Scientific Research and Essays

  • Abbreviation: Sci. Res. Essays
  • Language: English
  • ISSN: 1992-2248
  • DOI: 10.5897/SRE
  • Start Year: 2006
  • Published Articles: 2759

Full Length Research Paper

Haemoglobin genetic types and its association with qualitative traits in West African Dwarf sheep

Fajemilehin Samuel Oladipo
  • Fajemilehin Samuel Oladipo
  • Department of Animal Science, Faculty of Agricultural Science, Ekiti State University, Ado-Ekiti, Ekiti State, Nigeria.
  • Google Scholar
Adegun Maria Kikelomo
  • Adegun Maria Kikelomo
  • Department of Animal Science, Faculty of Agricultural Science, Ekiti State University, Ado-Ekiti, Ekiti State, Nigeria.
  • Google Scholar

  •  Received: 06 April 2020
  •  Accepted: 19 June 2020
  •  Published: 31 August 2020


This study was carried out to investigate the haemoglobin genetic types and their association with qualitative traits in West African Dwarf (WAD) sheep. The Modified Stratified Sampling Technique (MSST) was used to select the sampling sites within the selected state and animal samples within the sampling sites. A total of 280 adult sheep comprising 140 rams and 140 ewes aged 4 years were used for the study. Data were collected on Mendelian traits such as the horn status, wattle status and hair length on sex basis. Blood samples were collected from the animals for haemoglobin genetic types’ determination. The results showed that in ewes, the f(HbAA), f(HbAB) and f(HbBB) were 0.36, 0.28 and 0.36, respectively and the f(HbA) and f(HbB) was 0.50 in both alleles.  In rams, the f(HbAA), f(HbAB) and f(HbBB) were 0.68, 0.14 and 0.18, respectively and the f(HbA) and f(HbB) were 0.75 and 0.25, respectively. In the pooled data, f(HbAA), f(HbAB) and f(HbBB) were 0.68, 0.14 and 0.18, respectively and the f(HbA) and f(HbB) were 0.625 and 0.375, respectively.  The estimated heterozygosity was 0.47 and the estimated local inbreeding coefficient was 0.054. The hair length indicated sexual dimorphism with 12.79 to 12.98 cm in rams and 4.79 to 4.98 cm in ewes but was not dependent on the heamoglobin genetic types. The result shows that the status of wattles is not influenced by sex. The WAD sheep used had three haemoglobin genotypes under the control of two alleles at the haemoglobin locus.

Key words: Ewe, haemoglobin genetic type, inbreeding coefficient, Mendelian traits, ram.



West African Dwarf (WAD) sheep is owned in small stock by rural peasant farmers in southwestern Nigeria. It is an integral part of family units, features prominently in socio-cultural functions and as emergency source of fund. It is versatile and genetically-adapted to the zone where the basic conditions of its well-being are not compromised (Petazzi et al., 2009). This native livestock is not favored in industrial animal agriculture because of the characteristic small body size and low productivity. However, its tolerance to diseases and seasonal fluctuations in food and water availability are qualities that can be exploited in industrial animal agriculture.

The genetic quality of this livestock species is under threat of erosion by industrial breeds such that their morphologic and genotypic characterization is essential (FAO, 2011). Consequently,  a  lot  of  studies  had  been carried out in the area of morphological characterization (Salako and Ngere, 2002; Yakubu and Akinyemi, 2010; Yakubu and Ibrahim, 2011). Unfortunately, the morphological characteristics do not correspond to the genetic characteristics of blood protein and non-protein polymorphisms because the traits are complex in their mode of transmission and are influenced by the environment (Tsunoda et al., 2010). Blood protein characterization therefore may present higher accuracy procedures for a better measurement of genetic variation in sheep breeds because of their polymorphisms and simple mode of inheritance.

Blood protein characterization using electrophoretic method has been used as a tool for studying relationships among farm animals (Sun et al., 2009; Shahrbabak et al., 2010) and genetic differentiation among breeds and in phylogenetic studies (Ibeagha-Awemu and Erhardt, 2004; Camoglu and Elmaci, 2005). Although DNA-based technologies are now in vogue for genetic characterization, the analysis of genetic markers based on blood protein and non-protein variants remains useful because of its utility, ease, cost, amount of genetic information accessed, simplicity of data interpretation (Arora et al., 2011) and because genetic research in Africa is less fully developed as in Europe (Gifford-Gonzalez and Hanotte, 2011).

