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A Thesis Present.Ed to the Faculty of University of Manitoba in Partíal

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A Thesis Present.Ed to the Faculty of University of Manitoba in Partíal ENERGY METASOLISM IN PROGRESSIVE MUSCIILAR DYSTROPHY-- STI]DIES ON OXIDATIVE PHOS?HORY].ATION OF GENETICALLY DETERMINED MUSCIILAR DYSTROPHY ]N MICE AND HAMSTERS. A Thesis Present.ed to the Faculty of Graduate Studies and Research of the University of Manitoba In PartÍal Fulfillment of the Requirements for the Degree of Doctor of Philosophy by Klaus trnlrogemann Department of Biochemistry, Faculty of Medicine r969 c Klaus lnlrogemann 1969 1. TAsLE OF CONTENTS PAGE TABLE OF CONTENTS 2 LIST OF TABLES 9 LIST OF FIGURES T2 ACKNOT^TLEDGEMENTS t3 ABSTRACT 15 ORGANIZATION OF THE THESIS ... L9 GLOSSARY 20 ÏNTRODUCTION . 23 LITERATURE REVIEI^I 2B Tíssue high energy phosphate content 29 Glycolytic capacity and glycolytic enz)¡me activity in dystrophÍc muscle .. 32 Studies on mitochondrial structure and functÍon in muscular dystrophy . 34 SLudies on oxídative phosphorylation in muscular dystrophy 31 MATERIALS AND METHODS .. 39 A. MATERIALS (a) Chemicals 40 (b) Animals 44 B. METHODS (a) Assessment of the stage of the disease 45 X. Mice . 45 Hamsters 45 Iß. 1. Age. 45 2. Body weight, heart weight, heart weight body weight ratio 45 -2- TABLE OF CONTENTS (CONT'D) PAGB J, Macroscopic description of heart and skeletal muscle pathology . 46 His topathology 46 5. Serum creatine phosphokinase determination . 47 6. Protein, non-collagen protein, collagen protein in skeletal muscle 48 7. Acid phosphatase activíty in skeletal mus cle 49 8. f'gLucuronídase activity in skeletal mus cle 49 9. Cathepsin activity in skeleËal muscle 50 10. DNA and RNA content in skeleËal- muscle 51 (b) Dissection of animals and preparation of mitochondria . 52 L. Iulouse skeletal muscle mitochondria . 53 2. Mouse skeletal muscle mitochondria prepared in the presence of albumin . ,. s4 J, Hamster heart mitochondr|a . s4 4. Hamster skeletal muscle mitochondria: Standard Proteinase Preparation . .....;. 54 5. Hamster skeletal muscle mitochondria: Standard Proteinase Preparatíon wÍt.h l% albumin ... 55 6. Hamster skeletal muscle mitochondria: Modífied Proteinase Preparation .. 55 7. Hamster skeletal muscle mitochondria: Lochner Preparation .. 55 (.) Polarographic measurements 56 1. Apparatus 56 2. Measurements 56 -3- TABLE OF CONTENTS (CONT'D) PAGE 3. Oxygen calibration .. 59 4. Materials added durÍng the polarographic experiments 59 5. Calculations 59 (d) Polarographic measurements in the presence of a hexokinase trap . 60 (e) Manometric measurements 62 t. Oxidative phosphorylation with pyruvate/malate as substrate . 62 ) Respiration rates with palmitate as substrate . 64 3. Respiration rates with palmityl-L- carnitine as substrate . 64 (f) Assay procedures ... 65 1. ProteÍn estimation o¿,. direct 65 indirect Iß. 65 2. NADH concentration 66 3. ADP concentration 66 4. Acetyl-L-carnítine concentration 67 5. Palmityl-L-carniiine coneentration 67 (g) Substrate preparations 67 1. Acetyl-L-carnitine. 67 2, Palmityi--L-carnitine .. 67 3. Potassium palmitate 67 (h) Tissue mitochondrial content determinations 68 (i) Electron microscopic examination of isolated mitochondrLa , 69 -/,+ TABLE OF CONTENTS (CONT'D) PAGE (j) Calculations and statistics ... 69 (k) Preparation of distilled water .. .. 7A RESIILTS 7L Outline of the Results 72 A. OxídaÈive phosphorylatíon by skeletal muscle mitochondria from normal and dystrophic mice . 73 The effect of albumin . 80 Conclus ions B6 B. Oxidative phosphorylation by heart and skel-etal muscle mitochondria from normal and dystrophic hamsters 89 (a) HamsËer hearË mitochondrLa . 89 Characteristics of the dystrophic hamsters ... 90 Oxidative phosphorylation by heart mitochondria of dystrophic hamsters with a mean age of 110 days 90 Respiration with DL-ct-glycerophosphate and NA-DH as substrate 9B Oxidative phosphorylation by heart mitochondria from hamsters with a mean age of 160 days 1-05 The effect of a hexokinase trap L07 Oxidative phosphorylation by normal and dystrophic hamsËer mitochondria--isolation of the organelles by a different person 110 Oxidalive phosphorylation by heart mitochondria from normal and dystrophic hamsters more than 200 days old . IL4 The effect of higher reaction temperature LL4 1, With the standard substrate combination pyruvate/malaËe LL4 2. i{ith Dl-l3-hydroxybutyrate LLl Oxidative phosphorylation with palmityl-L- carnitine as substrate . L22 -5- TABLE OF CONTENTS (CONT'D) PAGE Mitochondrial yields . L24 Sunmary of the results obtaÍned from hamster heart mitochondria . .... L26 (b) Hamster skeletal muscle mitochondria . ... L27 Oxídative phosphorylaEion by skeletal muscle mitochondria from normal and dystrophic hamsters with a mean age of 110 days ..., LZB Respíration with Dl-ct-g lycerophosphate as substrate . 