Tropical Animal Health and Production, 3 1 (1 ...
Tropical Animal Health and Production, 3 1 (1 WI) 175-l 79
!Q: 1999 Kluwer Academic Pubhshers. Printed in the Netherlands
Blood Polymorphism in West African Breeds of Sheep
A. Missohou’*, T.C. Nguyen2, R. Sow3 and A, Gueye’
‘Service de Zootechnie-Alimentation, Ecole Inter-Etats des Sciences et Médecine
Vétérinaire.~ (EISMV), BP 5077 Dakur, Sénégal; “Laboratoire dAna1y.w Génétiques
pour les Espèces Animales (LABOGENA), INRA, 78352 Jouy-en-Josas, France; ‘Institut
National de Recherche Agronomique, Laboratoire de Hann, Dakar, Sénégal
Yorrespondence
and Gueye, A., 1999. Blood polymorphism in West African breeds of
31(3), 1755179
ABSTRACT
This paper reports the blood groups and blood protein distribution in West African sheep breeds.
About 100 animals of the Djallonke, Fulani and Touabire breeds were sampled for blood polymorph-
ism analysis. Their blood groups were typed by haemolytic and agglutination reactions. and their blood
proteins by starch gel electrophoresis. Almost dl the loti analysed showed variability in the three
breeds. with the Touabire and Fulani being closer to each other than to the Djallonke.
Keyltwdx blood groups, breed, haemoglobin, sheep, serum protein, transferrin
INTRODUCTION
In West Africa, two types of hairy thin-tailed sheep have been reported: the Savannah
and the forest types (Epstein, 1971). The Savannah type in the tsetse fly-free area is
large, long-legged and trypanosusceptible, while the dwarf forest type found in the
humid zone is trypanoresistant. Bradley (1995) demonstrated an increasing introgres-
sion of trypanosusceptible blood into trypanoresistant cattle as a consequence of the
increase in transhumance in recent decades due to drought and to farmers’ preference
for large animals.
Since genetic diversity is a basic requirement for animal improvement, it needs to be
preserved through characterization and management. Blood groups and blood
proteins have been used widely to characterize animal populations because of their
polymorphism and their simple mode of inheritance. Moreover, they are supposed to
be linked to productivity traits and environmental adaptation (Dally et al., 1980;
Vicovan and Rascu, 1989; Charon et a/., 1996). However, data on blood polymorphism
in tropical African sheep are scarce and refer only to their haemoglobin types
(Olusanya, 1975; Ndamukong, 1995).
This paper reports the distribution of blood groups and blood proteins in three West
African breeds of sheep.
176
MATERIALS AND METHODS
Animuls
Blood samples were collected from Senegalese Djallonke (dwarf forest type). Peul-peul
(Fulani) and Touabire (Savannah type) breeds of sheep. The Fulani sheep were sampled
in the Djoloff, located 250 km from Dakar in north Senegal, while the Djallonke
animals were chosen in Kolda, south Senegal, about 500 km from the capital. The
Touabire sheep, which originated from Mauritania, were sampled in Dakar market,
where they are brought for sale.
Blood samples (5 ml) were collect ed from approximately 100 animals per breed. The
animals were adults of both sexes and were chosen from different flocks. The blood
samples were collected into sodium citrate and sent to the Laboratoire d’Analyses
Génétiques pour les Espèces Animales (LABOGENA), Jouy-en-Josas. France, for
analysis.
Blood typing and data analysis
The blood protein polymorphism was analysed by starch gel electrophoresis and by
electrofocalisation in the case of haemoglobin (Nguyen and Bunch, 1980).
Six OEA (ovine erythrocyte antigen) systems were analysed by haemolytic and
haemagglutination reactions as described by Nguyen ( 1972). Blood protein alleles
were determined by direct counting; blood group ‘allele’ frequencies were calculated
after Hardy-Weinberg equilibrium had been established at the transferrin and
carbonic anhydrase loti.
RESULTS
Blood group polymorphism
The distribution of blood groups in the three sheep breeds is shown in Table 1. The
observed allelic frequencies may be biased because of the low number of animals used.
However, according to Nei (1978), the error is low in sample sizes of 100. Except for
locus B, which was very polymorphie, all the other alleles were present in the three
breeds at different frequencies. The differences between the Djallonke and Fulani sheep
were statistically significant 0, c 0.05) for alleles Aah, Ch, B“ and R, the Touabire sheep
being intermediate.