Among these, blood proteins is haemoglobin which is the red oxygen carrying pigment in red blood cells of vertebrates. It is a conjugated protein with two pairs of identical sub-units, each with a cleft that contains iron-porphyrin group which is the site of oxygen uptake and release. Osaiyuwu and Salako (2018) found two genotypes of Hb comprising 72% HbAA and 28% HbAB in WAD sheep while Dafur et al. (2019) reported 100% HbAA. Mabruka and Ahmed (2018) reported 22% of HbA and 78% of HbB as gene frequencies of alleles A and B in WAD sheep. While several other research reports are available on the influence of haemoglobin genetic types on disease resistance, ovulation rates and blood traits in local and exotic breeds of sheep (FAO, 1988; Di Stasio, 1997), little had been done on its relationship with Mendelian traits. It is therefore, the objective of this study to investigate the haemoglobin genetic types and its association with some qualitative traits in WAD sheep.



The Modified Stratified Sampling Technique (MSST) was adopted for the choice of sampling sites within Ekiti State, Nigeria and the animals sampled within the sampling sites. A total of 280 adult sheep comprising 140 rams and 140 ewes aged 4 years were used for the study. The age of the sheep was estimated using permanent teeth eruption (Gerald, 1994). In all the sampling sites, the animals were managed extensively and fed with kitchen wastes only when available.

Data collection and laboratory analysis

Data were collected on Mendelian traits  such  as  the  horn  status, wattle status and hair length on sex basis as described by Akinyemi and Salako (2010). Blood samples were collected from the adult sheep of both sexes by jugular venipuncture into well labeled Bijou bottles containing a speck of ethylenediaminetetraacetic acid (EDTA) as anticoagulant. Plasma and erythrocyte samples were separated from the heparinized whole blood by centrifugation. After centrifugation, red cells were washed three times in saline solution (0.155 M NaCl), and lysed with a four-fold volume of distilled H2O to release haemoglobin. The separated lysate was stored at 4°C prior to laboratory analysis.

Gel electrophoresis was carried out on cellulose acetate strips to analyze inherited biochemical differences at haemoglobin (Hb) locus. This involved the use of Tris EDTA Borate at pH 8.6 for haemoglobin as described by Riken (2006). The resultant gel was stained with Red Ponceau stain to visualize the protein bands. The frequency of the allele corresponding to each band was estimated by direct count.

Statistical analysis

Data on Hb alleles and genotypic frequencies were subjected to Chi-square goodness of fit (one sample test) to know whether the data conform to Hardy-Weinberg equilibrium using the formula:

Æ©(O - E)2 / E

where O denotes the observed data and E is the expected value.

The degree of heterozygosity was calculated as the expected proportion of heterozygotes in a population under Hardy-Weinberg equilibrium.

The PROC GLM and PROC T-test procedures of SPSS (1989) were used to analyze the effects of haemoglobin variants on the Mendelian traits in the WAD sheep.



The distribution of genotypic and allelic frequencies of haemoglobin genotypes of WAD sheep is shown in Table 1. In the rams, only 20 and 25 had HbAB and  HbBB  with genotypic frequencies of 0.14 and 0.18, respectively while those with HbAA were 95 with genotypic frequency of 0.68. The gene frequencies for the two co-dominant alleles A and B were 0.75 and 0.25, respectively. In the ewes, 50 had HbAA, 40 had HbAB and 50 had HbBB with genotypic frequencies of 0.36, 0.28 and 0.36, respectively. The gene frequencies for the two co-dominant alleles A and B were the same at 0.50. When the data were pooled, 145 HbAA, 60 HbAB and 75 HbBB with genotypic frequencies of 0.52, 0.21 and 0.27, respectively were obtained. The allelic frequencies for the two co-dominant alleles A and B were 0.625 and 0.375, respectively. The estimated heterozygosity was 0.47  and the estimated local inbreeding coefficient was 0.054.



The observed and expected numbers of Hb alleles in WAD sheep are shown in Table 2. The allelic frequencies observed in the population under consideration deviated significantly (c2 = 46.66; p<0.05) from Hardy-Weinberg equilibrium. Table 3 shows the distribution of haemoglobin genetic types in association with hair length in WAD sheep. The hair length in the Hb genetic types varied from 12.79 to 12.98 cm and 4.79 to 4.98 cm in male and female WAD sheep, respectively. The distribution of haemoglobin genetic types in association with wattles in WAD sheep is presented in Table 4. The were  80,  35  and  25, respectively in male and 50 and 50 and 40 in female.







