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Department of Sports and Exercise Medicine Clinicum Faculty of Medicine University of Helsinki Helsinki, Finland OXYGEN DELIVERY AND UTILIZATION DURING ACUTE DYNAMIC EXERCISE EFFECTS OF POLYCYSTIC OVARY SYNDROME, TYPE 1 DIABETES, AND EXERCISE TRAINING Antti-Pekka Rissanen ACADEMIC DISSERTATION To be presented, with the permission of the Faculty of Medicine of the University of Helsinki, for public examination in Auditorium PIII, Porthania, City Centre Campus, University of Helsinki, on May 20th 2017, at 10 o’clock in the morning. Helsinki 2017 Supervised by Docent Juha Peltonen, Ph.D. Department of Sports and Exercise Medicine Clinicum Faculty of Medicine University of Helsinki Helsinki, Finland Clinic for Sports and Exercise Medicine Foundation for Sports and Exercise Medicine Helsinki, Finland Professor Heikki Tikkanen, M.D., Ph.D. Institute of Biomedicine School of Medicine University of Eastern Finland Kuopio, Finland Department of Sports and Exercise Medicine Clinicum Faculty of Medicine University of Helsinki Helsinki, Finland Clinic for Sports and Exercise Medicine Foundation for Sports and Exercise Medicine Helsinki, Finland Reviewed by Senior Research Fellow J. Chris Baldi, Ph.D. Department of Medicine Dunedin School of Medicine University of Otago Dunedin, New Zealand Docent David E. Laaksonen, M.D., Ph.D., M.P.H. Internal Medicine Institute of Clinical Medicine Kuopio University Hospital Kuopio, Finland Opponent Professor Robert Petrella, M.D., Ph.D. Department of Family Medicine and Kinesiology Schulich School of Medicine and Dentistry Western University London, Canada Dissertationes Scholae Doctoralis Ad Sanitatem Investigandam Universitatis Helsinkiensis ISBN 978-951-51-3177-5 (paperback) ISSN 2342-3161 (print) ISBN 978-951-51-3178-2 (PDF) ISSN 2342-317X (online) Cover artwork: Liia Rissanen Press: Unigrafia Oy (Yliopistopaino Helsinki), Helsinki, Finland, 2017 http://ethesis.helsinki.fi Man, he took his time in the sun Had a dream to understand A single grain of sand - from the Nightwish masterpiece The Greatest Show on Earth (lyrics by Tuomas Holopainen) CONTENTS CONTENTS ..................................................................................................................... 4 LIST OF ORIGINAL PUBLICATIONS ........................................................................... 8 ABBREVIATIONS .......................................................................................................... 9 ABSTRACT ..................................................................................................................... 11 TIIVISTELMÄ (ABSTRACT IN FINNISH) ................................................................... 13 1 INTRODUCTION ......................................................................................................16 2 REVIEW OF THE LITERATURE ............................................................................ 18 2.1 OXYGEN DELIVERY AND UTILIZATION DURING ACUTE DYNAMIC EXERCISE ......................................................................................................... 18 2.1.1 SYSTEMIC OXYGEN DELIVERY DURING ACUTE DYNAMIC EXERCISE .................................................................................................. 20 2.1.1.1 Pulmonary function ......................................................................... 20 2.1.1.2 Cardiac pump function .................................................................... 21 Cardiac cycle ................................................................................... 21 Cardiac pump function during acute dynamic exercise .............. 22 Methods to quantify cardiac pump function ................................ 25 2.1.1.3 Blood oxygen carrying capacity ...................................................... 28 2.1.2 PERIPHERAL OXYGEN DELIVERY, EXTRACTION, AND UTILIZATION DURING ACUTE DYNAMIC EXERCISE .................................................. 29 2.1.2.1 Overview of blood flow distribution and regulation at the whole- body level ........................................................................................ 29 2.1.2.2 Skeletal muscle blood flow and oxygen extraction and utilization 31 Skeletal muscle blood flow ............................................................. 31 Skeletal muscle oxygen extraction and utilization ....................... 34 2.1.2.3 Cerebral blood flow and oxygen extraction and utilization ........... 35 2.1.2.4 Near-infrared spectroscopy (NIRS) as a method to evaluate tissue- specific (im)balance between oxygen delivery and utilization ...... 