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Topic:Genetic Diversity

Genetic diversity in horses refers to the variation in genetic characteristics within and among populations of horses. This diversity is shaped by factors such as breeding practices, geographic distribution, and historical population dynamics. Genetic diversity is measured using various molecular markers and provides insights into the evolutionary history, population structure, and adaptability of horse breeds. It influences traits such as disease resistance, fertility, and performance. This page compiles peer-reviewed research studies and scholarly articles that explore the assessment, implications, and conservation of genetic diversity in equine populations.
Immune selection of equine infectious anemia virus env variants during the long-term inapparent stage of disease.
Virology    February 27, 2007   Volume 363, Issue 1 156-165 doi: 10.1016/j.virol.2007.01.037
Sponseller BA, Sparks WO, Wannemuehler Y, Li Y, Antons AK, Oaks JL, Carpenter S.The principal neutralizing domain (PND) of equine infectious anemia virus (EIAV) is located in the V3 region of SU. Genetic variation in the PND is considered to play an important role in immune escape and EIAV persistence; however, few studies have characterized genetic variation in SU during the inapparent stage of disease. To better understand the mechanisms of virus persistence, we undertook a longitudinal study of SU variation in a pony experimentally inoculated with the virulent EIAV(Wyo). Viral RNA isolated from the inoculum and from sequential sera samples was amplified by RT-PCR, clon...
Molecular analysis using mitochondrial DNA and microsatellites to infer the formation process of Japanese native horse populations.
Biochemical genetics    February 1, 2007   Volume 45, Issue 3-4 375-395 doi: 10.1007/s10528-007-9083-0
Kakoi H, Tozaki T, Gawahara H.To assess the genetic diversity of Japanese native horse populations, we examined seven such populations using mitochondrial DNA (mtDNA) and microsatellite analyses. Four reference populations of Mongolian horses and European breeds were employed as other equids. In the mtDNA analysis, the control region (D-loop) of 411 bp was sequenced, and 12 haplotypes with 33 variable sites were identified in the Japanese native horses. The phylogenetic tree constructed by haplogrouping and using worldwide geographic references indicated that the haplotypes of the Japanese native horses were derived from s...
Genetic diversity and relationships of Portuguese and other horse breeds based on protein and microsatellite loci variation.
Animal genetics    January 30, 2007   Volume 38, Issue 1 20-27 doi: 10.1111/j.1365-2052.2006.01545.x
Luís C, Juras R, Oom MM, Cothran EG.There are three native Portuguese horse breeds: Lusitano, Sorraia and Garrano. This study compares diversity patterns of 17 protein and 12 microsatellite markers in these three as well as 30 other breeds to infer relationships among the breeds and to compare levels of polymorphism of these breeds for use in conservation efforts. The Garrano and the Lusitano showed a high level of genetic diversity, similar to that observed for most of the other analysed breeds, while the Sorraia and Friesian breeds showed low levels of variation for both genetic marker types. The combined protein and microsate...
Genetic structure and phylogenetic relationships of the Polish Heavy horse.
Journal of applied genetics    November 30, 2006   Volume 47, Issue 4 353-359 doi: 10.1007/BF03194645
Iwańczyk E, Juras R, Cholewiński G, Cothran EG.In this study a wide range of genetic markers (12 microsatellites, 7 blood-group loci, 10 blood-protein loci) and mitochondrial DNA (mtDNA) were used to assess genetic diversity in Polish Heavy horses. Three random samples were sequenced for 421 bp of the mitochondrial D-loop region, but no clear phylogenetic patterns were seen in mtDNA variation. Both heterozygosity and diversity levels are fairly high in Polish Heavy horses. In phylogenetic analysis the draught horses form a distinct cluster that pairs with the true pony breeds. Within this 'cold-blooded' group, the Polish Heavy Horse cluste...
Analysis of genetic diversity and the determination of relationships among western Mediterranean horse breeds using microsatellite markers.
Journal of animal breeding and genetics = Zeitschrift fur Tierzuchtung und Zuchtungsbiologie    September 13, 2006   Volume 123, Issue 5 315-325 doi: 10.1111/j.1439-0388.2006.00603.x
Marletta D, Tupac-Yupanqui I, Bordonaro S, García D, Guastella AM, Criscione A, Cañón J, Dunner S.The distribution of genetic diversity and the genetic relationships among western Mediterranean horse breeds were investigated using microsatellite markers. The examined sample included seven Spanish and three Italian local horse breeds and populations, plus a Spanish Thoroughbred outgroup. The total number of animals examined was 682 (on average 62 animals per breed; range 20-122). The microsatellite marker set analysed provided 128 alleles (10.7 alleles per locus). Within-breed genetic diversity was always high (>0.70), with breeds contributing about 8% of the total genetic variability. T...
