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A Proteomic Approach to Identify Biomarkers of Growth Hormone and Aging (312 Pp.)

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A Proteomic Approach to Identify Biomarkers of Growth Hormone and Aging (312 Pp.) A Proteomic Approach to Identify Biomarkers of Growth Hormone and Aging A dissertation presented to the faculty of the College of Arts and Sciences of Ohio University In partial fulfillment of the requirements for the degree Doctor of Philosophy Juan Ding August 2009 © 2009 Juan Ding. All Rights Reserved. 2 This dissertation titled A Proteomic Approach to Identify Biomarkers of Growth Hormone and Aging by JUAN DING has been approved for the Department of Biological Sciences and the College of Arts and Sciences by John J. Kopchick Goll-Ohio Professor of Molecular Biology Benjamin M. Ogles Dean, College of Arts and Sciences 3 ABSTRACT DING, JUAN, Ph.D., August 2009, Biological Sciences A Proteomic Approach to Identify Biomarkers of Growth Hormone and Aging (312 pp.) Director of Dissertation: John. J. Kopchick The growth hormone (GH)/ insulin-like growth factor-1 (IGF-1) axis is an important regulator of longitudinal growth, metabolism and aging. The physiological role of GH is manifested in mouse models with altered GH signaling. For example, bovine (b) GH transgenic mice are giant, lean, insulin resistant and short-lived. In contrast, GH receptor (R) gene knockout (GHR-/-) mice are dwarf, obese, insulin sensitive and long- lived. In order to uncover the biomarkers of GH and aging, two-dimensional gel electrophoresis (2-DE) has been used to analyze the mouse plasma proteome. The bGH, GHR-/- mice and their respective controls at different ages were analyzed. Female GHR- /- and control mice were also analyzed for gender differences. Several biomarkers of aging and GH actions have been discovered. Biomarkers for aging include proteins whose plasma levels increased (isoform(s) of transthyretin (TTR), haptoglobin (Hp), immunoglobulin kappa chain) and decreased (isoform(s) of albumin, serum amyloid A-1 (SAA-1) and peroxiredoxin-2). Biomarkers for chronic GH action include upregulation of apolipoprotein (apo) E, Hp, mannose-binding protein-C, specific isoform(s) of alpha-2 macroglobin (a2m), retinol-binding protein-4 (RBP-4), and downregulation of apoA4, TTR, specific isoforms of apoA1, a2m and RBP-4. Importantly, isoforms of Hp and clusterin increased markedly in bGH mice during aging compared to wild type (WT) mice, possibly indicating an accelerated aging phenotype of 4 these mice. Gender-specific biomarkers include plasma proteins that increased such as isoform(s) of apoE and RBP-4 or decreased such as isoform(s) of clusterin, albumin, Hp and hemoblogin-β in females. Interestingly, some proteins showed no change during male aging but significant increase (isoform(s) of apoA1, apoA4, TTR and albumin) or decrease (isoform(s) of RBP-4 and albumin) during female aging. In addition, several proteins (Hp, apoE and RBP-4) showed an interaction between genotype (GHR-/- and WT) and gender, suggesting interplay between GH and sex. Finally, short-term effects of GH/IGF-1 were studied in mice injected with GH and/or IGF-1. GH injection resulted in upregulation of isoform(s) of apoE, Hp, RBP-4, SAA-1, albumin and a2m; and downregulated isoform(s) of apoA4 and clusterin. For IGF-1, isoform(s) of Hp, albumin and TTR were upregulated. Further, biomarkers differentiating between GH and IGF-1 actions have been revealed, including isoform(s) of apoE and TTR. The biomarkers of GH/IGF-1 injection will provide potential candidates for detecting GH and IGF-1 doping. Approved: _____________________________________________________________ John J. Kopchick Goll-Ohio Professor of Molecular Biology 5 DEDICATION This work is dedicated to my mother, Shi Hongxia, who passed away in the summer 2005 when I was in the USA pursuing my Ph.D. degree, a Pacific Ocean away. I wish I could have been there with you in those last few days. Losing you has been the hardest thing in my life. Sorry, mother, for not being there for you, while you had been there for me for 23 years. May you rest in peace! 6 ACKNOWLEDGEMENTS I would like to give my special thanks to my advisor, Dr. John Kopchick. It has been a great pleasure and honor to work with him, a renowned scientist as well as a great mentor, or even a father, who has taught me dedication, profession and perfection in research, writing and everything else in life. I would also extend my thanks to my committee members, for guidance and support in the past four years. Dr. Darlene Berryman has helped me a lot, both in dissertation and everyday lab life; and I admire her as a scientist and a woman. Dr. Xiaozhuo Chen, the faculty advisor for Chinese students, has kindly helped me in transition to graduate student life in USA. He has also provided insight in understanding my research in relation with obesity