36 2.1.3 PEAK OXYGEN UPTAKE (V̇ O2peak) .......................................................... 40 2.1.3.1 V̇ O2peak as a concept ........................................................................ 40 2.1.3.2 V̇ O2peak as an integrated model ....................................................... 41 2.1.3.3 V̇ O2peak as a predictor of cardiovascular disease and mortality ..... 44 4 2.1.4 EFFECTS OF ENDURANCE TRAINING ON OXYGEN DELIVERY AND UTILIZATION DURING ACUTE DYNAMIC EXERCISE ......................... 46 2.1.4.1 Effects of endurance training on systemic oxygen delivery ........... 46 2.1.4.2 Effects of endurance training on skeletal muscle blood flow and oxygen extraction and utilization .................................................. 47 2.1.4.3 Endurance training effects in the context of Fick’s theories ......... 49 2.2 POLYCYSTIC OVARY SYNDROME (PCOS) .................................................... 50 2.2.1 DEFINITION AND EPIDEMIOLOGY ...................................................... 50 2.2.2 PATHOGENESIS ...................................................................................... 50 2.2.3 CLINICAL MANIFESTATIONS ................................................................. 51 2.2.4 PCOS AND EXERCISE ............................................................................. 52 2.2.4.1 Oxygen delivery and utilization during acute dynamic exercise in PCOS............................................................................................... 52 2.2.4.2 Effects of exercise training in PCOS .............................................. 53 2.3 TYPE 1 DIABETES ............................................................................................ 53 2.3.1 DEFINITION AND EPIDEMIOLOGY ...................................................... 53 2.3.2 PATHOGENESIS ...................................................................................... 54 2.3.3 CLINICAL MANIFESTATIONS ................................................................ 55 2.3.3.1 Microvascular complications ......................................................... 56 2.3.3.2 Macrovascular complications ........................................................ 56 2.3.3.3 Diabetic cardiomyopathy ............................................................... 57 2.3.4 TYPE 1 DIABETES AND EXERCISE ........................................................ 58 2.3.4.1 Oxygen delivery and utilization during acute dynamic exercise in type 1 diabetes ................................................................................ 58 2.3.4.2 Effects of exercise training in type 1 diabetes .................................61 3 AIMS OF THE STUDY ............................................................................................. 65 4 SUBJECTS AND STUDY DESIGN ........................................................................... 66 4.1 STUDY I ............................................................................................................. 66 4.2 STUDY II ........................................................................................................... 66 4.3 STUDY III .......................................................................................................... 67 4.4 STUDY IV .......................................................................................................... 68 4.5 ETHICAL APPROVAL ....................................................................................... 70 5 5 METHODS ................................................................................................................ 71 5.1 CLINICAL EVALUATION AND EXPERIMENTS .............................................. 71 5.1.1 MEDICAL EXAMINATION ........................................................................ 71 5.1.2 PRE-EXERCISE EXPERIMENTS .............................................................. 71 5.1.2.1 Anthropometry ................................................................................. 71 5.1.2.2 Flow-volume spirometry ................................................................. 71 5.1.2.3 Leisure-time physical activity (LTPA)............................................. 71 5.1.3 BLOOD SAMPLES ..................................................................................... 72 5.1.4 BLOOD OXYGEN CARRYING CAPACITY ............................................... 72 5.1.5 CARDIOPULMONARY EXERCISE TEST (CPET) ................................... 73 5.1.5.1 CPET protocols ................................................................................ 