Bloodproteins
177
TABLE 1
OEA allele frequencies in West African breeds of sheep
Frequencies
Systems
Allelles
Djallonke
Fulani
Touabire
A
a
0.526
0.55
0.499
b
0.087
0.035
0.068
a b
0.025
0.001
0.021
B
b
0.03
0.181
0.161
a b
0.005
0.027
0.108
0
0.022
0
abi
0
0.023
0
hi
0.020
0.051
0.148
d
0.05
0.007
0.027
di
0
0
0.046
e
0.05
0.014
0.013
ei
0
0
0.038
i
0.025
0.079
0.020
a
0
0.022
0
a
0.005
0.045
0.071
ab
0.005
0.056
0.063
D
a
0.529
0.341
0.471
M
a
1
0.99
0.975
R
r
0.326
0.44
0.408
Accordance with Hardy-Weinberg law”
Tf: x2: 4 df
4.64
3.25
4.92
CA: x2: ldf
0.038
0.756
0.277
‘Tf, transferrin; CA, carbonic anhydrase: df, degrees of freedom
Djallonke sheep in Cameroon (Ndamukong, 1995) but contradicts results obtained in
the same breed in Nigeria by Olusanya (1975). Concerning transferrin, seven
phenotypes (AA, CC, AD, AB, AC, AD and CD) were observed, the gene alleles
being TfA, TfB, TfC and TfD. The pattern of distribution of the B and D alleles
(respectively low and high in the Djallonke breed and medium in the other breeds)
rnnfirmecl thp recnltc nhtaind in trvnonncl~crm+;hl
P anrl tr.m~nnl.m~c.tr>n+
nntt1- L-7
178
TABLE II
Distribution of blood protein in West African breeds of sheep
Allele frequencies
Systems
Alleles
Djallonke
Fulani
Touabire
Transferrin
A
0.216
0.428
0.308
G
0
0.041
0.030
B
0.05
0.072
0.142
c
0.110
0.139
0.126
D
0.609
0.320
0.394
Haemoglobin
A
0
0
0.15
B
1
1
0.985
Carbonic anhydrase
M
0.232
0.08
0.05
s
0.768
0.920
0.95
Protein X
X
0.112
0.094
0.089
DISCUSSION
Although several sheep breeds have been described in West Africa (Doutressole, 1947;
Epstein, 197 l), little is known about the extent of genetic diversity between and within
these breeds (Rege, 1994). In this paper, we analyse the value of blood groups and
blood protein polymorphism in thc genetic study of sheep breeds. The overall results
showed significant differences between the Djallonke breed on the one hand and the
Touabire and Fulani breeds on the other. In a spatial autocorrelation study, Ordas and
Carriedo (1996) showed that the allele frequency differences among European sheep
breeds were due to migration and genetic drift. This could support the existence of
differences in the historic expansion into West Africa of Djallonke and the other breeds
and is consistent with the classification of Epstein (1971). In fact, this author pooled
the Touabire and Fulani breeds in the same group (the Savannah) while the Djallonke
breed was classified as a forest type.
CONCLUSION
Analysis of the biochemical polymorphism in West African sheep breeds showed
differences among breeds which reflected their historic expansion. However, this work
needs to be continued on a greater number of breeds and wirh other markers
I..-:------r-ll:r-\\ T-L- --l-L---L:, L,.* _.,__- cl..,,, -rn*o;rrn -,,q nnrxn*m~r~lllr m&=wJnt
1 7 9
REFERENCES
Braend, M. and Khana, N.D., 1968. Haemoglobin and transferrin type of some West African cattle.
AnimalProduction, 10, 129-134
Bradley, D., 1995. Genetie characterisation of cattle in West and Central Africa, (Department of Genetics,
Trinity College, Dublin 2, Annual report)
Charon, K.M., Lipecka, C., Siudek, T. and Swiderek, W., 1996. Relationship between transferrin and
globulin antigen polymorphism and sheep resistance to mastitis. Journal of Applied Genet& 37, 161-
l-71
Dally, M.R., Hohenboken, W., Thomas, D.L. and Craig, A.M., 1980. Relationships between hemoglobin
type and reproduction, lamb, wool and milk production and health-related traits in crossbred cwes.
Journal of Animal Science, 50,418-427
Doutressole, E.G., 1947. L’élevage en Afrique Occidentale Française, (Larousse, Paris)
Epstein, H., 1971. Origin of the Domestic Animais of Africa ZZ, (African Publishing Corporation, New York,
London, Munich)
Ndamukong, K.J.N., 1995. Haemoglobin polymorphism in grassland dwarf sheep and goats of the north
west province of Cameroon. Bulletin of Animal Health and Production in Africa, 43 53-56
Nei, M., 1978. Estimation of average heterozygosity and genetic distance from a small number of
individuals. Genetics, 89, 583-590
Nguyen, T.C., 1972. Les groupes sanguins des ovins 1. Relations entre les groupes sanguins des ovins et des
bovins. Annale de Génétique et de Sélection Animale, 4, 363-374
Nguyen, T.C. and Bunch, T.D., 1980. Blood groups and evolutionary relationships among sheep (Ovis
aries), domestic goat (Capra hircus) aoudad (Ammotragus lervia) and European mouflon (Ovis musimon).
