三种杂交鱼与纯种尼罗、奥利亚罗非鱼遗传关系研究
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摘要
罗非鱼是联合国粮农组织重点推广的水产养殖品种，它在国际贸易中成为继大马哈鱼和对虾之后的第三大水产品。在我国，罗非鱼的产量年年递增，到2000年我国罗非鱼产量已达63万吨，占世界产量的60％以上。在实际生产过程中，一般利用罗非鱼的正交一代(尼罗♀×奥利亚♂)进行养殖，但由于杂交罗非鱼很容易混在亲本纯种中，导致罗非鱼的二次杂交，如正交一代的兄妹间杂交、正交一代与奥利亚间的回交，由于二次杂交引起后代基因的分配不均匀，导致生产性能不均匀，影响了整体的产量和生产水平。在以往的实验中我们也发现，兄妹间杂交子代与回交子代在增重率，饵料转化率都没有正交一代高，且规格上出现严重的不整齐。
本论文试从形态学和分子水平研究三种不同杂交方式的杂交罗非鱼后代与纯种尼罗罗非鱼及奥利亚罗非鱼之间的遗传关系。比较正交一代、回交子代、兄妹交子代三种杂交鱼与纯种尼罗罗非鱼和奥利亚罗非鱼之间的种内、种间相似系数、遗传距离。从分子水平检验杂交鱼是否具有杂交优势，从分子水平解释三种杂交鱼生产性能存在的差异。同时在形态学上利用最新的统计学技术和软件对纯种鱼和杂交鱼进行区别，并建立新的形态学分析方法，对杂交鱼和纯种鱼进行区别，为选育和培养高品质的罗非鱼养殖品种打下理论基础和提供实践方法。
结果表明：尼罗罗非鱼和奥利亚罗非鱼之间有着较大的遗传距离，使正交一代基因杂合性增加，使其更有可能获得杂交优势，且正交一代与父母本间的种间相似系数没有明显的亲本偏向性，更进一步证明了其正交一代的杂交优势。回交子代和兄妹交子代与奥利亚有着更大的种间相似系数，在种质上更偏向于奥利亚罗非鱼，由于奥利亚在生产中表现出生长速度较慢、饵料转化率较低。故解释了回交子代、兄妹交子代生产性能较差的原因。同时建议对回交子代、兄妹交子代需进一步的选育和杂交，以其获得更高程度的基因的杂合性和优良的性状。
筛选出引物S_(323)可以作为尼罗罗非鱼和三种杂交鱼的分子标记；引物S_(326)和引物S_(336)可以作为奥利亚和三种杂交鱼的分子标记，可用来鉴定纯种和杂交鱼。
如何在外部形态上区分纯种和各种杂交鱼，在实际生产中有十分重要的意义。本研究表明：用聚类的方法可以将尼罗和杂交鱼、奥利亚和杂交鱼区分开来。尼罗和奥利亚在体形上有着较大的差别，三种杂交鱼在体形上更偏向于奥利亚。其中回交子代和兄妹交子代的体形最相似。
This study studied the genetic relations between three kinds of hybrid tilapia and oreochromics niloticus and O.aureus at the level of morphology and molecule. In order to detect the heterosis of three kinds of hybrid and to explain the difference of capability of production at the level of molecule we compared the between-population index of similarity and between-population genetic distance of three kinds of hybrid to oreochromics niloticus and O.aureus. At the same time we distinguished the hybrid from oreochromics niloticus and O.aureus.by new statistic technology and software in morphology and established new means of morphology. This study grounded for breeding and culturing of tilapia theoretically and practically.