73 5.1.5.2 Systemic oxygen delivery ................................................................ 74 Pulmonary function ....................................................................... 74 Cardiac pump function.................................................................. 74 Scaling of cardiopulmonary data ................................................. 75 5.1.5.3 Peripheral oxygen delivery, extraction, and utilization ................. 75 NIRS experiments .......................................................................... 75 Systemic arterial-venous oxygen difference .................................77 5.2 EXERCISE TRAINING INTERVENTION (STUDY IV) ....................................77 5.3 STATISTICAL ANALYSES ................................................................................ 78 5.3.1 STUDY I ..................................................................................................... 78 5.3.2 STUDY II ................................................................................................... 78 5.3.3 STUDY III .................................................................................................. 79 5.3.4 STUDY IV .................................................................................................. 79 5.3.5 STATISTICAL POWER CALCULATION (STUDIES II-IV)...................... 80 6 RESULTS .................................................................................................................. 81 6.1 DESCRIPTIVE CHARACTERISTICS AND BLOOD OXYGEN CARRYING CAPACITY .......................................................................................................... 81 6.1.1 STUDY I ...................................................................................................... 81 6.1.2 STUDY II .................................................................................................... 81 6.1.3 STUDY III .................................................................................................. 83 6.1.4 STUDY IV ................................................................................................... 83 6.2 OXYGEN DELIVERY AND UTILIZATION DURING ACUTE DYNAMIC EXERCISE ......................................................................................................... 85 6.2.1 RESPONSES TO CPET IN HEALTHY MEN (STUDY I) .......................... 85 6 6.2.2 RESPONSES TO CPET IN WOMEN WITH VERSUS WITHOUT PCOS (STUDY II) ................................................................................................. 89 6.2.3 RESPONSES TO CPET IN MEN WITH VERSUS WITHOUT TYPE 1 DIABETES (STUDY III) ............................................................................. 90 6.2.4 EFFECTS OF EXERCISE TRAINING ON RESPONSES TO CPET IN MEN WITH VERSUS WITHOUT TYPE 1 DIABETES (STUDY IV) ......... 94 6.3 STATISTICAL POWER ..................................................................................... 97 7 DISCUSSION ............................................................................................................ 99 7.1 LOCAL TISSUE-SPECIFIC IMBALANCE BETWEEN OXYGEN DELIVERY AND UTILIZATION AS PART OF WHOLE-BODY RESPONSE TO ACUTE DYNAMIC EXERCISE (STUDY I) ..................................................................... 99 7.2 ALTERATIONS IN PERIPHERAL RATHER THAN SYSTEMIC ADJUSTMENTS TO ACUTE DYNAMIC EXERCISE IN PCOS (STUDY II) .. 102 7.3 BOTH SYSTEMIC AND PERIPHERAL CARDIOVASCULAR IMPAIRMENTS ARE MANIFESTED DURING ACUTE DYNAMIC EXERCISE IN TYPE 1 DIABETES (STUDY III) .................................................................................. 107 7.4 EFFECTS OF LONG-TERM EXERCISE TRAINING ON RESPONSES TO ACUTE DYNAMIC EXERCISE AND GLYCEMIC CONTROL IN TYPE 1 DIABETES (STUDY IV) .................................................................................... 111 7.5 METHODOLOGICAL STRENGTHS AND LIMITATIONS ............................. 114 8 SUMMARY AND CONCLUSIONS .......................................................................... 116 9 FUTURE PERSPECTIVES ...................................................................................... 118 ACKNOWLEDGMENTS .............................................................................................. 119 APPENDICES .............................................................................................................. 121 REFERENCES ............................................................................................................ 