The haemoglobin genotypes, two homozygotes AA and BB and one heterozygote AB, detected in this study agree with the general observation that alleles A and B at the same locus are capable of producing three different genotypes AA, AB and BB, in different species of animals (Zaragoza et al., 1987; Tunon et al., 1989). The finding also agrees with Rodero et al. (1996) in Lebrijan Churro breed of Andalusia sheep but not in Merino sheep and Akinyemi and Salako (2010) in WAD sheep. The proportions of the haemoglobin genotypes at 52% HbAA, 22% HbAB and 27% HbBB in this study differ from the 100% HbAA reported by Dafur et al. (2019) in WAD sheep; the 72% HbAA and 28% HbAB obtained by Osaiyuwu and Salako (2018) in WAD sheep; the 88.89% HbAA and 11.11% HbAB in WAD sheep obtained by Akinyemi and Salako (2010); the 0.00% HbAA, 11.11% HbAB and 88.89% HbBB in Lebrijan Churro breed of Andalusia sheep and the 17% HbAA, 20.65% HbAB and 77.17% HbBB for the Grazalema Merino sheep reported by Rodero et al. (1996). However, the result supports the association of the highest frequency of HbAA in females (Akinyemi and Salako, 2010) and the HbAA genotype being the most frequent in WAD sheep (Osaiyuwu and Salako, 2018).

The preponderance of HbAA (52%) genotype can be attributed to an adaptive feature for survivability of the breed as reasoned by Agaviezor et al. (2013) that HbAA has a selective advantage in small ruminants. Also, Tella et al. (2000) reported that the HbAA genotype has selectable advantage in sheep at higher altitudes because its frequency increases towards the forest zone in the Southwestern, Nigeria. The HbAA is more preponderant in rams than in ewes while the HbAB and HbBB are more in ewes which is contrary to the report of Agaviezor et al. (2013) meaning that sex has an effect on the distribution of Hb types in WAD sheep. The and obtained in this study at 62.5 and 37.5%, respectively differed from 22% of HbA and 78% of HbB obtained by Mabruka and Ahmed (2018) and lower than the corresponding values of 94 and 6% obtained in WAD sheep (Akinyemi and Salako, 2010).

The at 0.22 is an indication of the level of genetic diversity at the Hb locus in the investigated sheep population. The estimated degree of heterozygosity (0.47) falls within the 0.30 and 0.80 range reported by Takezaki and Nei (1996) to be appropriate for markers to be used for measuring genetic variation. The rate of inbreeding was low and is in line with the high degree of heterozygosity value, suggesting that the sheep population could be undergoing assortative mating or may be experiencing a Wahund effect.

Comparison of observed and expected allelic frequencies is a test of the fulfillment of the conditions on which Hardy-Weinberg equilibrium depends. These conditions are random mating among the parents of the individuals observed, absence of migration and mutation which were not fulfilled in this study. This result is consistent with the reports of Musa et al. (2016) but contradicts the observation of Imumorin et al. (1999). The uniformity in hair length within the same sex among the different genetic types indicates that hair length was not dependent on the heamoglobin genetic types.

The hair length showed sexual dimorphism being longer in male than female which agrees with the observation of Akinyemi and Salako (2010). However, the hair length at 9.27±4.39 and 4.84±1.28 cm for ram and ewe, respectively reported by Akinyemi and Salako (2010) were shorter than the values obtained in this study while their postulation that animals with AA haemoglobin type would have longer hair than the other genotypes was not upheld.

The trend observed in the distribution of haemoglobin genetic types in association with wattle in WAD-sheep shows no indication that the different genotypes affect either the presence or absence of wattle in the population. Also, the status of wattle was not sex influenced as it was found in both rams and ewes.



Haemoglobin polymorphisms in WAD sheep is defined by two alleles, ‘A’ and ’B’ with the being 0.625 while the was 0.375. The hair length showed sexual dimorphism with longer length in male than female. However, the haemoglobin genetic type did not influence the trait. The status of wattle is not sex influenced as it occurred in some rams and ewes.






The authors have not declared any conflict of interests.



Agaviezor BO, Ajayi FO, Benneth HN (2013). Haemoglobin polymorphism in Nigerian indigenous goats in the Niger Delta Region, Nigeria. International Journal of Science and Nature 4(3):415-419.


Akinyemi MO, Salako AE (2010). Haemoglobin polymorphism and morphometrical correlates in the West African Dwarf Sheep of Nigeria. International Journal of Morphology. 28(1):205-208.


Arora R, Bhatia S, Mishra BP, Joshi BK (2011). Population structure in Indian sheep ascertained using microsatellite information. Animal Genetics 42:242-250.


Camoglu G, Elmaci C (2005). Carbonic Anhydrase (Ca) and X- Protein(X-p) Types in Turkish Sheep Breeds. Journal of Applied Animal Research 28:125-128.


Dafur BS, Darfur GS, Anwo OJ (2019). Haemoglobin polymorphism in Nigerian breeds of sheep and goats. Nigerian Journal of Animal Science 21(1).


Di Stasio L (1997). Biochemical genetics. In: Genetics of sheep. Piper L and Runvisky A (Eds), Oxon, CAB International, pp. 133-148.