Highly diverse type of equine arteritis virus (EAV) from the semen of a South African donkey: short communication.
Acta veterinaria Hungarica    July 18, 2006   Volume 54, Issue 2 263-270 doi: 10.1556/AVet.54.2006.2.12
Stadejek T, Mittelholzer Ch, Oleksiewicz MB, Paweska J, Belák S.Equine arteritis virus (EAV) was detected by RT-nested PCR in semen samples from a naturally infected South African donkey. Sequence analysis of the amplified ORF5 fragment revealed only 60 to 70% nucleotide identity to a panel of EAV reference sequences. The unique donkey EAV sequence was also found to be stable during passage in horses. The sequence data reported in this study indicate that the South African donkey variant might represent a new genotype of EAV. The distinct genetic properties of the South African asinine strain of EAV suggest a divergent evolution of this arterivirus in vari...
Genetic and permanent environmental variability of twinning in Thoroughbred horses estimated via three threshold models.
Journal of animal breeding and genetics = Zeitschrift fur Tierzuchtung und Zuchtungsbiologie    May 19, 2006   Volume 123, Issue 3 186-190 doi: 10.1111/j.1439-0388.2006.00575.x
Wolc A, Bresińska A, Szwaczkowski T.Twinning is a serious problem in Thoroughbred horses, as most of the multiple pregnancies are miscarried and live foals present lower performance values. The aim of this study was to estimate the heritability and repeatability of twinning in Thoroughbred horses. A total of 12648 pregnancies of 2033 Thoroughbred mares born between 1929 and 1994 in 11 studs were included. The analysis was performed by the use of AI-REML algorithm under three threshold animal models. The first model included effects of stud, mare's birth period and interaction between them as fixed. The second model included the ...
Genetic variation of Polish endangered Biłgoraj horses and two common horse breeds in microsatellite loci.
Journal of applied genetics    August 20, 2005   Volume 46, Issue 3 299-305 
Zabek T, Nogaj A, Radko A, Nogaj J, Słota E.Genetic variation of endangered Biłgoraj horses and two common Polish horse breeds was compared with the use of 12 microsatellite loci (AHT4, AHT5, ASB2, HMS2, HMS3, HMS6, HMS7, HTG4, HTG6, HTG7, HTG10, VHL20). Lower allelic diversity was detected in all investigated populations in comparison to other studies. Large differences in the frequencies of microsatellite alleles between Biłgoraj horses and two other horse breeds were discovered. In all polymorphic loci all investigated breeds were in the Hardy-Weinberg equilibrium. Mean Fis values and the results of a test for the presence of a rec...
Effect of moxidectin selection on the genetic variation within Cylicocyclus nassatus based on amplified fragment length polymorphism (AFLP).
International journal for parasitology    March 18, 2005   Volume 35, Issue 7 813-819 doi: 10.1016/j.ijpara.2005.02.004
Tandon R, Lyons ET, Tolliver SC, Kaplan RM.Cyathostomins are among the most important intestinal nematodes of horses, yet, the literature on the molecular genetics of these worms is scarce. In this study, the technique of amplified fragment length polymorphism (AFLP) was applied to study the genetic diversity as well as to determine the effect of moxidectin selection on the population genetic diversity for Cylicocyclus nassatus, one of the most common cyathostomin species. Genomic DNAs from 30 individual male worms were used from each of two populations: an avermectin-milbemycin (AM)-naive population (Population-S) and a population der...
Sequence analysis of the msp4 gene of Anaplasma phagocytophilum strains.