122 ORIGINAL PUBLICATIONS ....................................................................................... 156 7 LIST OF ORIGINAL PUBLICATIONS This thesis is based on the following original publications: I Rissanen AP, Tikkanen HO, Koponen AS, Aho JM, Hägglund H, Lindholm H, Peltonen JE. Alveolar gas exchange and tissue oxygenation during incremental treadmill exercise, and their associations with blood O2 carrying capacity. Front Physiol. 2012;3:265. II Rissanen AP, Koskela-Koivisto T, Hägglund H, Koponen AS, Aho JM, Pöyhönen-Alho M, Tiitinen A, Tikkanen HO, Peltonen JE. Altered cardiorespiratory response to exercise in overweight and obese women with polycystic ovary syndrome. Physiol Rep. 2016;4(4):e12719. III Rissanen AP, Tikkanen HO, Koponen AS, Aho JM, Peltonen JE. Central and peripheral cardiovascular impairments limit V̇ O2peak in type 1 diabetes. Med Sci Sports Exerc. 2015;47(2):223-230. IV Rissanen AP, Tikkanen HO, Koponen AS, Aho JM, Peltonen JE. One-year individualized exercise training intervention enhances cardiorespiratory fitness but not muscle deoxygenation or glycemic control in adults with type 1 diabetes. Submitted. The publications are referred to in the text by their Roman numerals and have been reprinted with permission of their copyright holders. In addition to the original publications, some unpublished data are presented. 8 ABBREVIATIONS ANS-EXE Autonomic Nervous System and EXErcise in gestational diabetes ARTEMIS Innovation to Reduce Cardiovascular Complications of Diabetes at the Intersection of Discovery, Prevention and Knowledge Exchange AT anaerobic threshold ATP adenosine triphosphate (a-v)O2 arterial-venous oxygen difference BMI body mass index BV blood volume CaO2 arterial oxygen content C(a-v)O2 systemic arterial-venous oxygen difference CO carbon monoxide CO2 carbon dioxide CPET cardiopulmonary exercise test CPO cardiac power output CPOi cardiac power output index CTI contractility index (= dZ/dtmax) CvO2 mixed venous oxygen content CVP central venous pressure DPF differential pathlength factor dZ/dt rate of variation of thoracic impedance dZ/dtmax largest rate of variation of thoracic impedance during systole (= CTI) EDGE Exercise, Diet and GEnes in T1D EDV end-diastolic volume EDVi end-diastolic volume index EF ejection fraction EIAH exercise-induced arterial hypoxemia ES standardized effect size FEV1 forced expiratory volume in one second FFM fat-free mass FinnDiane Finnish Diabetic Nephropathy Study FVC forced vital capacity Hb hemoglobin [Hb] hemoglobin concentration HbA1c glycosylated hemoglobin A1c [HbCO] carboxyhemoglobin concentration HHb deoxygenated hemoglobin ∆[HHb] relative concentration change in deoxygenated hemoglobin %∆[HHb] normalized relative concentration change in deoxygenated hemoglobin HOMA-IR homeostasis model assessment of insulin resistance HR heart rate IR insulin resistance 9 LTPA leisure-time physical activity MAP mean arterial blood pressure MET metabolic equivalent (= resting pulmonary oxygen uptake) NIP near-infrared spectroscopy inflection point NIPTissueAT/RC NIP observed in (leg muscle, arm muscle, or cerebral) tissue and being closest to anaerobic threshold or respiratory compensation point NIRS near-infrared spectroscopy NO nitric oxide O2 oxygen O2Hb oxygenated hemoglobin ∆[O2Hb] relative concentration change in oxygenated hemoglobin O2 pulse pulmonary oxygen uptake / heart rate O2 pulsei oxygen pulse index PaCO2 partial pressure of arterial carbon dioxide PaO2 partial pressure of arterial oxygen PCOS polycystic ovary syndrome PETCO2 end-tidal pressure of carbon dioxide of respired gases PvO2 partial pressure of muscle venous oxygen Q̇ cardiac output Q̇ i cardiac output index Q̇ VL local microvascular blood flow in the vastus lateralis muscle RC respiratory compensation point RER respiratory exchange ratio SD standard deviation SHBG sex hormone-binding globulin SpO2 arterial oxygen saturation SV stroke volume SVcal stroke volume during a PhysioFlow calibration procedure SVi stroke volume index SVR systemic vascular resistance SVRi systemic vascular resistance index tHb total hemoglobin ∆[tHb] relative concentration change in total hemoglobin tHb-mass total hemoglobin mass TSI tissue saturation index V̇ A alveolar ventilation V̇ CO2 pulmonary carbon dioxide output V̇ E minute ventilation V̇ O2 pulmonary oxygen uptake V̇ O2max maximal pulmonary oxygen uptake V̇ O2peak peak pulmonary oxygen uptake WEBWOMEX WEll-Being of WOMen during pregnancy: focus on individualized EXercise training Zmax highest thoracic impedance during systole Zmin lowest thoracic impedance during systole 10