FAO (1988). Food and Agricultural Organization. Animal genetic resources conservation by management, data banks and Training. FAO Animal Production and Health Paper 44/1. Rome, FAO. P. 186.


FAO (2011). Food and Agricultural Organization. Draft guidelines on molecular genetic characterization of animal genetic resources. Commission on Genetic Resources for Food and Agriculture, 13th Regular Session, 18- 22 July, 2011, Rome. 


Gerald W (1994). The tropical agriculturalist Macmillian Press Ltd. London, pp. 54-57


Gifford-Gonzalez D, Hanotte O (2011). Domesticating Animals in Africa: Implications of Genetic and Archaeological Findings. Journal of World Prehistory 24(1):1-23.


Ibeagha-Awemu EM, Erhardt G (2004). Genetic variations between African and German sheep breeds and description of a new variant of vitamin D-binding protein. Small Ruminant Research 55:33-43.


Imumorin IG, Ologun AG, Oyeyemi MO (1999). Preliminary observations on effects of hemoglobin genotype and estimate of genetic distance at the Hb locus in West African Dwarf and Red Sokoto goats. Tropical Journal of Animal Science 1:1-9.


Mabruka SHS, Ahmad AAS (2018). Preliminary investigation of hemoglobin polymorphism and its association with some morphometric traits and hematological parameters of sheep and goats in northeastern Libya. International Journal of Advanced Research 6(10):507-515.


Musa J, Garba H, Egena SSA, Aremu A (2016). Genetic variation of different phenotypes of West African Dwarf Goat based on haemoglobin polymorphism in Gulu, Niger State, Nigeria. Nigerian Journal of Agriculture, Food and Environment 12(3):108-113


Osaiyuwu HO, Salako EA (2018). Genetic structure of indigenous sheep breeds in Nigeria based on electrophoretic polymorphous systems of transferrin and haemoglobin. African Journal of Biotechnology 17(12):380-388.


Petazzi F, Rubino G, Alloggio I, Caroli A, Pieragostini E (2009). Relationships among functional markers, management, and husbandry in sheep: A Mediterranean case study. Veterinary Resource Communication 33:865-874.


RIKEN (2006). Institute of Physical and Chemical Research. Genetic quality monitoring by Biochemical isozymes. Riken Bioresource Centre. 


Rodero E, de la Haba MR, Rodero A, Herrera M (1996). Genetics and phenotypic profile of endangered Andalusian sheep and goat breeds. Rome, FAO, 1996.


Salako AE, Ngere LO (2002). Application of multifactorial discriminant analysis in the morphometric structural differentiation of West African Dwarf (WAD) and Yankassa sheep in South West Nigeria. Nigerian Journal of Animal Production 29(2):163-167.


Shahrbabak HM, Farahani AHK, Shahrbabak MM, Yeganeh HM (2010). Genetic variations between indigenous fat-tailed sheep populations. African Journal of Biotechnology 9:5993-5996.


SPSS (1989). Statistical Packages for Social Sciences. SPSS for Windows 6.1.3. Copyright Q SPSS Inc., v 1989-2000. Istanbul, Turkey.


Sun W, Musa, HH, Chang H, Tsunoda K (2009). Comparison of genetic detection efficiency of different markers under the same genetic background. African Journal of Biotechnology 8:2437-2442.


Takezaki N, Nei M (1996). Genetic distances and reconstruction of phylogenetic trees from microsatellite DNA. Genetics144:389-399.


Tella MA, Taiwo VO, Agbede SA, Alonge OD (2000). The influence of haemoglobin types on the incidence of babesiosis and anaplasmosis in West African Dwarf and Yankasa sheep. Tropical Veterinary Journal 18:121-127.


Tsunoda K, Chang H, Chang G, Sun W, Dorji T, Tshering G, Yamamoto Y, Namikawa T (2010). Phylogeny of Local Sheep Breeds in East Asia, Focusing on the Bayanbulak Sheep in China and the Sipsu Sheep in Bhutan. Biochemical Genetics 48:1-12.


Tunon MJ, Gonzalez P, Vallejo M (1989) Genetic relationships among 14 Spanish breeds of goats. Animal Genetics 20:205-212.


Yakubu A, Akinyemi MO (2010). An evaluation of sexual size dimorphism in Uda sheep using multifactorial discriminant analysis. Acta Agriculturae Scandinavica A-Animal Science 60:74-78.


Yakubu A, Ibrahim IA (2011). Multivariate analysis of morphostructural characteristics in Nigerian indigenous sheep. Italian Journal of Animal Science 10:83-86.


Zaragoza P, Arana A, Zaragoza I, Amorena B (1987). Blood biochemical polymorphisms in rabbits presently bred in Spain: Genetic variation and distances among populations. Australian Journal of Biological Sciences 40:275-286.