Annales Génétique Sélection Animale, 12, 169-180
Olusanya, S.K., 1975. Electrolyte concentration and haemoglobin types in the blood of Nigerian dwarf
sheep. Journal of the Nigerian Veterinart Medical Association, 4, 59-63
Ordas, J.G. and Carriedo, J.A., 1996. Spatial autocorrelation of ovine protein polymorphisms in Europe.
Genetics, Selection Evolution, 28, 531-536
Queval, R., 1982. Polymorphisme de la transferrine chez les bovins trypanosensibles et trypanotolérants de
l’Afrique de l’ouest. Répartition de fréquence des allèles. Revue d’Elevage et de Médecine Vétérinaire
Pays Tropicaux, 35, 373-380
Rege. J.E.O., 1994. Indigenous African small ruminants: a case of characterisation and improvement. In:
Small Ruminant Research and Development, Proc. 2nd Bi-annual Conference of the African Small
Ruminant Network, 1992, (AICC, Arusha, Tanzania)
Vicovan, G. and Rascu, D., 1989. Types of hdemoglobin in sheep related to environmental adaptation.
Archiva Zootechnica, 1, 33-44
(Accepted: 29 October 1998)
Polimorfismo sanguineo en razas ovinas de Africa Occidental
Resumen -, Este articula estudia la distribucibn de proteinas plasm&ticas y grupos sanguineos en razas
ovinas de Africa Occidental. Se obtuvieron muestras de sangre de alrededor de 100 animales de las razas
Djallonke, Fulani y Touabire, con objeto de estudiar su polimorfismo sanguineo. Los grupos sanguineos se
tipificaron mediante reacciones de hemolisis y aglutinacibn, y las proteinas sanguineas mediante
electroforesis en gel. Casi todos 10s loti analizados mostraron variabilidad en las 3 razas, siendo las razas
Touabire y Fulani mas parecidas entre si que a la raza Djallonke.
!Q: 1999 Kluwer Academic Pubhshers. Printed in the Netherlands
Blood Polymorphism in West African Breeds of Sheep
A. Missohou’*, T.C. Nguyen2, R. Sow3 and A, Gueye’
‘Service de Zootechnie-Alimentation, Ecole Inter-Etats des Sciences et Médecine
Vétérinaire.~ (EISMV), BP 5077 Dakur, Sénégal; “Laboratoire dAna1y.w Génétiques
pour les Espèces Animales (LABOGENA), INRA, 78352 Jouy-en-Josas, France; ‘Institut
National de Recherche Agronomique, Laboratoire de Hann, Dakar, Sénégal
Yorrespondence
and Gueye, A., 1999. Blood polymorphism in West African breeds of
31(3), 1755179
ABSTRACT
This paper reports the blood groups and blood protein distribution in West African sheep breeds.
About 100 animals of the Djallonke, Fulani and Touabire breeds were sampled for blood polymorph-
ism analysis. Their blood groups were typed by haemolytic and agglutination reactions. and their blood
proteins by starch gel electrophoresis. Almost dl the loti analysed showed variability in the three
breeds. with the Touabire and Fulani being closer to each other than to the Djallonke.
Keyltwdx blood groups, breed, haemoglobin, sheep, serum protein, transferrin
INTRODUCTION
In West Africa, two types of hairy thin-tailed sheep have been reported: the Savannah
and the forest types (Epstein, 1971). The Savannah type in the tsetse fly-free area is
large, long-legged and trypanosusceptible, while the dwarf forest type found in the
humid zone is trypanoresistant. Bradley (1995) demonstrated an increasing introgres-
sion of trypanosusceptible blood into trypanoresistant cattle as a consequence of the
increase in transhumance in recent decades due to drought and to farmers’ preference
for large animals.
Since genetic diversity is a basic requirement for animal improvement, it needs to be
preserved through characterization and management. Blood groups and blood
proteins have been used widely to characterize animal populations because of their
polymorphism and their simple mode of inheritance. Moreover, they are supposed to
be linked to productivity traits and environmental adaptation (Dally et al., 1980;
Vicovan and Rascu, 1989; Charon et a/., 1996). However, data on blood polymorphism
in tropical African sheep are scarce and refer only to their haemoglobin types
(Olusanya, 1975; Ndamukong, 1995).