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Department of Sports and Exercise Medicine Clinicum Faculty of Medicine University of Helsinki Helsinki, Finland OXYGEN DELIVERY AND UTILIZATION DURING ACUTE DYNAMIC EXERCISE EFFECTS OF POLYCYSTIC OVARY SYNDROME, TYPE 1 DIABETES, AND EXERCISE TRAINING Antti-Pekka Rissanen ACADEMIC DISSERTATION To be presented, with the permission of the Faculty of Medicine of the University of Helsinki, for public examination in Auditorium PIII, Porthania, City Centre Campus, University of Helsinki, on May 20th 2017, at 10 o’clock in the morning. Helsinki 2017 Supervised by Docent Juha Peltonen, Ph.D. Department of Sports and Exercise Medicine Clinicum Faculty of Medicine University of Helsinki Helsinki, Finland Clinic for Sports and Exercise Medicine Foundation for Sports and Exercise Medicine Helsinki, Finland Professor Heikki Tikkanen, M.D., Ph.D. Institute of Biomedicine School of Medicine University of Eastern Finland Kuopio, Finland Department of Sports and Exercise Medicine Clinicum Faculty of Medicine University of Helsinki Helsinki, Finland Clinic for Sports and Exercise Medicine Foundation for Sports and Exercise Medicine Helsinki, Finland Reviewed by Senior Research Fellow J. Chris Baldi, Ph.D. Department of Medicine Dunedin School of Medicine University of Otago Dunedin, New Zealand Docent David E. Laaksonen, M.D., Ph.D., M.P.H. Internal Medicine Institute of Clinical Medicine Kuopio University Hospital Kuopio, Finland Opponent Professor Robert Petrella, M.D., Ph.D. Department of Family Medicine and Kinesiology Schulich School of Medicine and Dentistry Western University London, Canada Dissertationes Scholae Doctoralis Ad Sanitatem Investigandam Universitatis Helsinkiensis ISBN 978-951-51-3177-5 (paperback) ISSN 2342-3161 (print) ISBN 978-951-51-3178-2 (PDF) ISSN 2342-317X (online) Cover artwork: Liia Rissanen Press: Unigrafia Oy (Yliopistopaino Helsinki), Helsinki, Finland, 2017 http://ethesis.helsinki.fi Man, he took his time in the sun Had a dream to understand A single grain of sand - from the Nightwish masterpiece The Greatest Show on Earth (lyrics by Tuomas Holopainen) CONTENTS CONTENTS ..................................................................................................................... 4 LIST OF ORIGINAL PUBLICATIONS ........................................................................... 8 ABBREVIATIONS .......................................................................................................... 9 ABSTRACT ..................................................................................................................... 11 TIIVISTELMÄ (ABSTRACT IN FINNISH) ................................................................... 13 1 INTRODUCTION ......................................................................................................16 2 REVIEW OF THE LITERATURE ............................................................................ 18 2.1 OXYGEN DELIVERY AND UTILIZATION DURING ACUTE DYNAMIC EXERCISE ......................................................................................................... 18 2.1.1 SYSTEMIC OXYGEN DELIVERY DURING ACUTE DYNAMIC EXERCISE .................................................................................................. 20 2.1.1.1 Pulmonary function ......................................................................... 20 2.1.1.2 Cardiac pump function .................................................................... 21 Cardiac cycle ................................................................................... 21 Cardiac pump function during acute dynamic exercise .............. 22 Methods to quantify cardiac pump function ................................ 25 2.1.1.3 Blood oxygen carrying capacity ...................................................... 28 2.1.2 PERIPHERAL OXYGEN DELIVERY, EXTRACTION, AND UTILIZATION DURING ACUTE DYNAMIC EXERCISE .................................................. 29 2.1.2.1 Overview of blood flow distribution and regulation at the whole- body level ........................................................................................ 29 2.1.2.2 Skeletal muscle blood flow and oxygen extraction and utilization 31 Skeletal muscle blood flow ............................................................. 31 Skeletal muscle oxygen extraction and utilization ....................... 34 2.1.2.3 Cerebral blood flow and oxygen extraction and utilization ........... 35 2.1.2.4 Near-infrared spectroscopy (NIRS) as a method to evaluate tissue- specific (im)balance between oxygen delivery and utilization ...... 