The results of this experiment showed that there were great between-population
distance between oreochromics niloticus and O.aureus. which indicated the gene of
oreochromics niloticus ♀ × O.aureus ♂ was more heterozygous and showed the heterosis
in oreochromics niloticus ♀ × O.aureus ♂ . The between-population index of similarity
between oreochromics niloticus ♀ × O.aureus ♂ and oreochromics niloticus and
O.aureus were equal nearly which also certified the heterosis of oreochromics niloticus
♀ × O.aureus ♂ .there were high between-population index of similarity between
(oreochromics niloticus ♀ × O.aureus ♂ ) ♀ × O.aureus ♂ and O.aureus which
indicated that (oreochromics niloticus ♀ × O.aureus ♂ ) ♀ × O.aureus ♂ resembled
O.aureus in idioplasm.it was the same to (oreochromics niloticus ♀ × O.aureus ♂ ) ?×
(oreochromics niloticus ♀ × O.aureus ♂) ♂. the result of this experiment explained the
poor quality of (oreochromics niloticus ♀ × O.aureus ♂ ) ♀ × O.aureus ♂ and
(oreochromics niloticus ♀ × O.aureus ♂ ) ♀ × (oreochromics niloticus ♀ × O.aureus
♂ ) ♂ .for example, slow growth, poor food transformation etc. The results of this
study suggested that (oreochromics niloticus ♀ ×O.aureus ♂ ) ♀ × O.aureus ♂ and
(oreochromics niloticus ♀ × O.aureus ♂ ) ♀ × (oreochromics niloticus ♀ × O.aureus
♂ ) ♂ need more selective breeding and hybridization in order to improve the quality of
them.
In addition, products of 8323 were found to be molecular markers for identifying oreochromics niloticus and three kinds of hybrid tilapia.products of 8326 and S336 were found to be markers for identifying O.aureus and three kinds of hybrid tilapia.
It is great meaningful to distinguish pure strain tilapia and hybrid in morphology. The results of this study showed that we could distinguish hybrid tilapia from pure strain tilapia by clustering conducting on metric characters, three kinds of hybrid resembled O.aureus from the result of clustering conducting on metric characters.
本论文试从形态学和分子水平研究三种不同杂交方式的杂交罗非鱼后代与纯种尼罗罗非鱼及奥利亚罗非鱼之间的遗传关系。比较正交一代、回交子代、兄妹交子代三种杂交鱼与纯种尼罗罗非鱼和奥利亚罗非鱼之间的种内、种间相似系数、遗传距离。从分子水平检验杂交鱼是否具有杂交优势，从分子水平解释三种杂交鱼生产性能存在的差异。同时在形态学上利用最新的统计学技术和软件对纯种鱼和杂交鱼进行区别，并建立新的形态学分析方法，对杂交鱼和纯种鱼进行区别，为选育和培养高品质的罗非鱼养殖品种打下理论基础和提供实践方法。
结果表明：尼罗罗非鱼和奥利亚罗非鱼之间有着较大的遗传距离，使正交一代基因杂合性增加，使其更有可能获得杂交优势，且正交一代与父母本间的种间相似系数没有明显的亲本偏向性，更进一步证明了其正交一代的杂交优势。回交子代和兄妹交子代与奥利亚有着更大的种间相似系数，在种质上更偏向于奥利亚罗非鱼，由于奥利亚在生产中表现出生长速度较慢、饵料转化率较低。故解释了回交子代、兄妹交子代生产性能较差的原因。同时建议对回交子代、兄妹交子代需进一步的选育和杂交，以其获得更高程度的基因的杂合性和优良的性状。
筛选出引物S_(323)可以作为尼罗罗非鱼和三种杂交鱼的分子标记；引物S_(326)和引物S_(336)可以作为奥利亚和三种杂交鱼的分子标记，可用来鉴定纯种和杂交鱼。
如何在外部形态上区分纯种和各种杂交鱼，在实际生产中有十分重要的意义。本研究表明：用聚类的方法可以将尼罗和杂交鱼、奥利亚和杂交鱼区分开来。尼罗和奥利亚在体形上有着较大的差别，三种杂交鱼在体形上更偏向于奥利亚。其中回交子代和兄妹交子代的体形最相似。
This study studied the genetic relations between three kinds of hybrid tilapia and oreochromics niloticus and O.aureus at the level of morphology and molecule. In order to detect the heterosis of three kinds of hybrid and to explain the difference of capability of production at the level of molecule we compared the between-population index of similarity and between-population genetic distance of three kinds of hybrid to oreochromics niloticus and O.aureus. At the same time we distinguished the hybrid from oreochromics niloticus and O.aureus.by new statistic technology and software in morphology and established new means of morphology. This study grounded for breeding and culturing of tilapia theoretically and practically.