Journal of clinical microbiology    March 8, 2005   Volume 43, Issue 3 1309-1317 doi: 10.1128/JCM.43.3.1309-1317.2005
de la Fuente J, Massung RF, Wong SJ, Chu FK, Lutz H, Meli M, von Loewenich FD, Grzeszczuk A, Torina A, Caracappa S, Mangold AJ, Naranjo V, Stuen S....The causative agent of human granulocytic ehrlichiosis was recently reclassified as Anaplasma phagocytophilum, unifying previously described bacteria that cause disease in humans, horses, dogs, and ruminants. For the characterization of genetic heterogeneity in this species, the homologue of Anaplasma marginale major surface protein 4 gene (msp4) was identified, and the coding region was PCR amplified and sequenced from a variety of sources, including 50 samples from the United States, Germany, Poland, Norway, Italy, and Switzerland and 4 samples of A. phagocytophilum-like organisms obtained f...
Genetic variation within and relationships among five subpopulations of Slovak Thoroughbred.
Acta veterinaria Hungarica    September 24, 2004   Volume 52, Issue 3 259-265 doi: 10.1556/AVet.52.2004.3.2
Sátková-Jakabová D, Trandzík J, Hudecová-Kvasnáková L, Hegedüsová-Zetochová E, Bugarský A, Buleca J, Zöldág L, Jakab F, Fl'ak P.Genetic variation at six microsatellite loci was analysed for five Thoroughbred subpopulations to determine the magnitude of genetic differentiation and the genetic relationships among the subpopulations. Significant deviations from Hardy-Weinberg equilibrium were shown for a number of locus-population combinations, with all subpopulations. The genetic diversities and relationships of five Thoroughbred subpopulations were evaluated using six microsatellites recommended by the International Society of Animal Genetics (ISAG). The allele frequencies, the effective numbers of alleles, and the obse...
Genetic diversity in German draught horse breeds compared with a group of primitive, riding and wild horses by means of microsatellite DNA markers.
Animal genetics    July 22, 2004   Volume 35, Issue 4 270-277 doi: 10.1111/j.1365-2052.2004.01166.x
Aberle KS, Hamann H, Drögemüller C, Distl O.We compared the genetic diversity and distance among six German draught horse breeds to wild (Przewalski's Horse), primitive (Icelandic Horse, Sorraia Horse, Exmoor Pony) or riding horse breeds (Hanoverian Warmblood, Arabian) by means of genotypic information from 30 microsatellite loci. The draught horse breeds included the South German Coldblood, Rhenish German Draught Horse, Mecklenburg Coldblood, Saxon Thuringa Coldblood, Black Forest Horse and Schleswig Draught Horse. Despite large differences in population sizes, the average observed heterozygosity (H(o)) differed little among the heavy ...
Genetic variation in Przewalski’s horses, with special focus on the last wild caught mare, 231 Orlitza III.
Cytogenetic and genome research    February 19, 2004   Volume 102, Issue 1-4 226-234 doi: 10.1159/000075754
Bowling AT, Zimmermann W, Ryder O, Penado C, Peto S, Chemnick L, Yasinetskaya N, Zharkikh T.In our continuing efforts to document genetic diversity in Przewalski's horses and relatedness with domestic horses, we report genetic variation at 22 loci of blood group and protein polymorphisms and 29 loci of DNA (microsatellite) polymorphisms. The loci have been assigned by linkage or synteny mapping to 20 autosomes and the X chromosome of the domestic horse (plus four loci unassigned to a chromosome). With cumulative data from tests of 568 Przewalski's horses using blood, hair or tooth samples, no species-defining markers were identified, however a few markers were present in the wild spe...
Microsatellite variation in Japanese and Asian horses and their phylogenetic relationship using a European horse outgroup.
The Journal of heredity    October 15, 2003   Volume 94, Issue 5 374-380 doi: 10.1093/jhered/esg079
Tozaki T, Takezaki N, Hasegawa T, Ishida N, Kurosawa M, Tomita M, Saitou N, Mukoyama H.The genetic relationships of seven Japanese and four mainland-Asian horse populations, as well as two European horse populations, were estimated using data for 20 microsatellite loci. Mongolian horses showed the highest average heterozygosities (0.75-0.77) in all populations. Phylogenetic analysis showed the existence of three distinct clusters supported by high bootstrap values: the European cluster (Anglo-Arab and thoroughbreds), the Hokkaido-Kiso cluster, and the Mongolian cluster. The relationships of these clusters were consistent with their geographical distributions. Basing our assumpti...
Greater genetic variability in Argentine Creole than in Thoroughbred horses based on serum protein polymorphisms.