36 2.1.3 PEAK OXYGEN UPTAKE (V̇ O2peak) .......................................................... 40 2.1.3.1 V̇ O2peak as a concept ........................................................................ 40 2.1.3.2 V̇ O2peak as an integrated model ....................................................... 41 2.1.3.3 V̇ O2peak as a predictor of cardiovascular disease and mortality ..... 44 4 2.1.4 EFFECTS OF ENDURANCE TRAINING ON OXYGEN DELIVERY AND UTILIZATION DURING ACUTE DYNAMIC EXERCISE ......................... 46 2.1.4.1 Effects of endurance training on systemic oxygen delivery ........... 46 2.1.4.2 Effects of endurance training on skeletal muscle blood flow and oxygen extraction and utilization .................................................. 47 2.1.4.3 Endurance training effects in the context of Fick’s theories ......... 49 2.2 POLYCYSTIC OVARY SYNDROME (PCOS) .................................................... 50 2.2.1 DEFINITION AND EPIDEMIOLOGY ...................................................... 50 2.2.2 PATHOGENESIS ...................................................................................... 50 2.2.3 CLINICAL MANIFESTATIONS ................................................................. 51 2.2.4 PCOS AND EXERCISE ............................................................................. 52 2.2.4.1 Oxygen delivery and utilization during acute dynamic exercise in PCOS............................................................................................... 52 2.2.4.2 Effects of exercise training in PCOS .............................................. 53 2.3 TYPE 1 DIABETES ............................................................................................ 53 2.3.1 DEFINITION AND EPIDEMIOLOGY ...................................................... 53 2.3.2 PATHOGENESIS ...................................................................................... 54 2.3.3 CLINICAL MANIFESTATIONS ................................................................ 55 2.3.3.1 Microvascular complications ......................................................... 56 2.3.3.2 Macrovascular complications ........................................................ 56 2.3.3.3 Diabetic cardiomyopathy ............................................................... 57 2.3.4 TYPE 1 DIABETES AND EXERCISE ........................................................ 58 2.3.4.1 Oxygen delivery and utilization during acute dynamic exercise in type 1 diabetes ................................................................................ 58 2.3.4.2 Effects of exercise training in type 1 diabetes .................................61 3 AIMS OF THE STUDY ............................................................................................. 65 4 SUBJECTS AND STUDY DESIGN ........................................................................... 66 4.1 STUDY I ............................................................................................................. 66 4.2 STUDY II ........................................................................................................... 66 4.3 STUDY III .......................................................................................................... 67 4.4 STUDY IV .......................................................................................................... 68 4.5 ETHICAL APPROVAL ....................................................................................... 70 5 5 METHODS ................................................................................................................ 71 5.1 CLINICAL EVALUATION AND EXPERIMENTS .............................................. 71 5.1.1 MEDICAL EXAMINATION ........................................................................ 71 5.1.2 PRE-EXERCISE EXPERIMENTS .............................................................. 71 5.1.2.1 Anthropometry ................................................................................. 71 5.1.2.2 Flow-volume spirometry ................................................................. 71 5.1.2.3 Leisure-time physical activity (LTPA)............................................. 71 5.1.3 BLOOD SAMPLES ..................................................................................... 72 5.1.4 BLOOD OXYGEN CARRYING CAPACITY ............................................... 72 5.1.5 CARDIOPULMONARY EXERCISE TEST (CPET) ................................... 73 5.1.5.1 CPET protocols ................................................................................ 