and diabetes, both subjects are his specialty. Last, but not least, Dr. Calvin James, who always put me at ease with his kind character, who always raised the ‘tough’ questions in molecular biology, reminding me to look at the big picture outside the limitation of my proteomic work. I would also like to thank my colleagues. Elahu Gosney and Amanda Palmer provided the mice necessary for this work. Dr. Edward List taught me careful design of animal experiments. I thank my fellow student Lucila Sackmann-Sala for much help and support. I thank Dr. Shigeru Okada and Dr. Bruce Kelder for kind guidance. And finally, thanks to all my lab mates who helped make the lab life at EBI much richer than just ‘lab life’. In addition to research, I have also enjoyed teaching undergraduate students. Thanks to Drs Joan Cunningham, Lorie LaPierre and John Zook, I have learned the art of 7 teaching and passing on not only knowledge, but also enthusiasm to others. Dr Robert Colvin, the MCB chair, has given me guidance over the years; many thanks to him as well. I would also like to take this opportunity to acknowledge fundings from Department of Biological Sciences, Molecular and Cellular Biology program, Diabetes Research Initiative, World Anti-Doping Agency and State of Ohio’s Eminent Scholars Program for support of my work. Lastly, I want to thank my parents, my brother and my husband Yueran Yan. They have always been there for me through the laughter and the tears. Thanks to them, I have had more laughter than tears!! 8 TABLE OF CONTENTS Page Abstract ............................................................................................................................... 3 Dedication ........................................................................................................................... 5 AcknowledgEments ............................................................................................................ 6 List of Tables .................................................................................................................... 11 List of Figures ................................................................................................................... 12 List of Abbreviations ........................................................................................................ 15 Chapter I. Background ...................................................................................................... 17 Review of GH and Mouse Models with Altered GH Signaling ................................... 17 GH/IGF-1 axis .......................................................................................................... 17 Characteristics of GH transgenic mice ..................................................................... 23 Characteristics of GHR-/- mice ................................................................................ 30 Overview of Plasma Proteome and Biomarkers ........................................................... 34 Plasma proteome ....................................................................................................... 34 Proteomic techniques to study plasma proteome ...................................................... 37 Plasma proteins as biomarkers .................................................................................. 43 Mouse plasma proteome ........................................................................................... 44 Introduction to 2-DE ..................................................................................................... 46 Sample preparation ................................................................................................... 46 Contaminant removal ................................................................................................ 47 Protein quantification, solubilization/denaturation/reduction/alkylation .................. 48 Electrophoresis .......................................................................................................... 49 Staining of protein spots ........................................................................................... 50 Plasma as a sample for 2-DE .................................................................................... 50 Review of GH Doping and Detection ........................................................................... 53 Recombinant human GH doping in sports ................................................................ 53 Current approaches to detect GH doping .................................................................. 55 Novel biomarkers for GH are needed ....................................................................... 58 Chapter II. Research Objectives ....................................................................................... 61 Chapter III. Biomarkers of
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