Genetics and molecular research : GMR    September 30, 2002   Volume 1, Issue 3 261-265 
Díaz S, Dulout FN, Peral-García P.Genetic polymorphism was analyzed for five blood proteins: albumin - Al, esterase - Es, alpha(1)B-glycoprotein - Xk, transferrin - Tf and hemoglobin - Hb in 200 Thoroughbred (TB) and 124 Argentine Creole (AC) horses. Of the five systems examined, Tf and Hb were not in Hardy-Weinberg equilibrium in either breed and Es was not in equilibrium in the Creole breed. Genetic variability, estimated as average heterozygosity, was higher in AC (H = 0.585 +/- 0.131) than in TB (H = 0.353 +/- 0.065). The genetic differentiation between these two populations (F(ST)) was 0.109. Thus, of the total genetic di...
[Microsatellite analysis in a population of Baudet du Poitou donkeys].
Schweizer Archiv fur Tierheilkunde    September 13, 2002   Volume 144, Issue 8 413-417 doi: 10.1024/0036-7281.144.8.413
Schelling C, Hagger C, Pieńkowska A, Siegfried JP, Stranzinger G.A population of Baudet du Poitou donkeys was genetically characterized using microsatellites. The results were used to verify the pedigrees and to estimate the genetic variability. It could be confirmed that a equine parentage test kit works well for donkeys and that by using 13 microsatellites more than 99% of wrong pedigree informations would be detected. The genetic variability was comparable to a representative group of Baudet du Poitou donkeys in France.
Evaluation of factors affecting individual assignment precision using microsatellite data from horse breeds and simulated breed crosses.
Animal genetics    July 26, 2002   Volume 33, Issue 4 264-270 doi: 10.1046/j.1365-2052.2002.00868.x
Bjørnstad G, Røed KH.Assignment tests have been utilized to investigate population classification, measure genetic diversity and to solve forensic questions. Using microsatellite data from 26 loci genotyped in eight horse breeds we examined how population differentiation, number of scored loci, number of scored animals per breed and loci variability affected individual assignment precision applying log likelihood methods. We found that both genetic differentiation and number of scored loci were highly important for recognizing the breed of origin. When comparing two and two breeds, a proportion of 95% of the most ...
Genetic diversity of Cheju horses (Equus caballus) determined by using mitochondrial DNA D-loop polymorphism.
Biochemical genetics    July 26, 2002   Volume 40, Issue 5-6 175-186 doi: 10.1023/a:1015884125507
Yang YH, Kim KI, Cothran EG, Flannery AR.We used sequence polymorphism of the mitochondrial DNA D-loop (968 bp excluding the tandem repeat region) to determine genetic diversity of horses inhabiting Cheju (a southern island of Korea). Seventeen haplotypes with frequencies from 1.5 to 21.5% were found among 65 Cheju horse samples. Genetic diversity (h) of the 17 haplotypes was calculated to be 0.91, indicating that the extant Cheju horse population consists of diverse genetic groups in their maternal lineage. Phylogenetic analysis showed that 17 types of Cheju (D-loop sequences determined), 5 Mongolian, 6 Arabian, 3 Belgian, 2 Tsushim...
Transient immune suppression of inapparent carriers infected with a principal neutralizing domain-deficient equine infectious anaemia virus induces neutralizing antibodies and lowers steady-state virus replication.
The Journal of general virology    May 25, 2002   Volume 83, Issue Pt 6 1353-1359 doi: 10.1099/0022-1317-83-6-1353
Craigo JK, Leroux C, Howe L, Steckbeck JD, Cook SJ, Issel CJ, Montelaro RC.The genetic variation of equine infectious anaemia virus (EIAV) clearly affects the antigenic properties of the viral envelope; however, effects on immunogenicity remain undefined, although widely assumed. Here, the immunogenicity is reported of a novel, neutralization-resistant, pony-isolate envelope EIAV(PV564DeltaPND) that contains a 14-residue deletion in the designated principal neutralizing domain (PND) of the gp90 protein. Two ponies inoculated with a chimeric virus, EIAV(DeltaPND), containing the EIAV(PV564DeltaPND) envelope in a reference provirus strain, remained asymptomatic through...
The genetic structure of Spanish Celtic horse breeds inferred from microsatellite data.