The results of this experiment showed that there were great between-population
distance between oreochromics niloticus and O.aureus. which indicated the gene of
oreochromics niloticus ♀ × O.aureus ♂ was more heterozygous and showed the heterosis
in oreochromics niloticus ♀ × O.aureus ♂ . The between-population index of similarity
between oreochromics niloticus ♀ × O.aureus ♂ and oreochromics niloticus and
O.aureus were equal nearly which also certified the heterosis of oreochromics niloticus
♀ × O.aureus ♂ .there were high between-population index of similarity between
(oreochromics niloticus ♀ × O.aureus ♂ ) ♀ × O.aureus ♂ and O.aureus which
indicated that (oreochromics niloticus ♀ × O.aureus ♂ ) ♀ × O.aureus ♂ resembled
O.aureus in idioplasm.it was the same to (oreochromics niloticus ♀ × O.aureus ♂ ) ?×
(oreochromics niloticus ♀ × O.aureus ♂) ♂. the result of this experiment explained the
poor quality of (oreochromics niloticus ♀ × O.aureus ♂ ) ♀ × O.aureus ♂ and
(oreochromics niloticus ♀ × O.aureus ♂ ) ♀ × (oreochromics niloticus ♀ × O.aureus
♂ ) ♂ .for example, slow growth, poor food transformation etc. The results of this
study suggested that (oreochromics niloticus ♀ ×O.aureus ♂ ) ♀ × O.aureus ♂ and
(oreochromics niloticus ♀ × O.aureus ♂ ) ♀ × (oreochromics niloticus ♀ × O.aureus
♂ ) ♂ need more selective breeding and hybridization in order to improve the quality of
them.
In addition, products of 8323 were found to be molecular markers for identifying oreochromics niloticus and three kinds of hybrid tilapia.products of 8326 and S336 were found to be markers for identifying O.aureus and three kinds of hybrid tilapia.
It is great meaningful to distinguish pure strain tilapia and hybrid in morphology. The results of this study showed that we could distinguish hybrid tilapia from pure strain tilapia by clustering conducting on metric characters, three kinds of hybrid resembled O.aureus from the result of clustering conducting on metric characters.
引文
1、李思发，蔡完其．我国尼罗罗非鱼和奥利亚罗非鱼养殖群体的遗传渐渗．水产学报，1995，19(2)：106～110．
2、淡水良种鱼类增养殖技术 北京：农业出版社，1995，265．
3、赖秋明 罗非鱼鱼苗群体性比测定方法研究．
4、朱泽进，李思发．不同亲本组合奥尼杂交鱼子代雄性率、生长率及早期成活率的研究水产科技情报，2000，27(2)：54～58．
5、夏德全，曹莹．罗非鱼杂交F1代与亲本的遗传关系及其杂种优势的利用 中国水产科学，1999，6(4)：29～32．
6、李家乐 李思发．尼奥鱼[尼罗罗非鱼(♀)X奥利亚罗非鱼(♂)]同其亲本的形态和判别 水产学报，1999，23(3)：261～265．
7、夏德全 2000中国罗非鱼养殖现状及发展前景。科学养鱼2000(1)
8、潘炯华 1989罗非鱼的养殖。广东科技出版社。
9、中国渔业统计年鉴1984-2000。农业部渔业局编。
10、李思发 1993主要养殖鱼类种质资源研究进展。水产学报 17(4)：344-358。
11、陈素芝等 1994我国引进的罗非鱼类的初步研究动物学杂志 29(3)：18-23。
12 蔡仁逵等 1991名特优水产品养殖实用技术。中央广播电视大学出版社。
