30]付茂忠，何治平，林小伟．瘦肉系专门化父系杜洛克遗传参数估测[J]．四川畜牧兽医，1997(1)：8-10．
[31]王楚端，马卫东，陈清明等．利用多性状混合模型估计猪繁殖性状的遗传参数[J]．中国农业大学学报，2001，6(3)：96-100．
[32]郑友民，王立贤，张沅．大白猪生产和繁殖性状遗传趋势的分析[J]．中国农业科学，1999，32(3)：85-89．
[33]徐来仁，王维洪，仇志军等．约克夏猪繁殖性状直接的和母体的加性遗传效应分析[J]．畜牧与兽医，1999，31(1)：16-17．
[34]连林生，鲁绍雄．撒坝猪繁殖性状的表型和遗传参数分析[J]．畜牧与兽医，2001，33(2)：6-8．
[35]曹洪战，吴常信，师守堃．利用MTDFREML估计大约克双肌臀种猪活体背膘厚
遗传力[J]．畜牧与兽医，2003，35(1)：6-7．
[36] Henderson.C.R. Estimation of variance and cavariance components Biometrics[J]. 1953, 9:226-252.
[37] Henderson.C.R. Appliations of liner models in animal breeding[J].Univ of Guelph, Ontario. 1984, Canada.
[38] Rao.C.R. Estimation of heteroscedastic variances in linear models[J]. J.Am.Stat Assoc, 1970,65:161-172.
[39] Hartley.H.O, J.N.K.Rao. Maximum-likelihood estimation for the mixed analysis of variance model [J].Biomtrika,1967,54:93-108.
[40] Wang.C.S J,J.Rutledgye, D.Gianola. Marshal inferences about variance components in a mixed linear model using Gibbs sampling[J].Genet Sel Evol, 1993,25:41-64.
[41] Jensen.J, C.S.wang, D.A.Sorenson, et al. Beyesian inference on variance and covariance components for traits influenced by maternal and direct genetic effects using the Gibbs sampler[J]. Acta. Agric. Stand, 1994,44:193-201.
[42] Van.Tassell, C.P.G.Casella, E.J.Pollak. Effects of selection on estimates of variance componects using Gibbs samply and restricted maximum likelihood[J]. J. Dairy Sci, 1995,78:678-692.
[43] Reverter.A.B, L.Golden, R.M.Bourdon, et al. Method R. variance components procedure Application on the simple breeding value model[J]. J.Anim.Sci, 1994,72:2247-2253.
[44] See,J.W.Mabry, J.K.Bertrand. Restricted Maximum likelihood Estimation of variance components from Field Data for Number of pigs Born Alive[J]. J.Anim Sci, 1993,71:2905-2909.
[45] S.Culbertson,J.W.Mabry, I.Miszatal,et.al. estimation of Dominance in purebred Yorkshire swine[J].J.Anim Sci, 1998,76:448-45.
[46] Druet,I.Misztal,M.Duangjinda,et.al. Estimation of genetic covariances with Mothed R[J].J.Anim Sci ,2001,79:605-615.
[47] Crump R E,Haley C S,Thompson R et..al,Individual animal model estimates of genetic parameters for reproduction traits of landrace pigs performance test in a commercial nucleus herd[J]. Animal sci, 1997,65:285-290.
[48] Adamec,V.and Johnson,R.K. Genetic analysis kf rebreeding intervals,litter traits,and production traits in sows kf the National Czech nucleus[J]. 1997,48(1):13-22.
[49] Alfonso,L,Noguera,J.L,Babot,D.Estany,j.Estimation of genetic parameters for
litter size at different parities in pigs[J].Livst.Prod.Sci. 1997, 47(2): 149-156.
[50] Roche R,kermedy B W.Estimation of genetic parameters for litter size in canadian Yorkshine and landrace Swine with each parity of farrowing freated as a different trait[J]. J Anim sci. 1995,73:2959-2970.
[51] Mercer J,T, et.al.Genetic parameter estimates for reproduction traits in purebred landrace pigs[J].proc 4th world congress Genet Apply in Ani. Prod.xv:489-492.
[52] T Ishida, T kuroki H Harada,et..al.Estimation of Additine and Dominance Genetic Variances in Line Breeding Swine[J].Anim Sci.2001,vol 14(1): 1-6.
[53] Bidanel,J.P, Ducos A, Labroue F, et.al.Genetic parameters of backfat thickness,age at 100 kg and meat quality traits in pietrain pigs[J].Ann Zootech.43:141-149.
