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The Effect of Β-Alanine Supplementation on Neuromuscular Performance

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The Effect of Β-Alanine Supplementation on Neuromuscular Performance The Effect of β-alanine Supplementation on Neuromuscular Performance by Rebecca Louise Jones (née Stannard) A thesis submitted in partial fulfilment of the requirements of Nottingham Trent University for the degree of Doctor of Philosophy May 2017 Copyright statement Copyright statement This work is the intellectual property of the author. You may copy up to 5% of this work for private study, or personal, non-commercial research. Any re-use of the information contained within this document should be fully referenced, quoting the author, title, university, degree level and pagination. Queries or requests for any other use, or if a more substantial copy is required, should be directed in the owner(s) of the Intellectual Property Rights. i Publications Abstract Carnosine (β-alanyl-L-histidine), a histidine containing dipeptide, is one of the most abundant small-molecular compounds in human skeletal muscle. Supplementation with the rate limiting amino acid, β-alanine, has resulted in significant improvements to high-intensity exercise performance. The role of carnosine as an intracellular pH buffer is undisputable, yet other physiological roles have been proposed, including the potential influence of increased carnosine content on regulation of skeletal muscle calcium (Ca2+) kinetics. The movement of Ca2+ is vital during both skeletal muscle contraction and relaxation phases. The overall aim of this thesis was to investigate the effect of β-alanine supplementation on voluntary and electrically evoked contractile properties of in-vivo human skeletal muscle. To examine this research question, there were several aims of this thesis, initially to examine the effect of β-alanine supplementation on intrinsic in-vivo isometric knee extensor force production and skeletal muscle contractility in both fresh and fatigued human skeletal muscle in young (Studies 1 and 2; Chapters 4 and 5) and older (Study 3; Chapter 6) adults. The distribution of the carnosine molecule across subcellular fractions within rat skeletal muscle tissue was explored, as well as the impact of increased carnosine availability on ATPase activity, a measure associated with skeletal muscle relaxation, estimated by Pi generation. In young adults, 28-days of β-alanine supplementation did not significantly influence voluntary and evoked force responses, or the force-frequency relationship, the in-vivo analogue of the force-Ca2+ relationship, in either fresh (Studies 1 and 2; Chapters 4 and 5) or fatigued (Study 2; Chapter 5) skeletal muscle. Furthermore, older adults experiencing pre-existing declines in skeletal muscle function due to ageing, demonstrated no beneficial effect of 28-days β-alanine supplementation on voluntary or electrically evoked skeletal muscle contractions (Study 3; Chapter 6). In young adults, there was, however, a significant decline in skeletal muscle half-relaxation time (HRT) during electrically evoked octet contractions, resting and potentiated twitches (Studies 1 and 2; Chapters 4 and 5). Two possible steps influence skeletal muscle relaxation speed include the Ca2+ removal from the myoplasm and Ca2+ dissociation from troponin followed by cross-bridge detachment. Based on the in-vivo research, it was proposed that there was a direct or indirect mechanism associated with activity of the sarcoplasmic/endoplasmic reticulum Ca2+ ATPase (SERCA) pump, the proposed rate-limiting step of muscle relaxation. In-vitro analysis of ATPase activity demonstrated that SERCA activity was unaffected by increased carnosine concentrations, although there was a significant increase in overall ATPase activity (Study 4; Chapter 7). The results in this thesis showed that β-alanine supplementation was effective in improving skeletal muscle HRT in young adults, although not in healthy older adults. The exact mechanism associated with the in-vivo decline in skeletal muscle HRT remains unclear, yet raising the availability of carnosine in-vitro, does increase overall ATPase activity, although not Ca2+-dependent or SERCA activity. ii List of equations Publications Journal articles (Appendix 2) Jones, R.L., Barnett, C.T., Davidson, J., Maritza, B., Fraser, W.D., Harris, R.C., & Sale, C. (2017). β-alanine supplementation improves in-vivo fresh and fatigued muscle relaxation speed. European Journal of Applied Physiology, 1-13. DOI: 10.1007/s00421-017-3569-1 Hannah, R., Stannard, R.L., Minshull, C., Artioli, G.G., Harris, R.C., Sale, C. (2015). β-alanine Supplementation Enhances Human Skeletal Muscle Relaxation Speed but Not Force Production Capacity. Journal of Applied Physiology. 