13、吴婷婷等 1993鳜鱼和奥利亚罗非鱼乳酸脱氢酶谱的比较。水产学报17(4)：325-329。
14、杨兴棋等 1984几种罗非鱼乳酸脱氢酶的电泳研究。遗传学报11(2)：123-140。
15、李志忠 1997奥利亚罗非鱼和尼罗罗非鱼同工酶的研究南京农业大学硕土论文。
16、郑光明等 1995鱼类线粒体DNA的研究及其应用。中国水产科学 2(5)：159-163。
17、夏德全等 1997两种不同地方种群尼罗罗非鱼遗传差异和分子遗传标记。中国水产科学 4(3)：7-12。
18、曹萤等 1997尼罗非鲫和奥利亚罗非鱼非鲫线粒体DNA遗传差异性研究。水产学报 21(4)：360-365。
19、王文君 1996奥利亚罗非鱼和尼罗罗非鱼类线粒体DNA遗传差异性研究。南京农业大学硕士论文。
20、王天奇 1997随机多态性DNA法分析奥利亚罗非鱼和尼罗罗非鱼基因组多态性的研究。南京农业大学硕士论文。
21、王希道 2000不同鳗鱼种群遗传结构和种质鉴定分子遗传标记研究。南京农业大学硕士论文。
22、王慧等 1997 RAPD及其在动物育种中的应用。山东农业大学学报 28(1)：79-82。
23、刘春林等 1998利用RAPD标记技术对远缘杂交后代的分子遗传学研究。湖南农业科学 1：22-23。
24、夏德全等 1999用RAPD分析对罗非鱼遗传变异的研究及其对杂种优势的应用。内蒙古农牧学院学报 20：27-31。
25、龙继蓉 1999 RAPD技术及其在动物遗传育种中应用及前景。四川畜牧兽医5。
26、贾永红 1998遗传标记及其在动物遗传育种中的应用(上)。黄牛杂志 24(6)：45-46。
27、贾永红 1998遗传标记及其在动物遗传育种中的应用(下)。黄牛杂志 25(1)：50-55。
28、李春丽 1997应用分子标记差异性预测作物杂种优势的研究进展。遗传19(1)：46-48。
29、邱高峰 1996分析基因组DNA多态性RAPD技术在水产生物技术研究中的应用前景。上海水产大学学报 2(5)：101-108。
30、马中军等 1997动物RFLP分子遗传标记及其应用。甘肃畜牧兽医 2(27)：27-29。
31、赵君等 1999几种常用的分子标记的比较。内蒙古农业科技 3：32-33。
32、J.萨姆布鲁克等分子克隆指南(第二版)[M] 金冬雁等译 北京：科学出版社 1996 464-467。
33、李思发等 1998尼罗罗非鱼品系间形态差异研究。动物学报 44(4)：450-457。
34、Artalion, R.R. 1982 Genetic markers in Sarothrrodon and their use for sex and species identification In: The Biology and Culture of Tilapias (ed.) by R. S. V. Pullin et al. ICLARM, manila, Philippines.
35、Capili, J. B. et al. 1990 Isozyme and Mitochondrial DNA Restriction Endonuclease Analysis and Subspecies Identification. Heredity 73: 117-123.
36、Chen,F. Y. et al. 1970 Zone electrophoresis studies on the proteins of Tilapia mossambicus and Thornorum and their F1 hybrids, T. zillii and T. melanopleura J. Fish Res. Board can., 27:2167-2177.
37、Cruz, T. a. et al. 1982 Enzyme electrophoresis in Tilapia ziuii: a pattern for determining biochemical genetic markers for use in Tilapia stock identification. Aquaculture。29:311-329.
38、FAO Fisheries Statistics 1997 Food and Agriculture Organization of the United Nations, Rome.
39、Herzberg, A. 1978 Electrophoretic esterase patterns of the surface mucus for the identification of Tilapia species Aquaculture 13:81-83.
40、OJen-leih Wu, et al. 1983 Esterase Isozymes in Oreochromis niloticus, O. aureus, O. mossambicus and Red Tilapia In: International symposium on Tilapia in Aquaculture, Proceedings, Nazarety, Isreal.
41、Kornfield, I. 1984 Descriptive genetics of cichild fishes In: B. J. Truner(Editor), Evolutionary Genetics of Fishes, Plenum Press, New York, NY, PP. 590-616.
42、Koifield, I. 1991 Genetics In: M. Keenleyside(Editor), Cichild Fishes: Behavior, Biology and Evolution Champman and Hall, London, PP: 590-616.