135 Respiration wÍth NADH as substrate .. 138 Tissue mitochondrial content of normal and dystrophic hamster skeletal muscle ... 138 Oxidative phosphorylation by skeletal muscle mitochondria from hamsters with a mean age of 160 days. Isol-ation of the organelles by a different person ..... 1"42 The effect of a hexokínase trap .. .. L43 Oxidative phosphorylaËion by skeletal muscle mitochondria from severely affecled dystrophic hamsters. Comparison of different methods of mitochondrial preparations .. .. L46 1. Standard Proteinase Preparation . .. L46 Recoupling of uncoupled oxidative phos- phorylation in dystrophic hamster mito- chondria by NI9CL2 L49 ) Standard Proteinase Preparation withalbumin. .L54 J. Lochner Preparation .. L57 Comparison of polarographic and manomeEric techniques for the determination of oxidative phosphorylation parameters ... .. L66 Oxídative phosphorylation by hamster skeletal muscle mitochondria prepared by the Modified Proteinase Procedure .. LlL -6- TA3LE 0F CONTENTS (CONT!D) PAGE Reproduction of the manometrÍc experiments of Lochner using hamsters with a mean age oÍ. 2L5 days L7 4 Respiration with palmitate and palmityl-L- carnitine as substrate . L75 Confirmation of the manometrically detected mitochondrial respíration defect by polaro- graphic techniques ... L79 A single or a t\'/ofold defect? LB2 The localízatíon of the respiration defecl t84 1. Impaired respiration wiLh acetyl-L-carnitine LB4 2. Normal respiratíon with succinate 185 3. Normal respiration with NADH . T87 Is the observed mitochondríal respiration defect also present in viri-g? 189 Studies on the tissue compositíon of normal and dystrophic muscle L92 1, Collagen, non-collagen protein, DNA, RNA L92 2. Hydrolytic enz)¡rne activities ... 194 Sunrnary of the results obËained from hamster skeletal muscle 198 DISCUSSION ... 200 On theoretical AnP/0 ratios and the relation between ADP/O and respiratory control ratios .. zOL Oxidative phosphoryLaËion by skeletal muscle mitochondria from normal and dystrophic mice .. 208 Oxidative phosphorylation by heart mitochondria from normal and dystrophic hamsters . .. ... 2L0 Oxidative phosphorylation by skeletal muscle mitochondria from normal and dystrophic hamsters ... 2L4 -7 TABLE OF CONTENTS (CONT'D) PAGE LIST OF REFERENCES .,. 222 APPENDIX 247 -B- II. LIST OF TABLES TABLE PAGE Characteristics of dystrophic mice and their littermate controls ... 74 Characteristics of mice used in the preparation of mitochondria in the presence of albumin ... 75 3a RespÍration and oxidative phosphorylation by mouse muscle mitochondría . 79 4a Respiration and oxidative phosphorylation by muscle mitochondria prepared from 4 control mice and the effect of ATP 81 5a Respiration and oxidative phosphorylation by mouse muscl-e mitochondría prepared in the presence oî. L% albumin B3 6a Yield of mouse muscle mitochondria prepared in the presence of. L% albumin 87 7a Characteristics of the hamster Group t ... 9L Ba Respíration and oxidative phosphorylation by heart mitochondria from hamsters of Group I (subsÈrate added prior to firsE ADP) 93 9a Respiration and oxidative phosphorylation by heart mitochondria from hamsters of Group 1 (indígenous substrate depleted) 97 10 Multiple comparison of the means of the different periods of Tables 8 and 9 99 lla Respiration by heart mitochondria from hamsters of Group I (substrates: DL-ct-glycerophosphate and NADH) 100 L2a Characteristics of the hamster Group 2 ... 106 l3a Respiration and oxidative phosphorylation by heaÉ mitochondria from hamster Group 2 . 1-08 L4a Characteristics of Ëhe hamster Group 3 ... 111 15a Respiration and oxidative phosphorylation by heart mitochondria frsm hamsters of Group 3 ... 113 L6a Characteristics of the hamster Group 6 .., 1l-5 -9- Lrsr 0F TABLES (CONT'D) TABLE PAGE L7 a Respiration and oxidative phosphorylation by heart mitochondria from hamsters of Group 6 (substrate: pyruvate/malare) 116 18a Respiration and oxidative phosphorylation by heart mitochondria from hamsters of Group 6 (substrate: Dl-p-hydroxyburyrare) ll8 L9a Respiration and oxidative phosphorylation by heart mitochondria from hamsters of Group 6 (substrate: palmityl-L-carnÍtíne -l- malate) L23 20a Yield of hamster hearr mitochondría . LZs ZLa Respíration and oxidative phosphorylation by skeletal muscle mitochondria from hamsters of Group 1 (substrate added príor to first ADp) 131 22a Respiration and oxídative phosphorylation by skeletal muscle mitochondria from hamsters of Group 1 (indigenous substrate depleted) 133 23 Multiple comparison of the means of the dífferent periods of Tables 2L and 22 L34 24a Respiratíon by skeletal muscle miËochondria from hamsters of Group I (substrates: Dl-ct-glycerophosphate and NADH) 136 25a Succinic dehydrogenase activíty and tissue mitochondrial content in skeletal muscle of hamsters of Group I 140 : . 26a Yield of hamster skeletal
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