Animal genetics    February 26, 2000   Volume 31, Issue 1 39-48 doi: 10.1046/j.1365-2052.2000.00591.x
Cañon J, Checa ML, Carleos C, Vega-Pla JL, Vallejo M, Dunner S.Partition of the genetic variability, genetic structure and relationships among seven Spanish Celtic horse breeds were studied using PCR amplification of 13 microsatellites on 481 random individuals. In addition, 60 thoroughbred horses were included. The average observed heterozygosity and the mean number of alleles were higher for the Atlantic horse breeds than for the Balearic Islands breeds. Only eight percentage of the total genetic variability could be attributed to differences among breeds (mean FST approximately 0.08; P < 0.01). Atlantic breeds clearly form a separate cluster from th...
Single nucleotide polymorphisms in the equine transferrin gene.
Animal genetics    December 28, 1999   Volume 30, Issue 6 439-443 doi: 10.1046/j.1365-2052.1999.00546.x
Brandon RB, Giffard JM, Bell K.Single nucleotide polymorphisms (SNPs) in exons 13, 15 and 16 of equine transferrin for common, rare and mutant variants were investigated. Compared with previous work a further 13 SNPs have been identified, allowing for the two previously identified clades to be subdivided into 11 groups. A combination of one or more of eight SNPs can be used to classify the equine variants into these 11 groups, since most are co-inherited. Putative sites of glycosylation in exons 13 and 16 showed no polymorphism, suggesting that presence or absence of sugar moieties does not lead to electrophoretic variation...
Genetic variation and phylogenetic analysis of open reading frames 3 and 4 of various equine arteritis virus isolates.
Advances in experimental medicine and biology    October 23, 1998   Volume 440 813-819 doi: 10.1007/978-1-4615-5331-1_106
Archambault D, Laganière G, St-Laurent G.The genetic variation in equine arteritis virus (EAV) nonstructural (NS) protein-encoding open reading frames (ORF) 3 and 4 genes was investigated. Nucleotide and deduced amino acid sequences from seven different EAV isolates (one European, one American and five Canadian isolates) and the Arvac vaccine strain were compared with those of the Bucyrus reference strain. ORF 3 nucleotide and amino acid sequence identities amongst these isolates (including the Arvac vaccine strain) and the Bucyrus reference strain ranged from 85.6 to 98.8%, and 85.3 to 98.2%, respectively, whereas ORF 4 nucleotide a...
Directional and anteroposterior asymmetry of common white markings in the legs of the Arabian horse: response to selection.
Genetica    January 1, 1997   Volume 101, Issue 3 199-208 doi: 10.1023/a:1018366122913
Woolf CM.Arabian bay horses manifest, on the average, more common white markings in their hind legs than their forelegs (anteroposterior asymmetry) and more common white markings in their left legs than their right legs (directional asymmetry). To determine if genetic variation exists for these types of asymmetry, the phenotypic response was studied in bay foals when their dams and sires were selected for the directions of fore-hind and left-right differences. In the fore-hind studies, the quantitative shifts in the bay foals were in the direction specified by the selection scheme and the observed devi...
[Preservation of genetic variation in domestic animals using biotechnical methods].
DTW. Deutsche tierarztliche Wochenschrift    August 1, 1996   Volume 103, Issue 8-9 336-340 
Falge R, Ehling C, Niemann H.The conservation of endangered breeds as live animals is at present the main national strategy of the government and breeding organizations to maintain genetic diversity. Fourty-three breeds and some old strains of cattle, pig, sheep, goat and horses are currently involved. Cryopreservation and banks for sperm, embryos or DNA are another type of genetic material which could subsequently be used for breeding and production in agriculture. Present semen banks involve 9 endangered cattle breeds and also a small amount of deep-frozen sperm of some endangered sheep and horse breeds. Only 2 embryo b...
Phylogenetic analysis of open reading frame 5 of field isolates of equine arteritis virus and identification of conserved and nonconserved regions in the GL envelope glycoprotein.
Virology    December 20, 1995   Volume 214, Issue 2 690-697 doi: 10.1006/viro.1995.0087
Balasuriya UB, Timoney PJ, McCollum WH, MacLachlan NJ.The variation and phylogenetic relationship of open reading frame 5 (ORF5) of 3 different laboratory strains of the original prototype Bucyrus strain of equine arteritis virus (EAV), the modified live virus vaccine (ARVAC, Fort Dodge Laboratories), and 18 field isolates of EAV from North America and Europe were determined by comparison of their gene sequences. The viruses differed from the published sequence by between 3 (99.6% homology) and 94 (87.8%) nucleotides and by between 3 (98.8%) and 24 (90.6%) amino acids. The field isolates differed from each other by between 2 (99.7%) and 110 (85.7...