43、Koifield, I. Et al. 1979 Biochemical and cytological differentiation among cichild fishes of sea of Galilee Evolution 33(1):1-14
44、Lynch M. 1991 Analysis of population genetic structure by DNA fingerprinting. In: DNA fingerprinting Approaches and applications (ed. By T. Burke, G. Dolf, A. J. Jeffreys & R. Wolf), pp:113-126. Basel.
45、Macaranas, J. M. et al. 1986 Electrophoretic evidence for extensive hybrid gene introgression into
commercial O.niliticus stocks in the Philippines Aqua. Fish Manag. 17: 249-268．
46、 Moav, R. Et al. 1976 Applications of electrophoretic markers to Fish breeding. In: Advantages and methods Aquaculture, 9:217-228．
47、 McAndrew, B. J. 1981 Tilapia stock identification using electrophoretic markers, Aquaculture 30:249-261．
48、 Mckaye, K. R. et al. 1984 Genetics evidence for allopatric differentiation among color morphs of a lake Malawi cichild fish Evolution 38: 215-219．
49、 McAndrew, B. J. et al. 1984 Evolutionary relationships within three Tilapia genera (Pisces: Cichildae) Zool. J. Linn: Soc. 80:421-434．
50、 Naish, K. A. et al. 1995 Multi-locus DNA fingerprinting and RAPD reveal similar genetic relationships between strains of Oreochromis niloticus (Pisces:Cichlidae) Mol. Ecol. 4:271-274．
51、 Oberst, S. et al. 1996 Biochemical and immunological markers for the discrimination of three tilapia species: T.zillii Gervais, T.guineemis Bleeker and T.daget, Thys v.d. Audenaede(pisces:cichlidae) from west Africa Aquaculture Research 27:235-244．
52、 Pullin, R. S.V. (ed.) 1988 Tilapia genetic resources for Aquaculture, 108, ICLARM.
53、 Pullin, R. S. V. 1988 Genetics improvement of Tilapias: problems and prospects In: Pullin R. S. V. et al. (ed.) The second international Symposium on Tilapia in aquaculture . ICLARM conference proceeding 15, Manila Philippines .
54、 Pullin 1982 General Discussion on the Biology and Culture of Tilapias,In: The Biology and culture of Tilapias(ed,)by R.S.V.pullin et al.ICLARM,manila,Philippines.
55、 Philippart, J. C. L. et al. 1982 Ecology and Distribution of Tilapia,In:The Biology and culture of Tilapias (ed.) by R. S. V. Pullin et al. ICLARM, manila, Philippins.
56、 Rognon, X. et al. 1996 Allozyme variation in natural and cultured populations in two Tilapia Species: Oreochromis .niloticus and Tilapia.zillii Heredity 76:640-650．
57、 Seyoum, S. 1989 Stock identification and evolutionary relationships of the tilapiine fishes of the genera Oreochromis, Sarothrodon and Tilapia (Pisces:Cichildae) using allozyme analysis and restriction endonuclease analysis of mitochonrial DNA PhD. Thesis, University of Waterloo, Waterloo, Ont., Canada.
58、 Seyoum, S. 1990 Allozyme variation in Subspecies of Oreochromis, niloticus Isozyme Bull. 23:97．
59、 Sage, R. D. et al. 1975 Tropic radiation through polymorphism in cichild fishes Proc. Natl. Acad. Sci. U.S.A. 72:4669-4673．
60、 Sage, R. D. et al. 1984 Molecular versus morphological change among cichild fishes (Pisces:Cichildae) of Lake Victoria. In: A. A. Echelle and I. Komfield (Editors) Evolution in Fish Species Flocks, University of Maine, OronoPress, Orono, ME, pp. 185-202．
61、 Seyoum, S. et al. 1992 Identification of the subspecies of Oreochromis niloticus (Pisces:Cichildae) using restriction endonuclease analysis of mitochondrial DNA. Aquaculture 102: 29-42．
62、 Sodsuk, P. et al. 1992 Molecular systematics of three tilapiine genera Oreochromis, Sarothrodon and Tilapia using allozyme data J. Fish Biol. 39: 301-308．
63、 Williames, J. G. K. et al. 1990 DNA polymorphisms amplified by arbitrary primers are useful as genetic markers [J]. Nucleic acids Res. 18:6531-6535．
64、 Lynch, M. 1990 The similarity index and DNA fingerprinting. Mol. Biol. Evol. 7: 478-484．
65、 Trewavas, E. 1982 Tilapiine Fishes of the Genera Sarotherodon, Oreochromis and Danakilia. British Museum(Natural History),London.
