Further Evidence of Linkage at the tva and tvc Loci in the Layer Lines and a Possibility of Polyallelism at the tvc Locus

  • Ghosh, A.K. (Indian Veterinary Research Institute) ;
  • Pani, P.K. (Central Avian Research Institute)
  • Received : 2004.08.17
  • Accepted : 2004.12.18
  • Published : 2005.05.01


Three lines of White Leghorn (WL) chickens (IWJ, IWG and IWC) maintained at Central Avian Research Institute, Izatnagar (UP), were used for chorioallantoic membrane (CAM) and liver tumour (LT) assay. Eleven-day-old embryos of each line were partitioned into three groups and inoculated with 0.2 ml of subgroup A, subgroup C and an equal mixture of subgroup A and C Rous sarcoma virus (RSV). Subgroup virus receptor on the cell surface membrane for subgroup A is coded for by tumour virus a (tva) locus and for subgroup C by tumour virus c (tvc) locus. The random association of the genes at the tva and tvc loci in IWJ and IWC line was assessed and the $x^2$-values for phenotypic classes were found to be significant, indicating the linkage between the tva and tvc loci. The linkage value was estimated to be 0.09 on pooled sex and pooled line basis. On the basis of four subclass tumour phenotypes a 4-allele model was proposed for tva locus having $a^{s1}$, $a^{s2}$, $a^{r1}$ and $a^{r2}$ alleles and the frequencies were calculated as 0.47, 0.13, 0.13 and 0.27 for IWJ line, 0.31, 0.33, 0.14 and 0.22 for IWG line and 0.44, 0.11, 0.21 and 0.24 for IWC line, respectively. Similarly, for tvc locus the frequencies of four alleles i.e. $c^{s1}$, $c^{s2}$, $c^{r1}$ and $c^{r2}$ were calculated as 0.42, 0.20, 0.21 and 0.17 for IWJ line, 0.42, 0.17, 0.27 and 0.14 for IWG line and 0.30, 0.21, 0.16 and 0.33 for IWC line, respectively. The $x^2$-values for all classes of observations were not significant (p>0.05), indicating a good fit to the 4-allele model for the occurrence of 4-subclass tumour phenotypes for tva and tvc loci. On the basis of the 2-allele model both tva and tvc locus carries three genotypes each. But, on the basis of the 4-allele model tva and tvc locus carries 10 genotypes each. The interaction between A-resistance and C-resistance (both CAM and LT death) was ascertained by taking the 10 genotypes of tva locus and 3 genotypes of tvc locus by pooling the lines and partitioning the observations into 3 classes. The $x^2$-values for the genotypic classes of CAM (-) LT (+) and CAM (-) LT (-) phenotypes to mixed virus (A+C) infection were found to be highly significant (p<0.01), indicating increased resistance, which indicates the joint segregation of $a^r$ and $c^r$ genes, suggesting the existence of close linkage between the tva and tvc loci. Therefore, an indirect selection approach using subgroup C viruses can be employed to generate stocks resistant to subgroup A LLV, obviating contamination with the most common agent causing LL in field condition.


Chorioallantoic Membrane;Liver Tumour;Linkage;Polyallelism;Tumour Virus


  1. Ghosh, A. K. and P. K. Pani. 2002. Induced liver tumour further support to a genetic marker with its high correlation with chorioallantoic membrane phenotypes in selected layer lines. Indian J. Exp. Biol. 40:144-150.
  2. Pani, P. K. and S. Naithani. 1990. Inheritance of fibrosarcomatous liver tumours in chorioallantoic membrane inoculated WL chicks by retrovirus subgroup A (BS-RSV), a 4-allele genetic model. Avian Pathol. 19:669-685.
  3. Pani, P. K. and S. Naithani. 1991. Genetics of post-hatching survival potential of Australorp chicks infected as embryos by subgroup A Rous sarcoma virus: further support to 4-allele model. Indian J. Exp. Biol. 29:416-421.
  4. Snedecor, W. G. and G. W. Cochran. 1967. Statistical methods. Sixth Edition. The Iowa State University Press, Iowa, (USA).
  5. Dren, C. N. and P. K. Pani. 1977. Genetic control of resistance of subgroup A and subgroup C tumour viruses in Rhode Island Red fowl, evidence for linkage between the tumour virus a (tva) and tumour virus c (tvc) loci. J. Gen. Virol. 35:13-23.
  6. Johari, D. C., M. C. Kataria, D. Sharma, D. P. Singh, R. Gopal and B. B. Dash. 1992. Improvement of poultry for egg production. Annu. Sci. Rep. CARI, Izatnagar. pp. 7-13.
  7. Falconer, D. S. and T. F. C. Mackay. 1989. Introduction to Quantitative Genetics. Third Edition. Longman Scientific and Technical, Essex, England.
  8. Dougherty, R. M., J. A. Stewart and A. R. Morgan. 1960. Quantitative studies of the relationship between infecting dose of Rous sarcoma virus, antiviral immune response and tumour growth in chickens. Virol. 11:349-370.
  9. Pani, P. K., S. Kishore and S. Naithani. 1988. Hatchability and mortality following infection of chick embryos with subgroup A Rous sarcoma virus. Avian Pathol. 17:533-555.
  10. Pani, P. K. 1976. Genetics of resistance of fowl to infection by RNA tumour viruses. Proc. Royal Soc. Med. 69:43-48.
  11. Payne, L. N. and P. K. Pani. 1971. Evidence for linkage between genetic loci controlling response of fowl to subgroup A and subgroup C sarcoma viruses. J. Gen. Virol. 13:253-259.
  12. Payne, L. N. 1985. Interaction between host genome and avian RNA tumour viruses. In: RNA viruses and host genome in oncogenesis (Ed. P. Emmelot and P. Bentvelgen). North Holand Publishing Company, Amsterdom. pp. 1-16.
  13. Pani, P. K. 1974. Close linkage between genetic loci controlling response of fowl to subgroup A and subgroup C sarcoma viruses. J. Gen. Virol. 22:187-195.