Comp. Bimhem Physiol., Vol. 62B, pp. 305 to 308. 0305-0491 79 0401-0305502.00/0
© Pergamon Press Ltd 1979. Printed in Great Britain
HEMOGLOBIN POLYMORPHISM IN EQUUS PRZEWALSKII
AND E. CABALLUS ANALYZED BY
ISOELECTRIC FOCUSING
OLIVER A. RYDER
Zoological Society of San Diego, Research Department, PO Box 551,
San Diego. California 92112 U.S.A.:
ROBERT S. SPARKES and MARYELLEN C. SPARKES
School of Medicine, Department of Medicine, University of California, Los Angeles,
California 90024 U.S.A.
and
JOHN B. CLEGG
Oxford University, Nuffield Department of Clinical Medicine, Radcliffe Infirmary,
Oxford OX2 6HE, U.K.
(Receired 30 August 19781
Abstract 1. Through the use of isoelectric focusing and peptide analysis, the hemoglobins of Prze-
walski's horse, Equus przewalskii and the domestic horse, E. caballus have been compared.
2. Przewalski's horses have two separate ~-globin chain polymorphisms similar to domestic horses.
Each hemoglobin phenotype could be accurately determined by isoelectric focusing.
3. Confirmation of the electrofocusing hemoglobin determinations was made by comparison to amino
acid composition analyses of purified tryptic peptides and by analysis of the rare hemoglobins pheno-
types observed in a family of Norwegian trotting horses.
4. Hemoglobin genotypes of fifteen Przewalski's horses were determined and inheritance of hemo-
globin haplotypes has been observed.
INTRODUCTION
Przewalski's horse, Equus przewalskii is the only true
wild horse surviving to the present day. Restricted
to the western Takin-Shara-Nuru range in the Mon-
golian Peoples Republic in recent times, its former
range extended throughout Europe and Asia (Mohr,
1970). Some 270 of this endangered species survive
in captivity (Volf, 1976) in zoos and animal parks.
Comparative analyses of E. przewalskii and the
domestic horse, E. caballus, utilizing a variety of mor-
phological and biochemical criteria have established
the close similarity of these two species (Simpson,
1950: Trommershausen-Smith et al., 1979). Cytogene-
tic studies of domestic and Przewalski's horses have
shown that the karyotypes of these two species differ
by a Robertsonian translocation. Comparative
chromosome banding studies of these two species
have revealed that fusion of four acrocentric chromo-
somes of E. przewalskii forms two metacentric
chromosomes (pair 5) in E. caballus (Short et al.,
1974; Ryder et al., 1978).
Domestic horse hemoglobins have been shown by
Kilmartin & Clegg (1967) to exhibit two distinct
~t-chain polymorphisms. Substitution of lysine (lys) or
gutamine (glnl at position 60 of the ct-globin chains
(ct60) produce, respectively, the slow or fast hemo-
globins detectable by starch gel electrophoresis (Kil-
martin & Clegg, 1967). Normally, the fast form
accounts for 60°4, of the total hemoglobin, and the
slow form 40~o. Additionally, analysis of amino acid
compositions of tryptic peptides of purified ~-globin
chains revealed a polymorphism at position 24 (ct24)
in which both phenylalanine (phe) and tyrosine (tyr)
residues may occur. Further genetic and biochemical
studies by Clegg (1970) have shown that four distinct,
heritable, a-chain sequences exist, depending on the
substitutions at positions 24 and 60. All /3-chain
sequences examined were apparently identical.
Isoelectric focusing of hemoglobins has proved to
be a powerful tool in separating sequence variants
in humans (Drysdale et al., 1971: Jeppson et al., 1972:
Monte et al., 1976) and other species as well (Butcher
& Hawkey, 1977). We have utilized isoelectric focus-
ing techniques to compare the hemoglobins of Prze-
walskFs and domestic horses and investigate whether
similar polymorphisms occur in these two species.
These studies have revealed that the four ~-chain
sequences present in E. caballus are also present in
E. przewalskii and, furthermore, each phenotype can
be accurately determined by isoelectric focusing with-
out resorting to amino acid composition analysis of
separated tryptic peptides. Electrofocusing analysis
was also able to accurately identify the rare hemo-
globin variants found in Norwegian trotting horses
by Braend (19671 and further characterized by Clegg
(1970).
MATERIALS AND METHODS
Blood samples from E. caballus were obtained from nor-
mal adult animals by venipuncture. Blood samples from
E. przewalskii were obtained by venipuncture of adult ani-
mals which had been chemically immobilized for routine
veterinary procedures.
Isoelectric focusing was performed in thin polyacryl-
amide slab gels on a commercial apparatus (LKB Multi-
305