66、 Artalion,R,R.Genetic markers in Sarothrrodon and their use for sex and speciess identification ln:The Biology and Culture of Tilapias (ed)by R.S.V.Pullin et al.iclarm,manila,Philippines.
67、 J.M.Myers et al.Iinduction of diploid androgenetic and mitotic gynogenetic Nile tilapia (oreochromis niloticus L.) .Ther Appl Genet (1995) 90: 205-210．
68、 Graham C.Mair .Chromosone-set manipulation in tilapia-techniques,problems and prspects.Aquaculture,Ⅲ (1993) 227-244．
69、 Yehheskel,O.and Avtalion,R.R,1988． Artificial fertilization of tilapia eggs,a preliminary study.Colloq.INRA,44: 169-175．
2、淡水良种鱼类增养殖技术 北京：农业出版社，1995，265．
3、赖秋明 罗非鱼鱼苗群体性比测定方法研究．
4、朱泽进，李思发．不同亲本组合奥尼杂交鱼子代雄性率、生长率及早期成活率的研究水产科技情报，2000，27(2)：54～58．
5、夏德全，曹莹．罗非鱼杂交F1代与亲本的遗传关系及其杂种优势的利用 中国水产科学，1999，6(4)：29～32．
6、李家乐 李思发．尼奥鱼[尼罗罗非鱼(♀)X奥利亚罗非鱼(♂)]同其亲本的形态和判别 水产学报，1999，23(3)：261～265．
7、夏德全 2000中国罗非鱼养殖现状及发展前景。科学养鱼2000(1)
8、潘炯华 1989罗非鱼的养殖。广东科技出版社。
9、中国渔业统计年鉴1984-2000。农业部渔业局编。
10、李思发 1993主要养殖鱼类种质资源研究进展。水产学报 17(4)：344-358。
11、陈素芝等 1994我国引进的罗非鱼类的初步研究动物学杂志 29(3)：18-23。
12 蔡仁逵等 1991名特优水产品养殖实用技术。中央广播电视大学出版社。
13、吴婷婷等 1993鳜鱼和奥利亚罗非鱼乳酸脱氢酶谱的比较。水产学报17(4)：325-329。
14、杨兴棋等 1984几种罗非鱼乳酸脱氢酶的电泳研究。遗传学报11(2)：123-140。
15、李志忠 1997奥利亚罗非鱼和尼罗罗非鱼同工酶的研究南京农业大学硕土论文。
16、郑光明等 1995鱼类线粒体DNA的研究及其应用。中国水产科学 2(5)：159-163。
17、夏德全等 1997两种不同地方种群尼罗罗非鱼遗传差异和分子遗传标记。中国水产科学 4(3)：7-12。
18、曹萤等 1997尼罗非鲫和奥利亚罗非鱼非鲫线粒体DNA遗传差异性研究。水产学报 21(4)：360-365。
19、王文君 1996奥利亚罗非鱼和尼罗罗非鱼类线粒体DNA遗传差异性研究。南京农业大学硕士论文。
20、王天奇 1997随机多态性DNA法分析奥利亚罗非鱼和尼罗罗非鱼基因组多态性的研究。南京农业大学硕士论文。
21、王希道 2000不同鳗鱼种群遗传结构和种质鉴定分子遗传标记研究。南京农业大学硕士论文。
22、王慧等 1997 RAPD及其在动物育种中的应用。山东农业大学学报 28(1)：79-82。
23、刘春林等 1998利用RAPD标记技术对远缘杂交后代的分子遗传学研究。湖南农业科学 1：22-23。
24、夏德全等 1999用RAPD分析对罗非鱼遗传变异的研究及其对杂种优势的应用。内蒙古农牧学院学报 20：27-31。
25、龙继蓉 1999 RAPD技术及其在动物遗传育种中应用及前景。四川畜牧兽医5。
26、贾永红 1998遗传标记及其在动物遗传育种中的应用(上)。黄牛杂志 24(6)：45-46。
27、贾永红 1998遗传标记及其在动物遗传育种中的应用(下)。黄牛杂志 25(1)：50-55。
28、李春丽 1997应用分子标记差异性预测作物杂种优势的研究进展。遗传19(1)：46-48。
29、邱高峰 1996分析基因组DNA多态性RAPD技术在水产生物技术研究中的应用前景。上海水产大学学报 2(5)：101-108。
30、马中军等 1997动物RFLP分子遗传标记及其应用。甘肃畜牧兽医 2(27)：27-29。
31、赵君等 1999几种常用的分子标记的比较。内蒙古农业科技 3：32-33。
32、J.萨姆布鲁克等分子克隆指南(第二版)[M] 金冬雁等译 北京：科学出版社 1996 464-467。
33、李思发等 1998尼罗罗非鱼品系间形态差异研究。动物学报 44(4)：450-457。
34、Artalion, R.R. 1982 Genetic markers in Sarothrrodon and their use for sex and species identification In: The Biology and Culture of Tilapias (ed.) by R. S. V. Pullin et al. ICLARM, manila, Philippines.
35、Capili, J. B. et al. 1990 Isozyme and Mitochondrial DNA Restriction Endonuclease Analysis and Subspecies Identification. Heredity 73: 117-123.
36、Chen,F. Y. et al. 1970 Zone electrophoresis studies on the proteins of Tilapia mossambicus and Thornorum and their F1 hybrids, T. zillii and T. melanopleura J. Fish Res. Board can., 27:2167-2177.
