- Email: [email protected]
Allelic Frequencies in Eight Alloantigen Systems of Chickens Selected for High and Low Antibody Response to Sheep Red Blood Cells
Allelic Frequencies in Eight Alloantigen Systems of Chickens Selected for High and Low Antibody Response to Sheep Red Blood Cells E. A. DUNNINGTON,1 R. W. BRILES, 2 W. E. BRILES, 2 W. B. GROSS, 1 and P. B. SIEGEL 1 Poultry Science Department, Virginia Polytechnic Institute and State University, Blacksburg, Virginia 24061 and Department of Biological Sciences, Northern Illinois University, Dekalb, Illinois 60115
ABSTRACT Parents and offspring from lines of chickens selected for high (HA) and low (LA) antibody response to sheep red blood cell antigen(s) were blood typed for systems A, B, C, D, E, H, I, and L. All birds were homozygous for allele L2, and allelic frequencies for the / system were essentially identical in both lines. In contrast, allelic frequencies of the other blood group systems differed rather markedly between the HA and LA lines. (Key words: alleles, alloantigens, chickens, antibody, selection) 1984 Poultry Science 63:1470-1472 INTRODUCTION
Studies of antigen recognition in antibody production of the fowl have generally focused on effects of specific loci and responses to infectious agents (Gavora and Spencer, 1983; van der Zijpp, 1983). Responses to divergent selection for antibody production to erythrocyte antigens have been demonstrated in mice (Biozzi et al, 1982), guinea pigs (Ibanez et al, 1980), and chickens (Siegel et al, 1982). Such selection has resulted in correlated responses in resistance to infectious agents, growth, and reproductive traits (Biozzi et al, 1982; Siegel et al, 1982). Presented here are comparisons of allelic frequencies for specific blood group systems in lines of chickens selected for high (HA) and low (LA) antibody response to the antigens of sheep red blood cells (SRBC). MATERIALS AND METHODS
Beginning with a sample of the Cornell randombred population (Lowe and Garwood, 1974), selection for high and low antibody response to antigens of SRBC was used to develop lines HA and LA, respectively (Siegel et al, 1982). Individual selection within each line
1
Virginia Polytechnic Institute and State University. 2 Northern Illinois University.
was based solely on antibody titers 5 days postchallenge to a .1 ml intravenous injection of a .25% suspension of SRBC. Parents (aged 386 to 407 days) from selected Generation 9 and their offspring were blood typed for Systems A, B, C, D, E, H, I, and L. Number of tested sires, dams, and offspring were 8, 21, and 141 in Line HA and 8, 19, and 138 in Line LA, respectively. Approximately .5 ml of blood was collected directly into a plastic test tube containing 2.0 ml of chilled isotonic solution of sodium citrate (2%) and sodium chloride (.42%) as anticoagulant. Tubes were coded, packed in ice, and immediately shipped from Blacksburg to DeKalb for blood typing. Reagents specific for alloantigens of each system were selected mostly on the basis of previous experience with the Cornell randombred or lines derived from that population (Briles et al, 1976, 1980; Briles and Briles, 1982). An allele of the A system not previously encountered in Cornell randombred-derived lines (Briles et al, 1977) was temporarily designated Ax. RESULTS Frequencies (%) of alleles and of homozygotes for each blood group system were similar for males and females; therefore, data were pooled (Table 1). All blood-typed individuals appeared to be homozygous for L 2 , as the L 1 allele which exists in other populations, was absent. Frequencies of alleles and homo-
1470
Downloaded from http://ps.oxfordjournals.org/ at North Dakota State University on May 25, 2015
(Received for publication March 19, 1984)
1471
RESEARCH NOTE TABLE 1. Frequencies (%) of genes and homozygotes at each locus in Generation 10 in lines selected for high (HA) and low (LA) antibody response Blood group system Allele
A
1 2 3 4 5 13 21 31 X
19/0 1 20/4
B
C
E
D
H
/
L
0/26 100/74
32/33
100/100
Gene frequencies 29/51 40/5
58/96
9/0
74/88 26/12
71/39 0/10
68/67
51/95
5/0 3/0
Homozygotes 11/28
3/0 6/0
1 2 3 4 5 13 21 31
11/0
52/83
0/5
5/0
100/53
9/11
100/100
53/17 44/97
0/0
44/45
0/2
25/89 2/98 61/0 0/0 1
(HA/LA).