This paper reports the distribution of blood groups and blood proteins in three West
African breeds of sheep.
176
MATERIALS AND METHODS
Animuls
Blood samples were collected from Senegalese Djallonke (dwarf forest type). Peul-peul
(Fulani) and Touabire (Savannah type) breeds of sheep. The Fulani sheep were sampled
in the Djoloff, located 250 km from Dakar in north Senegal, while the Djallonke
animals were chosen in Kolda, south Senegal, about 500 km from the capital. The
Touabire sheep, which originated from Mauritania, were sampled in Dakar market,
where they are brought for sale.
Blood samples (5 ml) were collect ed from approximately 100 animals per breed. The
animals were adults of both sexes and were chosen from different flocks. The blood
samples were collected into sodium citrate and sent to the Laboratoire d’Analyses
Génétiques pour les Espèces Animales (LABOGENA), Jouy-en-Josas. France, for
analysis.
Blood typing and data analysis
The blood protein polymorphism was analysed by starch gel electrophoresis and by
electrofocalisation in the case of haemoglobin (Nguyen and Bunch, 1980).
Six OEA (ovine erythrocyte antigen) systems were analysed by haemolytic and
haemagglutination reactions as described by Nguyen ( 1972). Blood protein alleles
were determined by direct counting; blood group ‘allele’ frequencies were calculated
after Hardy-Weinberg equilibrium had been established at the transferrin and
carbonic anhydrase loti.
RESULTS
Blood group polymorphism
The distribution of blood groups in the three sheep breeds is shown in Table 1. The
observed allelic frequencies may be biased because of the low number of animals used.
However, according to Nei (1978), the error is low in sample sizes of 100. Except for
locus B, which was very polymorphie, all the other alleles were present in the three
breeds at different frequencies. The differences between the Djallonke and Fulani sheep
were statistically significant 0, c 0.05) for alleles Aah, Ch, B“ and R, the Touabire sheep
being intermediate.
Bloodproteins
177
TABLE 1
OEA allele frequencies in West African breeds of sheep
Frequencies
Systems
Allelles
Djallonke
Fulani
Touabire
A
a
0.526
0.55
0.499
b
0.087
0.035
0.068
a b
0.025
0.001
0.021
B
b
0.03
0.181
0.161
a b
0.005
0.027
0.108
0
0.022
0
abi
0
0.023
0
hi
0.020
0.051
0.148
d
0.05
0.007
0.027
di
0
0
0.046
e
0.05
0.014
0.013
ei
0
0
0.038
i
0.025
0.079
0.020
a
0
0.022
0
a
0.005
0.045
0.071
ab
0.005
0.056
0.063
D
a
0.529
0.341
0.471
M
a
1
0.99
0.975
R
r
0.326
0.44
0.408
Accordance with Hardy-Weinberg law”
Tf: x2: 4 df
4.64
3.25
4.92
CA: x2: ldf
0.038
0.756
0.277
‘Tf, transferrin; CA, carbonic anhydrase: df, degrees of freedom
Djallonke sheep in Cameroon (Ndamukong, 1995) but contradicts results obtained in
the same breed in Nigeria by Olusanya (1975). Concerning transferrin, seven
phenotypes (AA, CC, AD, AB, AC, AD and CD) were observed, the gene alleles
being TfA, TfB, TfC and TfD. The pattern of distribution of the B and D alleles
(respectively low and high in the Djallonke breed and medium in the other breeds)
rnnfirmecl thp recnltc nhtaind in trvnonncl~crm+;hl
P anrl tr.m~nnl.m~c.tr>n+
nntt1- L-7
178
TABLE II
Distribution of blood protein in West African breeds of sheep
Allele frequencies
Systems
Alleles
Djallonke
Fulani
Touabire
Transferrin
A
0.216
0.428
0.308
G
0
0.041
0.030
B
0.05
0.072
0.142
c
0.110
0.139
0.126
D
0.609
0.320
0.394
Haemoglobin
A
0
0
0.15
B
1
1
0.985
Carbonic anhydrase
M
0.232
0.08
0.05
s
0.768
0.920
0.95
Protein X
X
0.112
0.094
0.089
DISCUSSION
Although several sheep breeds have been described in West Africa (Doutressole, 1947;
Epstein, 197 l), little is known about the extent of genetic diversity between and within
these breeds (Rege, 1994). In this paper, we analyse the value of blood groups and
blood protein polymorphism in thc genetic study of sheep breeds. The overall results
showed significant differences between the Djallonke breed on the one hand and the
Touabire and Fulani breeds on the other. In a spatial autocorrelation study, Ordas and
Carriedo (1996) showed that the allele frequency differences among European sheep
breeds were due to migration and genetic drift. This could support the existence of
differences in the historic expansion into West Africa of Djallonke and the other breeds
and is consistent with the classification of Epstein (1971). In fact, this author pooled
the Touabire and Fulani breeds in the same group (the Savannah) while the Djallonke
breed was classified as a forest type.