37、Cruz, T. a. et al. 1982 Enzyme electrophoresis in Tilapia ziuii: a pattern for determining biochemical genetic markers for use in Tilapia stock identification. Aquaculture。29:311-329.
38、FAO Fisheries Statistics 1997 Food and Agriculture Organization of the United Nations, Rome.
39、Herzberg, A. 1978 Electrophoretic esterase patterns of the surface mucus for the identification of Tilapia species Aquaculture 13:81-83.
40、OJen-leih Wu, et al. 1983 Esterase Isozymes in Oreochromis niloticus, O. aureus, O. mossambicus and Red Tilapia In: International symposium on Tilapia in Aquaculture, Proceedings, Nazarety, Isreal.
41、Kornfield, I. 1984 Descriptive genetics of cichild fishes In: B. J. Truner(Editor), Evolutionary Genetics of Fishes, Plenum Press, New York, NY, PP. 590-616.
42、Koifield, I. 1991 Genetics In: M. Keenleyside(Editor), Cichild Fishes: Behavior, Biology and Evolution Champman and Hall, London, PP: 590-616.
43、Koifield, I. Et al. 1979 Biochemical and cytological differentiation among cichild fishes of sea of Galilee Evolution 33(1):1-14
44、Lynch M. 1991 Analysis of population genetic structure by DNA fingerprinting. In: DNA fingerprinting Approaches and applications (ed. By T. Burke, G. Dolf, A. J. Jeffreys & R. Wolf), pp:113-126. Basel.
45、Macaranas, J. M. et al. 1986 Electrophoretic evidence for extensive hybrid gene introgression into
commercial O.niliticus stocks in the Philippines Aqua. Fish Manag. 17: 249-268．
46、 Moav, R. Et al. 1976 Applications of electrophoretic markers to Fish breeding. In: Advantages and methods Aquaculture, 9:217-228．
47、 McAndrew, B. J. 1981 Tilapia stock identification using electrophoretic markers, Aquaculture 30:249-261．
48、 Mckaye, K. R. et al. 1984 Genetics evidence for allopatric differentiation among color morphs of a lake Malawi cichild fish Evolution 38: 215-219．
49、 McAndrew, B. J. et al. 1984 Evolutionary relationships within three Tilapia genera (Pisces: Cichildae) Zool. J. Linn: Soc. 80:421-434．
50、 Naish, K. A. et al. 1995 Multi-locus DNA fingerprinting and RAPD reveal similar genetic relationships between strains of Oreochromis niloticus (Pisces:Cichlidae) Mol. Ecol. 4:271-274．
51、 Oberst, S. et al. 1996 Biochemical and immunological markers for the discrimination of three tilapia species: T.zillii Gervais, T.guineemis Bleeker and T.daget, Thys v.d. Audenaede(pisces:cichlidae) from west Africa Aquaculture Research 27:235-244．
52、 Pullin, R. S.V. (ed.) 1988 Tilapia genetic resources for Aquaculture, 108, ICLARM.