73 5.1.5.2 Systemic oxygen delivery ................................................................ 74 Pulmonary function ....................................................................... 74 Cardiac pump function.................................................................. 74 Scaling of cardiopulmonary data ................................................. 75 5.1.5.3 Peripheral oxygen delivery, extraction, and utilization ................. 75 NIRS experiments .......................................................................... 75 Systemic arterial-venous oxygen difference .................................77 5.2 EXERCISE TRAINING INTERVENTION (STUDY IV) ....................................77 5.3 STATISTICAL ANALYSES ................................................................................ 78 5.3.1 STUDY I ..................................................................................................... 78 5.3.2 STUDY II ................................................................................................... 78 5.3.3 STUDY III .................................................................................................. 79 5.3.4 STUDY IV .................................................................................................. 79 5.3.5 STATISTICAL POWER CALCULATION (STUDIES II-IV)...................... 80 6 RESULTS .................................................................................................................. 81 6.1 DESCRIPTIVE CHARACTERISTICS AND BLOOD OXYGEN CARRYING CAPACITY .......................................................................................................... 81 6.1.1 STUDY I ...................................................................................................... 81 6.1.2 STUDY II .................................................................................................... 81 6.1.3 STUDY III .................................................................................................. 83 6.1.4 STUDY IV ................................................................................................... 83 6.2 OXYGEN DELIVERY AND UTILIZATION DURING ACUTE DYNAMIC EXERCISE ......................................................................................................... 85 6.2.1 RESPONSES TO CPET IN HEALTHY MEN (STUDY I) .......................... 85 6 6.2.2 RESPONSES TO CPET IN WOMEN WITH VERSUS WITHOUT PCOS (STUDY II) ................................................................................................. 89 6.2.3 RESPONSES TO CPET IN MEN WITH VERSUS WITHOUT TYPE 1 DIABETES (STUDY III) ............................................................................. 90 6.2.4 EFFECTS OF EXERCISE TRAINING ON RESPONSES TO CPET IN MEN WITH VERSUS WITHOUT TYPE 1 DIABETES (STUDY IV) ......... 94 6.3 STATISTICAL POWER ..................................................................................... 97 7 DISCUSSION ............................................................................................................ 99 7.1 LOCAL TISSUE-SPECIFIC IMBALANCE BETWEEN OXYGEN DELIVERY AND UTILIZATION AS PART OF WHOLE-BODY RESPONSE TO ACUTE DYNAMIC EXERCISE (STUDY I) ..................................................................... 99 7.2 ALTERATIONS IN PERIPHERAL RATHER THAN SYSTEMIC ADJUSTMENTS TO ACUTE DYNAMIC EXERCISE IN PCOS (STUDY II) .. 102 7.3 BOTH SYSTEMIC AND PERIPHERAL CARDIOVASCULAR IMPAIRMENTS ARE MANIFESTED DURING ACUTE DYNAMIC EXERCISE IN TYPE 1 DIABETES (STUDY III) .................................................................................. 107 7.4 EFFECTS OF LONG-TERM EXERCISE TRAINING ON RESPONSES TO ACUTE DYNAMIC EXERCISE AND GLYCEMIC CONTROL IN TYPE 1 DIABETES (STUDY IV) .................................................................................... 111 7.5 METHODOLOGICAL STRENGTHS AND LIMITATIONS ............................. 114 8 SUMMARY AND CONCLUSIONS .......................................................................... 116 9 FUTURE PERSPECTIVES ...................................................................................... 118 ACKNOWLEDGMENTS .............................................................................................. 119 APPENDICES .............................................................................................................. 121 REFERENCES ............................................................................................................ 122 ORIGINAL PUBLICATIONS ....................................................................................... 