CONCLUSION
Analysis of the biochemical polymorphism in West African sheep breeds showed
differences among breeds which reflected their historic expansion. However, this work
needs to be continued on a greater number of breeds and wirh other markers
I..-:------r-ll:r-\\ T-L- --l-L---L:, L,.* _.,__- cl..,,, -rn*o;rrn -,,q nnrxn*m~r~lllr m&=wJnt
1 7 9
REFERENCES
Braend, M. and Khana, N.D., 1968. Haemoglobin and transferrin type of some West African cattle.
AnimalProduction, 10, 129-134
Bradley, D., 1995. Genetie characterisation of cattle in West and Central Africa, (Department of Genetics,
Trinity College, Dublin 2, Annual report)
Charon, K.M., Lipecka, C., Siudek, T. and Swiderek, W., 1996. Relationship between transferrin and
globulin antigen polymorphism and sheep resistance to mastitis. Journal of Applied Genet& 37, 161-
l-71
Dally, M.R., Hohenboken, W., Thomas, D.L. and Craig, A.M., 1980. Relationships between hemoglobin
type and reproduction, lamb, wool and milk production and health-related traits in crossbred cwes.
Journal of Animal Science, 50,418-427
Doutressole, E.G., 1947. L’élevage en Afrique Occidentale Française, (Larousse, Paris)
Epstein, H., 1971. Origin of the Domestic Animais of Africa ZZ, (African Publishing Corporation, New York,
London, Munich)
Ndamukong, K.J.N., 1995. Haemoglobin polymorphism in grassland dwarf sheep and goats of the north
west province of Cameroon. Bulletin of Animal Health and Production in Africa, 43 53-56
Nei, M., 1978. Estimation of average heterozygosity and genetic distance from a small number of
individuals. Genetics, 89, 583-590
Nguyen, T.C., 1972. Les groupes sanguins des ovins 1. Relations entre les groupes sanguins des ovins et des
bovins. Annale de Génétique et de Sélection Animale, 4, 363-374
Nguyen, T.C. and Bunch, T.D., 1980. Blood groups and evolutionary relationships among sheep (Ovis
aries), domestic goat (Capra hircus) aoudad (Ammotragus lervia) and European mouflon (Ovis musimon).
Annales Génétique Sélection Animale, 12, 169-180
Olusanya, S.K., 1975. Electrolyte concentration and haemoglobin types in the blood of Nigerian dwarf
sheep. Journal of the Nigerian Veterinart Medical Association, 4, 59-63
Ordas, J.G. and Carriedo, J.A., 1996. Spatial autocorrelation of ovine protein polymorphisms in Europe.
Genetics, Selection Evolution, 28, 531-536
Queval, R., 1982. Polymorphisme de la transferrine chez les bovins trypanosensibles et trypanotolérants de
l’Afrique de l’ouest. Répartition de fréquence des allèles. Revue d’Elevage et de Médecine Vétérinaire
Pays Tropicaux, 35, 373-380
Rege. J.E.O., 1994. Indigenous African small ruminants: a case of characterisation and improvement. In:
Small Ruminant Research and Development, Proc. 2nd Bi-annual Conference of the African Small
Ruminant Network, 1992, (AICC, Arusha, Tanzania)
Vicovan, G. and Rascu, D., 1989. Types of hdemoglobin in sheep related to environmental adaptation.
Archiva Zootechnica, 1, 33-44
(Accepted: 29 October 1998)
Polimorfismo sanguineo en razas ovinas de Africa Occidental
Resumen -, Este articula estudia la distribucibn de proteinas plasm&ticas y grupos sanguineos en razas
ovinas de Africa Occidental. Se obtuvieron muestras de sangre de alrededor de 100 animales de las razas
Djallonke, Fulani y Touabire, con objeto de estudiar su polimorfismo sanguineo. Los grupos sanguineos se
tipificaron mediante reacciones de hemolisis y aglutinacibn, y las proteinas sanguineas mediante
electroforesis en gel. Casi todos 10s loti analizados mostraron variabilidad en las 3 razas, siendo las razas
Touabire y Fulani mas parecidas entre si que a la raza Djallonke.