Mitochondrial DNA sequences of various species of the genus Equus with special reference to the phylogenetic relationship between Przewalskii’s wild horse and domestic horse.
Journal of molecular evolution    August 1, 1995   Volume 41, Issue 2 180-188 doi: 10.1007/BF00170671
Ishida N, Oyunsuren T, Mashima S, Mukoyama H, Saitou N.The noncoding region between tRNAPro and the large conserved sequence block is the most variable region in the mammalian mitochondrial DNA D-loop region. This variable region (ca. 270 bp) of four species of Equus, including Mongolian and Japanese native domestic horses as well as Przewalskii's (or Mongolian) wild horse, were sequenced. These data were compared with our recently published Thoroughbred horse mitochondrial DNA sequences. The evolutionary rate of this region among the four species of Equus was estimated to be 2-4 x 10(-8) per site per year. Phylogenetic trees of Equus species demo...
FTIR analysis of the interaction of azide with horse heart myoglobin variants.
Biochemistry    June 21, 1994   Volume 33, Issue 24 7600-7608 doi: 10.1021/bi00190a013
Bogumil R, Hunter CL, Maurus R, Tang HL, Lee H, Lloyd E, Brayer GD, Smith M, Mauk AG.The interaction of azide with variants of horse heart myoglobin (Mb) has been characterized by Fourier transform infrared (FTIR), electron paramagnetic resonance (EPR), and UV-VIS absorption spectroscopy and by molecular modeling calculations. Distal histidine variants (His64Thr, His64Ile, His64Lys) and charged surface variants (Val67Arg, Lys45Glu, Lys45Glu/Lys63Glu) were included in this study. All variants, with the exception of Val67Arg, have a lower azide affinity than the wild-type protein. Analysis of the temperature dependence of the FTIR spectra (277-313 K) revealed that the wild-type ...
Genomic variability among globally distributed isolates of equine arteritis virus.
Veterinary microbiology    September 1, 1992   Volume 32, Issue 2 101-115 doi: 10.1016/0378-1135(92)90099-f
Murphy TW, McCollum WH, Timoney PJ, Klingeborn BW, Hyllseth B, Golnik W, Erasmus B.Equine arteritis virus (EAV), a non-arthropod borne togavirus, has been shown to have a global distribution. To date, no major antigenic variation has been demonstrated between EAV isolates from different geographic origins. In this study, the genomic RNA of EAV isolates obtained from horses of different breeds in various countries around the world was oligonucleotide fingerprinted. Comparisons of these fingerprints were used to determine the extent of genomic variation among such isolates. Comparisons among isolates from North American horses revealed, for the most part, oligonucleotide homol...
[Genetic differentiation between breeds of horses by polymorphic blood protein loci].
Genetika    April 1, 1992   Volume 28, Issue 4 152-165 
Dubrovskaia RM, Starodumov IM, Bannikova LV.The estimation of genetic differentiation between 27 horses breeds originated in USSR, based on serum proteins polymorphism (loci Tf, Al, Es) is shown. Genetical variability among aborigine breeds is higher then among cultural ones. The erosion of gene pool of Przewalski's Horse is explained by special history of this population and a few horses in analyzing group. Genetic distances reflect the directions and intensity of breeding. High genetical distances between Przewalski's Horse, Shetland Pony and other horses obtained could be explained by overcoming the "bottle neck" of selections in bre...
Cloning of highly polymorphic microsatellites in the horse.
Animal genetics    January 1, 1992   Volume 23, Issue 2 133-142 doi: 10.1111/j.1365-2052.1992.tb00032.x
Ellegren H, Johansson M, Sandberg K, Andersson L.We have isolated equine microsatellites by screening a genomic library with (TG)n and (TC)n probes. TG microsatellites were found to be more abundant than TC repeats, with an estimated frequency of one per 100,000bp. Sequence analysis of eight TG-positive clones revealed varying structures of the repeat regions; perfect stretches of TG repeats, imperfect stretches of TG repeats and compound regions of TG and TC repeats. Five loci were analysed by PCR and showed extensive polymorphism; three to seven alleles and heterozygosities of 0.40-0.76 were observed when screening 20-30 unrelated individu...