156 7 LIST OF ORIGINAL PUBLICATIONS This thesis is based on the following original publications: I Rissanen AP, Tikkanen HO, Koponen AS, Aho JM, Hägglund H, Lindholm H, Peltonen JE. Alveolar gas exchange and tissue oxygenation during incremental treadmill exercise, and their associations with blood O2 carrying capacity. Front Physiol. 2012;3:265. II Rissanen AP, Koskela-Koivisto T, Hägglund H, Koponen AS, Aho JM, Pöyhönen-Alho M, Tiitinen A, Tikkanen HO, Peltonen JE. Altered cardiorespiratory response to exercise in overweight and obese women with polycystic ovary syndrome. Physiol Rep. 2016;4(4):e12719. III Rissanen AP, Tikkanen HO, Koponen AS, Aho JM, Peltonen JE. Central and peripheral cardiovascular impairments limit V̇ O2peak in type 1 diabetes. Med Sci Sports Exerc. 2015;47(2):223-230. IV Rissanen AP, Tikkanen HO, Koponen AS, Aho JM, Peltonen JE. One-year individualized exercise training intervention enhances cardiorespiratory fitness but not muscle deoxygenation or glycemic control in adults with type 1 diabetes. Submitted. The publications are referred to in the text by their Roman numerals and have been reprinted with permission of their copyright holders. In addition to the original publications, some unpublished data are presented. 8 ABBREVIATIONS ANS-EXE Autonomic Nervous System and EXErcise in gestational diabetes ARTEMIS Innovation to Reduce Cardiovascular Complications of Diabetes at the Intersection of Discovery, Prevention and Knowledge Exchange AT anaerobic threshold ATP adenosine triphosphate (a-v)O2 arterial-venous oxygen difference BMI body mass index BV blood volume CaO2 arterial oxygen content C(a-v)O2 systemic arterial-venous oxygen difference CO carbon monoxide CO2 carbon dioxide CPET cardiopulmonary exercise test CPO cardiac power output CPOi cardiac power output index CTI contractility index (= dZ/dtmax) CvO2 mixed venous oxygen content CVP central venous pressure DPF differential pathlength factor dZ/dt rate of variation of thoracic impedance dZ/dtmax largest rate of variation of thoracic impedance during systole (= CTI) EDGE Exercise, Diet and GEnes in T1D EDV end-diastolic volume EDVi end-diastolic volume index EF ejection fraction EIAH exercise-induced arterial hypoxemia ES standardized effect size FEV1 forced expiratory volume in one second FFM fat-free mass FinnDiane Finnish Diabetic Nephropathy Study FVC forced vital capacity Hb hemoglobin [Hb] hemoglobin concentration HbA1c glycosylated hemoglobin A1c [HbCO] carboxyhemoglobin concentration HHb deoxygenated hemoglobin ∆[HHb] relative concentration change in deoxygenated hemoglobin %∆[HHb] normalized relative concentration change in deoxygenated hemoglobin HOMA-IR homeostasis model assessment of insulin resistance HR heart rate IR insulin resistance 9 LTPA leisure-time physical activity MAP mean arterial blood pressure MET metabolic equivalent (= resting pulmonary oxygen uptake) NIP near-infrared spectroscopy inflection point NIPTissueAT/RC NIP observed in (leg muscle, arm muscle, or cerebral) tissue and being closest to anaerobic threshold or respiratory compensation point NIRS near-infrared spectroscopy NO nitric oxide O2 oxygen O2Hb oxygenated hemoglobin ∆[O2Hb] relative concentration change in oxygenated hemoglobin O2 pulse pulmonary oxygen uptake / heart rate O2 pulsei oxygen pulse index PaCO2 partial pressure of arterial carbon dioxide PaO2 partial pressure of arterial oxygen PCOS polycystic ovary syndrome PETCO2 end-tidal pressure of carbon dioxide of respired gases PvO2 partial pressure of muscle venous oxygen Q̇ cardiac output Q̇ i cardiac output index Q̇ VL local microvascular blood flow in the vastus lateralis muscle RC respiratory compensation point RER respiratory exchange ratio SD standard deviation SHBG sex hormone-binding globulin SpO2 arterial oxygen saturation SV stroke volume SVcal stroke volume during a PhysioFlow calibration procedure SVi stroke volume index SVR systemic vascular resistance SVRi systemic vascular resistance index tHb total hemoglobin ∆[tHb] relative concentration change in total hemoglobin tHb-mass total hemoglobin mass TSI tissue saturation index V̇ A alveolar ventilation V̇ CO2 pulmonary carbon dioxide output V̇ E minute ventilation V̇ O2 pulmonary oxygen uptake V̇ O2max maximal pulmonary oxygen uptake V̇ O2peak peak pulmonary oxygen uptake WEBWOMEX WEll-Being of WOMen during pregnancy: focus on individualized EXercise training Zmax highest thoracic impedance during systole Zmin lowest thoracic impedance during systole 10

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