53、 Pullin, R. S. V. 1988 Genetics improvement of Tilapias: problems and prospects In: Pullin R. S. V. et al. (ed.) The second international Symposium on Tilapia in aquaculture . ICLARM conference proceeding 15, Manila Philippines .
54、 Pullin 1982 General Discussion on the Biology and Culture of Tilapias,In: The Biology and culture of Tilapias(ed,)by R.S.V.pullin et al.ICLARM,manila,Philippines.
55、 Philippart, J. C. L. et al. 1982 Ecology and Distribution of Tilapia,In:The Biology and culture of Tilapias (ed.) by R. S. V. Pullin et al. ICLARM, manila, Philippins.
56、 Rognon, X. et al. 1996 Allozyme variation in natural and cultured populations in two Tilapia Species: Oreochromis .niloticus and Tilapia.zillii Heredity 76:640-650．
57、 Seyoum, S. 1989 Stock identification and evolutionary relationships of the tilapiine fishes of the genera Oreochromis, Sarothrodon and Tilapia (Pisces:Cichildae) using allozyme analysis and restriction endonuclease analysis of mitochonrial DNA PhD. Thesis, University of Waterloo, Waterloo, Ont., Canada.
58、 Seyoum, S. 1990 Allozyme variation in Subspecies of Oreochromis, niloticus Isozyme Bull. 23:97．
59、 Sage, R. D. et al. 1975 Tropic radiation through polymorphism in cichild fishes Proc. Natl. Acad. Sci. U.S.A. 72:4669-4673．
60、 Sage, R. D. et al. 1984 Molecular versus morphological change among cichild fishes (Pisces:Cichildae) of Lake Victoria. In: A. A. Echelle and I. Komfield (Editors) Evolution in Fish Species Flocks, University of Maine, OronoPress, Orono, ME, pp. 185-202．
61、 Seyoum, S. et al. 1992 Identification of the subspecies of Oreochromis niloticus (Pisces:Cichildae) using restriction endonuclease analysis of mitochondrial DNA. Aquaculture 102: 29-42．
62、 Sodsuk, P. et al. 1992 Molecular systematics of three tilapiine genera Oreochromis, Sarothrodon and Tilapia using allozyme data J. Fish Biol. 39: 301-308．
63、 Williames, J. G. K. et al. 1990 DNA polymorphisms amplified by arbitrary primers are useful as genetic markers [J]. Nucleic acids Res. 18:6531-6535．
64、 Lynch, M. 1990 The similarity index and DNA fingerprinting. Mol. Biol. Evol. 7: 478-484．
65、 Trewavas, E. 1982 Tilapiine Fishes of the Genera Sarotherodon, Oreochromis and Danakilia. British Museum(Natural History),London.
66、 Artalion,R,R.Genetic markers in Sarothrrodon and their use for sex and speciess identification ln:The Biology and Culture of Tilapias (ed)by R.S.V.Pullin et al.iclarm,manila,Philippines.
67、 J.M.Myers et al.Iinduction of diploid androgenetic and mitotic gynogenetic Nile tilapia (oreochromis niloticus L.) .Ther Appl Genet (1995) 90: 205-210．
68、 Graham C.Mair .Chromosone-set manipulation in tilapia-techniques,problems and prspects.Aquaculture,Ⅲ (1993) 227-244．
69、 Yehheskel,O.and Avtalion,R.R,1988． Artificial fertilization of tilapia eggs,a preliminary study.Colloq.INRA,44: 169-175．