zygotes for the / system were essentially identical in both lines. The HA line was 100% homozygous for allele H2. In Line LA, however, frequency of H2 was 74% with 5 3% of the chickens homozygous; frequency of H1 was 26% with 5% homozygotes. For the B system, 99% of the individuals were B 1 3 and 1% B 2 1 in Line LA, while in HA, frequencies were 15, 80, and 5 for B 1 3 , B21, and B 3 1 , respectively. Ninety-five percent of the C system alleles were C 5 in Line LA, while in Line HA 51% of the alleles were Cs. Conversely, LA chickens had only 5% allele C2 and 0% allele C 4 compared to 40% and 9% in the HA line. For the D system, frequencies for O 1 were 29 and 51%, D3 were 71 and 39%, and D 4 were 0 and 10% for Lines HA and LA, respectively. Systems A and Et which are linked, also showed differences between the selected lines; alleleic frequencies are given separately for each locus. Frequencies of alleles A1, A2, and ^44 in the HA line were 19, 20, and 58%, respectively, whereas in Line LA the corresponding frequencies were 0, 4, and 96%, respectively. Allelic frequencies of E1 and E2 were 74 and 26% in the HA line and 88 and 12% in LA line. Frequencies of segregating A-E haplotype combinations (Table 2) show
that Line LA chickens homozygous for E1 were also homozygous for A4. The results show considerable changes in allelic frequencies for several blood group systems concomitant to divergent selection for antibody response to SRBC. Although symmetry of response and drift effects are not known because allelic frequencies were not measured in the base generation, the data provide a basis for designing genotype-comparison experiments to evaluate effects of those systems exhibiting allelic segregation. This reasoning is consistent with the observations of
TABLE 2. Frequency of A-E haplotype combinations in lines selected for high (HA) and low (LA) antibody response Haplotype
HA
LA
A'-E 1 A 2 -E 2 A 4 -E' A 4 -E 2 A x -E' A X -E J
16 21 56 5 2 1
0 3 89 8 0 0
Downloaded from http://ps.oxfordjournals.org/ at North Dakota State University on May 25, 2015
15/99 80/1
1472
DUNNINGTON ET AL.
Benedict et al. (1975) that one homozygous B genotype failed to initiate response to a synthetic polypeptide whereas two other homozygotes and the three possible heterozygous genotypes responded fully. Our data also show for the A, B, C, and E systems unusually high frequencies of homozygotes in Line LA and a more normal frequency of heterozygotes in HA, suggesting evaluation of alleles in both heterozygous and homozygous states for antibody response to the SRBC antigens.
Downloaded from http://ps.oxfordjournals.org/ at North Dakota State University on May 25, 2015
REFERENCES Benedict, A. A., L. W. Pollard, P. R. Morrow, H. A. Abplanalp, P. H. Maurer, and W. E. Briles, 1975. Genetic control of immune responses in chickens. Immunogenetics 2:313—324. Biozzi, G., D. Mouton, A. M. Heumann, and Y. Bouthillier, 1982. Genetic regulation of immunoresponsiveness in relation to resistance against infectious diseases. 2nd World Congr. Genet. Appl. Livest. Prod. 5:150-163. Briles, W. E., and R. W. Briles, 1982. Identification of haplotypes to the chicken major histocompatibility complex (B). Immunogenetics 15: 441-447. Briles, W. E., R. W. Briles, W. H. McGibbon, V. A. Garwood, and P. C. Lowe, 1977. Erythrocyte alloantigens in Regional Cornell Randombred
stock. Poultry Sci. 56:1698. (Abstr.) Briles, W. E., R. W. Briles, W. H. McGibbon, and H. A. Stone, 1980. Identification of B alloalleles association with resistance to Marek's disease, Pages 395—413 in Resistance and Immunity to Marek's Disease. P. B. Biggs, ed. Coram. Eur. Commun., Directorate-Gen. Sci. Tech. Info., Luxembourg. Briles, W. E., W. H. McGibbon, and H. A. Stone, 1976. Effects of B alloalleles from Regional Cornell Randombred stock on mortality from Marek's disease. Poultry Sci. 55:2011-2012. Gavora, J. S., and J. L. Spencer, 1983. Breeding for immune responsiveness and disease resistance. Anim. Blood Groups Biochem. Genet. 14: 159-180. Ibanez, O. M., M. S. Reis, M. Gennari, V.C.A. Ferreira, O. A. Sant' Anna, M. Siqueira, and G. Biozzi, 1980. Selective breeding of high and low antibody responder lines of guinea pigs. Immunogenetics 10:283-293. Lowe, P. C , and V. A. Garwood, 1974. Inbreeding in 16 generations of the Regional Cornell Control populations of chickens. Poultry Sci. 53:514— 517. Siegel, P. B., W. B. Gross, and J. A. Cherry, 1982. Correlated responses of chickens to selection for production of antibodies to sheep erythrocytes. Anim. Blood Groups Biochem. Genet. 13: 291-297. van der Zijpp, A. J., 1983. Breeding for immune responsiveness and disease resistance. World's Poult. Sci. J. 39:118-131.
