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Download (8MB) https://theses.gla.ac.uk/ Theses Digitisation: https://www.gla.ac.uk/myglasgow/research/enlighten/theses/digitisation/ This is a digitised version of the original print thesis. Copyright and moral rights for this work are retained by the author A copy can be downloaded for personal non-commercial research or study, without prior permission or charge This work cannot be reproduced or quoted extensively from without first obtaining permission in writing from the author The content must not be changed in any way or sold commercially in any format or medium without the formal permission of the author When referring to this work, full bibliographic details including the author, title, awarding institution and date of the thesis must be given Enlighten: Theses https://theses.gla.ac.uk/ [email protected] INVESTIGATIONS OF ENVIRONMENTAL AND GENETIC INFLUENCES ON EAST AFRICAN DISTANCE RUNNING SUCCESS Robert A. Scott BSc (Hons) Submitted for the degree of Doctor of Philosophy (PhD) in the Faculty of Science. University of Glasgow. March, 2006 ProQuest N um ber: 10390988 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. uest ProQuest 10390988 Published by ProQuest LLO (2017). C o pyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code Microform Edition © ProQuest LLO. ProQuest LLO. 789 East Eisenhower Parkway P.Q. Box 1346 Ann Arbor, Ml 4 8 1 0 6 - 1346 G l a s g o w "% UNIVERSITY JLIBRARY; Abstract The primary objective of these experiments was to assess the environmental and genetic contributors to east African running success. Elite Ethiopian and Kenyan athletes, as well as matched controls, consented to participate in the current series of experiments by completing a questiomiaire on their etlinic and environmental background, and by providing a buccal swab for analysis of novel and existing candidate genes for human performance. The aim of the first two studies presented in this thesis (Chapters three and four) was to compare the demographic characteristics of elite Ethiopian and Kenyan athletes with those of their source populations. This was to assess the validity of reports linking their success to altitude inhabitation and having ran long distances to school each day during childhood. In both experiments, it was found that the elite athletes were clustered in regions of high altitude, proximal to the Rift Valley. However, in Ethiopia, non-endurance athletes were also found to cluster in these areas, so the linlc between altitude and elite athlete status may not simply be as a result of haematological adaptations conferring benefit in endurance performance. It was also found that elite athletes had travelled farther to school each day as children, and more of them had done so by running. In addition, the athletes displayed a distinct ethnicity relative to the general population. The associative nature of the experiments did not allow causality to be assigned to any one factor, but the results highlight the importance of environmental factors in the determination of elite east African athlete status. The aim of Chapter five was to compare the frequency of mtDNA haplogroups between elite Ethiopian athletes with the general Ethiopian population. mtDNA is inherited in a matrilineal fashion, making it useful in population genetics. A number of studies have shown a maternal influence on the heritability of aerobic performance phenotypes, and more recent studies have suggested that mtDNA polymorphisms are associated with inter-individual variation in aerobic performance. Given the likely origin of modern humans in east Africa, Ethiopians display a wide degree of mtDNA haplogroup variation. Many of the haplogroups found in east Africa are rarely found elsewhere, while others are similar to those found commonly outwith east Africa. Given that mtDNA codes for components of mitochondria, mtDNA is a potential candidate gene for physical performance. However, elite Ethiopian endurance athletes did not differ in their mtDNA haplogroup distribution from the control group. Elite Ethiopian athletes were not a genetically distinct group, as defined by their mtDNA. This does not support hypotheses that east African athletes arise from a limited genetic isolate. Chapter six aimed to compare the mtDNA haplogroup frequency between elite Kenyan athletes and the Kenyan control group. Kenya also displays a wide degree of mtDNA variation, but displays a different haplogroup distribution from Ethiopia, with a higher frequency of typically African ‘L’ haplogroups (Ethiopia = 54 %, Kenya = 90 %). Kenyan athletes displayed a distinct distribution from controls. National standard athletes displayed an excess of M haplogroups relative to controls, and a tendency toward a lower frequency of L2 haplogroups, while international Kenyan athletes displayed an excess of LG haplogroups, a lower frequency of L2 haplogroups and a tendency toward an excess of M haplogroups. Despite the associations of mtDNA haplogroups with elite Kenyan athlete status, the elite athletes retained a similar level of diversity to the general population, which attests to the complexity of the elite athlete phenotype. Findings of ethnic variation in mtDNA haplogroup frequency meant that influences of population stratification could not be ruled out. The experiment in Chapter seven compared the frequency of Y chromosome haplogroups in elite Ethiopian athletes with that of the general Ethiopian population. Y chi'omosomal DNA is inherited solely down the paternal line and is confined solely to males. Y chromosome variants have been associated with a diverse range of phenotypes in health and disease, making the Y chr omosome a viable candidate gene for physical performance in males. Similar to mtDNA, the non-recombinant pattern of inheritance makes the Y cliromosome useful in population genetics to determine the ancestry of individuals or populations. Certain Y chi'omosome haplogroups were associated with elite Ethiopian athlete status. Athletes showed an excess of E3*, a lower frequency of E3b, and a tendency toward an excess of K(xP) haplogroups relative to controls. Although no influence of region or ethnicity was found on haplogroup distribution, effects of population stratification could not be ruled out. These results suggest that Y chromosome variation may influence elite Ethiopian athlete status. Perhaps the most studied variant of the candidate genes for human performance is the ACE I/D polymorphism. A number of studies have associated the ACE I allele with endurance performance, while the D allele has been associated with power type performance. The experiment in Chapter eight aimed to compare ACE I/D genotype frequencies between elite Ethiopian and Kenyan athletes and their source populations. It was found that male Ethiopian marathon runners showed an excess of II genotypes relative to controls. However, this association was not replicated in 5-10 km Ethiopian runners, or in Kenyan distance runners. It is unlikely that the I/D polymorphism is functional in influencing variation in the ACE phenotype. Other variants in the ACE gene are more strongly associated with serum ACE levels in African populations, where levels of linkage disequilibrium differ from Caucasian populations. For this reason, A22982G was genotyped in elite athletes and controls. This variant showed a stronger association with serum ACE levels than the I/D polymorphism in the Kenyan control group. However, there was no association between A22982G genotype and elite athlete status in either Ethiopian or Kenyan athletes. These results do not support a role for ACE gene variation in the determination of elite East African athlete status. 'S 5 Table of Contents Abstract....................................................................................................... 2 Table of Contents...................................................................................................................... 5 List of Tables.................................................................................................................................9 List of Figures..............................................................................................................................10 Acknowledgements...................................................................... 14 Author's Declaration ..................................................................................................... 16 List of Abbreviations.................................................................................................................17 Chapter 1 General Introduction....................................................................................... 19 1.1 East African Running........................................................................................................20 1.2 Explanations for east African domination...................................................................... 24 1.2.1 Psychological and socio-cultural explanations...................................................... 24 1.2.2 Physiological explanations........................................................................................25
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