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Taurine Depletion and Schwann Cell Dysfunction in Diabetic Neuropathy

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Taurine Depletion and Schwann Cell Dysfunction in Diabetic Neuropathy Taurine Depletion and Schwann Cell Dysfunction in Diabetic Neuropathy By Trevor Askwith A thesis submitted to The University of Birmingham for the degree of DOCTOR OF PHILOSOPHY Department of Clinical and Experimental Medicine University of Birmingham Medical School The University of Birmingham June 2010 University of Birmingham Research Archive e-theses repository This unpublished thesis/dissertation is copyright of the author and/or third parties. The intellectual property rights of the author or third parties in respect of this work are as defined by The Copyright Designs and Patents Act 1988 or as modified by any successor legislation. Any use made of information contained in this thesis/dissertation must be in accordance with that legislation and must be properly acknowledged. Further distribution or reproduction in any format is prohibited without the permission of the copyright holder. Abstract It is estimated that 2.6 million people in the UK suffer from diabetes, 50% of whom suffer from diabetic neuropathy. Patients with diabetes have low levels of platelet and plasma taurine and in animal models taurine supplementation ameliorates neuropathic symptoms. The mechanisms behind taurine depletion and taurine supplementation are not well understood. Schwann cells are highly vulnerable to hyperglycaemia-induced stress which plays a key role in the pathogenesis of diabetic neuropathy, however, the mechanisms behind these effects are not well understood. In these studies I have elucidated the effect of hyperglycaemia on taurine transport in isolated human Schwann cells and the mechanisms behind the beneficial effects of taurine supplementation. I demonstrated that high glucose reduces TauT expression in a dose-dependent manner and that high glucose inhibited the pro-oxidant increase in TauT expression and taurine uptake. This high glucose response was ablated by inhibition of aldose reductase, nitric oxide synthase as well as antioxidant treatment. Taurine supplementation reduced glucose-induced increases in oxidative stress, lipid peroxidation nitrosative stress and poly(ADP-ribosyl)ation and these effects were not accompanied by changes in antioxidant defence. Taurine also restored glucose-induced increases in iNOS and nNOS expression along with phospho-p38 MAPK abundance. ii Acknowledgements Firstly I would like to express my sincere thanks to Martin Stevens for his guidance and support during my PhD studies; it has been an honour to be his first UK PhD student and his clinical expertise and guidance has been much appreciated. Many thanks to Margaret Eggo for her much appreciated help, support and technical guidance through-out the duration of my time in the laboratory and also her ongoing support during the write up process. A big thank you to everyone within the laboratory. Special thanks to Wei Zeng and Sharon Hughes for their practical and technical support at the lab bench, and to Kiran Dubb and Elena Gillespie for their clinical guidance. Sincere thanks to the University of Birmingham for providing the funding for this project. I would also like to thank all my family and in-law family for their help, encouragement, and patience during my studies.Thank you to my friends for their support, in person and via email and supportive phonecalls. Special thanks to my wife Louise for her love, care and continual support during the good times and bad. Thank you for your, faith, courage, and sense of humour, helping to keep us smiling and laughing, even during the challenging days. iii Contents CHAPTER 1 INTRODUCTION ................................................................................................ 1 1.1 TYPE 1 DIABETES-INSULIN DEPENDENT DIABETES MELLITUS .............................................................. 2 1.2 TYPE 2 DIABETES ................................................................................................................................... 3 1.3 EPIDEMIOLOGY OF DIABETES ................................................................................................................ 3 1.4 COMPLICATIONS OF DIABETES ............................................................................................................... 4 1.4.1 Microvascular Complications of Diabetes ........................................................................................ 6 1.4.1.1 Retinopathy ..............................................................................................................................................6 1.4.1.2 Nephropathy ............................................................................................................................................6 1.4.1.3 Neuropathy ..............................................................................................................................................7 1.4.1.4 Diabetic Foot Ulceration .........................................................................................................................11 1.4.1.5 Vascular Deficits in Diabetic Neuropathy ...............................................................................................11 1.4.1.6 Schwann Cells in Diabetic Neuropathy .............................................................................................13 1.4.2 Metabolic Deficits of Diabetic Neuropathy .................................................................................................16 1.4.2.1 Polyol Pathway ..................................................................................................................................16 1.4.2.2 Advanced Glycated End Products .....................................................................................................19 1.4.2.3 Hexosamine Pathway ........................................................................................................................21 1.4.3 Oxidative stress ...........................................................................................................................................24 1.4.3.1 The Biochemistry of Oxidative Stress ...............................................................................................24 1.4.3.2 Oxidative Stress in Diabetes ...............................................................................................................26 1.4.3.3 Antioxidant Treatment and Neuropathy ............................................................................................27 1.4.3.4 Antioxidant Defence System .............................................................................................................29 1.4.3.4.1 Glutathione ....................................................................................................................................29 1.4.3.4.2 Superoxide Dismutase....................................................................................................................30 1.4.3.4.3 Catalase ..........................................................................................................................................31 1.4.4 Nitric Oxide ..................................................................................................................................................32 1.4.4.1 Reactive Nitrogen Species .................................................................................................................34 1.4.4.2 3-Nitrotyrosine ..................................................................................................................................34 1.4.4.3 S-nitrosylation ...................................................................................................................................36 1.4.5 Poly(ADP-ribose) polymerase.......................................................................................................................36 1.4.6 Mitogen Activated Protein Kinase ...............................................................................................................39 1.4.6 Akt/ Protein Kinase B ..................................................................................................................................42 1.5 TAURINE .............................................................................................................................................. 45 1.5.1 Taurine Biosynthesis ...................................................................................................................... 46 1.5.2 Taurine Transporter ........................................................................................................................ 47 1.5.3 Taurine Deficient Animal Models .................................................................................................. 48 1.5.4 Molecular Identity of the Taurine Transporter ............................................................................... 49 1.5.4.1 Gating of Transport ................................................................................................................................50 1.5.4.2 Regulation of Expression ........................................................................................................................50 1.5.5 Taurine Efflux Pathway .................................................................................................................. 51 1.5.6 Taurine and Diabetes ...................................................................................................................... 52 1.5.6.1 TauT Regulation in Diabetes
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