ABSTRACT Parents and offspring from lines of chickens selected for high (HA) and low (LA) antibody response to sheep red blood cell antigen(s) were blood typed for systems A, B, C, D, E, H, I, and L. All birds were homozygous for allele L2, and allelic frequencies for the / system were essentially identical in both lines. In contrast, allelic frequencies of the other blood group systems differed rather markedly between the HA and LA lines. (Key words: alleles, alloantigens, chickens, antibody, selection) 1984 Poultry Science 63:1470-1472 INTRODUCTION
Studies of antigen recognition in antibody production of the fowl have generally focused on effects of specific loci and responses to infectious agents (Gavora and Spencer, 1983; van der Zijpp, 1983). Responses to divergent selection for antibody production to erythrocyte antigens have been demonstrated in mice (Biozzi et al, 1982), guinea pigs (Ibanez et al, 1980), and chickens (Siegel et al, 1982). Such selection has resulted in correlated responses in resistance to infectious agents, growth, and reproductive traits (Biozzi et al, 1982; Siegel et al, 1982). Presented here are comparisons of allelic frequencies for specific blood group systems in lines of chickens selected for high (HA) and low (LA) antibody response to the antigens of sheep red blood cells (SRBC). MATERIALS AND METHODS
Beginning with a sample of the Cornell randombred population (Lowe and Garwood, 1974), selection for high and low antibody response to antigens of SRBC was used to develop lines HA and LA, respectively (Siegel et al, 1982). Individual selection within each line
1
Virginia Polytechnic Institute and State University. 2 Northern Illinois University.
was based solely on antibody titers 5 days postchallenge to a .1 ml intravenous injection of a .25% suspension of SRBC. Parents (aged 386 to 407 days) from selected Generation 9 and their offspring were blood typed for Systems A, B, C, D, E, H, I, and L. Number of tested sires, dams, and offspring were 8, 21, and 141 in Line HA and 8, 19, and 138 in Line LA, respectively. Approximately .5 ml of blood was collected directly into a plastic test tube containing 2.0 ml of chilled isotonic solution of sodium citrate (2%) and sodium chloride (.42%) as anticoagulant. Tubes were coded, packed in ice, and immediately shipped from Blacksburg to DeKalb for blood typing. Reagents specific for alloantigens of each system were selected mostly on the basis of previous experience with the Cornell randombred or lines derived from that population (Briles et al, 1976, 1980; Briles and Briles, 1982). An allele of the A system not previously encountered in Cornell randombred-derived lines (Briles et al, 1977) was temporarily designated Ax. RESULTS Frequencies (%) of alleles and of homozygotes for each blood group system were similar for males and females; therefore, data were pooled (Table 1). All blood-typed individuals appeared to be homozygous for L 2 , as the L 1 allele which exists in other populations, was absent. Frequencies of alleles and homo-
1470
Downloaded from http://ps.oxfordjournals.org/ at North Dakota State University on May 25, 2015
(Received for publication March 19, 1984)
1471
RESEARCH NOTE TABLE 1. Frequencies (%) of genes and homozygotes at each locus in Generation 10 in lines selected for high (HA) and low (LA) antibody response Blood group system Allele
A
1 2 3 4 5 13 21 31 X
19/0 1 20/4
B
C
E
D
H
/
L
0/26 100/74
32/33
100/100
Gene frequencies 29/51 40/5
58/96
9/0
74/88 26/12
71/39 0/10
68/67
51/95
5/0 3/0
Homozygotes 11/28
3/0 6/0
1 2 3 4 5 13 21 31
11/0
52/83
0/5
5/0
100/53
9/11
100/100
53/17 44/97
0/0
44/45
0/2
25/89 2/98 61/0 0/0 1
(HA/LA).