118 (5), 604-612. DOI: 10.1152/japplphysiol.00991.2014 Poster presentations Stannard, R.L., Hannah, R., Minshull, C., Artioli, G.G., Harris, R.C., Sale, C. (2015). β-alanine Supplementation Enhances Human Skeletal Muscle Relaxation Speed but Not Force Production Capacity. In Proceedings of the 62nd America College of Sport Medicine (ACSM) Annual Meeting, San Diego, USA. Stannard, R.L., Hannah, R., Minshull, C., Artioli, G.G., Harris, R.C., Sale, C. (2015). β-alanine Supplementation Enhances Human Skeletal Muscle Relaxation Speed but Not Force Production Capacity. Science and Technology Annual Research (STAR) conference, Nottingham Trent University. Oral presentations Stannard, R.L., Barnett, C.T., Davidson, J., Maritza, B., Fraser, W.D., Harris, RC., & Sale, C. (2016). β-alanine supplementation improves in-vivo fresh and fatigued muscle relaxation speed. In Proceedings of the 21st Annual European College of Sport Science Annual Conference, Vienna, Austria. Stannard, R.L. (2015). Effects of β-alanine supplementation on human skeletal muscle contractile properties and voluntary muscle performance. STAR conference, Nottingham Trent University. iii Acknowledgements Acknowledgements This thesis is the accumulation of countless hours (years!) worth of work. It is an immense pleasure to acknowledge the roles of those individuals who were instrumental for the completion of this Ph.D. research. Firstly, to Professor Craig Sale, without your endless guidance, encouragement, critical analysis, and expertise, I would never have started, let alone completed this research! Thank you for the numerous hours spent discussing, reading and reviewing every aspect of this thesis. You have provided me with so many brilliant opportunities and experiences over the last 5 years. My hope is to take a small part of your vast knowledge away with me. I would also like to acknowledge my Ph.D. supervisors, Dr. Cleveland Barnett and Prof. William Fraser for your helpful contributions to this programme of work. Further thanks go to all those who participated in my experimental research; I am very grateful for your time, commitment and willingness to encounter these unpleasant experiences on more than one occasion. To all those who have steered me in the right direction, given me advice and new opinions, taught me how to use equipment, answered my endless (and repeated) questions, helped with data collection, and given me a spare 5 minutes, even when you didn’t have it, this immense thank you goes to you; Professor Roger Harris, Dr. Ruth James, Dr. Chris Lloyd-Mills, Dr. Carl Nelson, Dr. Guilherme Artioli, Dr. Eimear Dolan, Dr. Bryan Saunders, Dr. Ricci Hannah, Dr. Kirsty Elliott- Sale; Andrew Marr, Michael Shaw, Joel Davidson and Billy Maritza. To all the staff and academics at Nottingham Trent University, it has been an absolute pleasure working with you. To all my family and friends who have helped me in so many ways to achieve this dream. You haven’t always fully understood what I was doing/talking about, yet, you have always been there to listen and guide me. I could not have done this without you all. Finally, to my husband, David Jones, you believe in me when I struggle to believe in myself, with your unwavering constant support and encouragement, you have allowed me to become the person I am today, I am eternally grateful, thank you. iv Contents Contents Copyright statement .............................................................................................................................i Abstract .............................................................................................................................................. ii Publications ....................................................................................................................................... iii Journal articles (Appendix 2) ...................................................................................................... iii Poster presentations .................................................................................................................... iii Oral presentations ....................................................................................................................... iii Acknowledgements ............................................................................................................................iv Contents .............................................................................................................................................. v List of tables ........................................................................................................................................ x List of figures ................................................................................................................................... xii List of equations ...............................................................................................................................xix
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