[54] Bidanel,J.P, Dueos A,Gueblez R,et..al.Genetie parameters of back_fat thickness,age at 100 kg and ultimate pH in on-farm tested French Landrace and Large White pigs[J]. Livestock Sei. 1994,40:291-301.
[55] Cameron N D,Methodologies for estimation of genotype with environment interaction[J].Livest Sci. 1993,35(3-4):237-249.
[56] Cho Y C,Kim G W, Park H Y. Estimation of genetic parameters for major economic traits in swine[J].Korean Journal of Anita. Sci. 1998,40(5):447-454.
[57] Ducos A,Bidanel J P.Genetic coorelations between production and reproduction traits measured in the farm,on the Large White and French Landrace pigs breeds[J].Breed Genet. 1996,113(6):493-504.
[58] Estany J,Sorensen D.Estimation of genetic parameters for litter size in Danish Landrace and Yorkshire pigs[J].Anim Sci. 1995,60(2):315-324.
[59] Falconer D S,Mackey T F C. Introduction to quantitative genetics edition.Longman Harlon. 1996,Essex.
[60] Ferraz J B S Johnson R K. Animal model estimation of genetic parameters and response to selection for litter size and weight,growth and backfat in closed seedstock productions of Large White and Landrace swine[J].J of Anim Sci. 1993,71(4):850-858.
[61] Garreau H,Tribowt T Bidanel J P. New estimation of genetic parameters of production traits in Large White and French Landrace pigs[J], pig News and Information. 1999,20(2): 113.
[62] Hagger C. Litter permanent environmental,ramflock,and genetic effects on early weight gain of lambs[J].J of Anim Sci. 1998,76:452-457.
[63] Hofer A. Variance compenent estimation in animal breeding:a review[J].J Anim Breed genet. 1998,115:247-265.
[64] Keele J W, Long T E, Johnson R K. Comparison of methods of. estimating variance components in pig[J].J Anita Sci. 1991,69(4): 1428-1434.
[65] Kerr J C, Cameron N D. Genetic and phenotypic relationships between performance test and reproduction traits in Large White pigs[J]. Anim Sci. 1996,62(3):531-540.
[66] Meyer K, Smith S P, Restricted Maximum Likelihood estimation for animal models using derivatives of the likelihood[J].Genet sel Evol. 1997,28:23-49.
[67] Meyer K. Estimating variance and covariance for multivariate animal models by restricted maximum likelihood[J].Genet Sel Evol. 1991,23(1):67-83.
[68] Meyer K. Biased and sampling covarianee of estimates of variance components due to maternal effects[J].Genet. Sel. Evol. 1992,24(6):487-509.
[69] Meyer K.variance components due to direct and maternal effects for growth traits of Australian beef cattle[J].Livest.Prod.Sci. 1992,31:197-204.
[70] Misztel I.,Perez-Enciso,M. Sparse matrix inversion for REML estimates of variance components by expectation-maximization[J].J.Dairy.Sci.1993,76(5): 1479-1483.
[71] Mrode,R.A. Kennedy, B.W. Genetic variation in measures of food efficiency in pigs and their genetic relation ships with growth rate and backfat[J].Anim.Prod. 1993,56(2):225-232.
[72] Roehe,R,Kennedy, B.W. Estimation of Genetic parameters for litter size in Canadian Yorkshire and Landrace swine with each parity of farrowing treated as a different trait[J].J, of Anim. Sci. 1995,73 (10):2959-2970.
[73] Roehe,R.and Kennedy, B.W.The influence of maternal effects on accuracy of evaluation of litter size in swine[J].J.Anim.Sci. 1993,71 (10):2353.
[74] See,M.T.Mabry, J.W. and Bertrand,J.K. REML estimates of variance components from field data for number of pigs born alive[J]. J.Anim.Sci. 1993, 71 (11): 2905-2909.
[75] Skorupski,M.K.,Garrick,D.J.,Blair, H.J. Estimation of Genetic parameters for production and reproduction trains in three breeds of pigs[J].New zealand Journal of Agricultural Reseaeh.1996,39(3):387-395.
[76] Southwood, O.I. and Kennedy, B.W. Estimation of direct and maternal genetic variance for litter size in Canadian Yorkshire and Landrace swine using an animal model[J]. J.Anim.Sci. 1990,68:1841-1847.
[77] Southwood,O.I.and Kennedy, B.W. Genetic and environmental trends for litter size in swine[J].J. Anim.Sci. 1991,69:3117-3182.