zygotes for the / system were essentially identical in both lines. The HA line was 100% homozygous for allele H2. In Line LA, however, frequency of H2 was 74% with 5 3% of the chickens homozygous; frequency of H1 was 26% with 5% homozygotes. For the B system, 99% of the individuals were B 1 3 and 1% B 2 1 in Line LA, while in HA, frequencies were 15, 80, and 5 for B 1 3 , B21, and B 3 1 , respectively. Ninety-five percent of the C system alleles were C 5 in Line LA, while in Line HA 51% of the alleles were Cs. Conversely, LA chickens had only 5% allele C2 and 0% allele C 4 compared to 40% and 9% in the HA line. For the D system, frequencies for O 1 were 29 and 51%, D3 were 71 and 39%, and D 4 were 0 and 10% for Lines HA and LA, respectively. Systems A and Et which are linked, also showed differences between the selected lines; alleleic frequencies are given separately for each locus. Frequencies of alleles A1, A2, and ^44 in the HA line were 19, 20, and 58%, respectively, whereas in Line LA the corresponding frequencies were 0, 4, and 96%, respectively. Allelic frequencies of E1 and E2 were 74 and 26% in the HA line and 88 and 12% in LA line. Frequencies of segregating A-E haplotype combinations (Table 2) show
that Line LA chickens homozygous for E1 were also homozygous for A4. The results show considerable changes in allelic frequencies for several blood group systems concomitant to divergent selection for antibody response to SRBC. Although symmetry of response and drift effects are not known because allelic frequencies were not measured in the base generation, the data provide a basis for designing genotype-comparison experiments to evaluate effects of those systems exhibiting allelic segregation. This reasoning is consistent with the observations of
TABLE 2. Frequency of A-E haplotype combinations in lines selected for high (HA) and low (LA) antibody response Haplotype
HA
LA
A'-E 1 A 2 -E 2 A 4 -E' A 4 -E 2 A x -E' A X -E J
16 21 56 5 2 1
0 3 89 8 0 0
Downloaded from http://ps.oxfordjournals.org/ at North Dakota State University on May 25, 2015
15/99 80/1
1472
DUNNINGTON ET AL.
Benedict et al. (1975) that one homozygous B genotype failed to initiate response to a synthetic polypeptide whereas two other homozygotes and the three possible heterozygous genotypes responded fully. Our data also show for the A, B, C, and E systems unusually high frequencies of homozygotes in Line LA and a more normal frequency of heterozygotes in HA, suggesting evaluation of alleles in both heterozygous and homozygous states for antibody response to the SRBC antigens.
Downloaded from http://ps.oxfordjournals.org/ at North Dakota State University on May 25, 2015
REFERENCES Benedict, A. A., L. W. Pollard, P. R. Morrow, H. A. Abplanalp, P. H. Maurer, and W. E. Briles, 1975. Genetic control of immune responses in chickens. Immunogenetics 2:313—324. Biozzi, G., D. Mouton, A. M. Heumann, and Y. Bouthillier, 1982. Genetic regulation of immunoresponsiveness in relation to resistance against infectious diseases. 2nd World Congr. Genet. Appl. Livest. Prod. 5:150-163. Briles, W. E., and R. W. Briles, 1982. Identification of haplotypes to the chicken major histocompatibility complex (B). Immunogenetics 15: 441-447. Briles, W. E., R. W. Briles, W. H. McGibbon, V. A. Garwood, and P. C. Lowe, 1977. Erythrocyte alloantigens in Regional Cornell Randombred
stock. Poultry Sci. 56:1698. (Abstr.) Briles, W. E., R. W. Briles, W. H. McGibbon, and H. A. Stone, 1980. Identification of B alloalleles association with resistance to Marek's disease, Pages 395—413 in Resistance and Immunity to Marek's Disease. P. B. Biggs, ed. Coram. Eur. Commun., Directorate-Gen. Sci. Tech. Info., Luxembourg. Briles, W. E., W. H. McGibbon, and H. A. Stone, 1976. Effects of B alloalleles from Regional Cornell Randombred stock on mortality from Marek's disease. Poultry Sci. 55:2011-2012. Gavora, J. S., and J. L. Spencer, 1983. Breeding for immune responsiveness and disease resistance. Anim. Blood Groups Biochem. Genet. 14: 159-180. Ibanez, O. M., M. S. Reis, M. Gennari, V.C.A. Ferreira, O. A. Sant' Anna, M. Siqueira, and G. Biozzi, 1980. Selective breeding of high and low antibody responder lines of guinea pigs. Immunogenetics 10:283-293. Lowe, P. C , and V. A. Garwood, 1974. Inbreeding in 16 generations of the Regional Cornell Control populations of chickens. Poultry Sci. 53:514— 517. Siegel, P. B., W. B. Gross, and J. A. Cherry, 1982. Correlated responses of chickens to selection for production of antibodies to sheep erythrocytes. Anim. Blood Groups Biochem. Genet. 13: 291-297. van der Zijpp, A. J., 1983. Breeding for immune responsiveness and disease resistance. World's Poult. Sci. J. 39:118-131.