The Cardiovascular System

Description

The Cardiovascular System: Phenotypic and Physiological Responses, Volume 37, part of a two-volume set, provides comprehensive coverage of the current state of knowledge in this very active and growing field of research, also highlighting the tremendous diversity in cardiovascular morphology and function among the various fish taxa and the anatomical and physiological plasticity shown by this system when faced with various abiotic and biotic challenges. Specific chapters in this updated book include Research Technologies/Methodology for Studying Fish Cardiovascular Function, Cardiovascular Development in Embryonic and Larval Fishes, Cardiovascular Responses to Limiting Oxygen Levels, and Temperature and the Cardiovascular System.

The book's chapters integrate molecular and cellular data with the growing body of knowledge on heart and in vivo cardiovascular function, and as a result, provide insights into some of the most interesting, and important, questions that still need to be answered in this field.

Presents a comprehensive overview of cardiovascular structure and function in fish
Provides a valuable resource for researchers in fish physiology and audiences within the fields of comparative morphology and histology, aquaculture and ecophysiology
Highlights the tremendous diversity in cardiovascular morphology and function among the various fish taxa

Front Cover 2
The Cardiovascular System: Development, Plasticity and Physiological Responses: Development, Plasticity and Physiological R... 5
Copyright 6
Contents 7
Contributors 9
Abbreviations 11
Preface 23
Chapter 1: The O2 and CO2 Transport System in Teleosts and the Specialized Mechanisms That Enhance Hb-O2 Unloading to Tissues 27
- 1. Introduction 28
- 2. Blood O2 Transport 29
  - 2.1. Hematocrit 31
  - 2.2. Hemoglobin O2-Binding 32
    - 2.2.1. The Oxygen Equilibrium Curve 32
    - 2.2.2. The Optimal P50 35

Front Cover 2
The Cardiovascular System: Development, Plasticity and Physiological Responses: Development, Plasticity and Physiological R... 5
Copyright 6
Contents 7
Contributors 9
Abbreviations 11
Preface 23
Chapter 1: The O2 and CO2 Transport System in Teleosts and the Specialized Mechanisms That Enhance Hb-O2 Unloading to Tissues 27
- 1. Introduction 28
- 2. Blood O2 Transport 29
  - 2.1. Hematocrit 31
  - 2.2. Hemoglobin O2-Binding 32
    - 2.2.1. The Oxygen Equilibrium Curve 32
    - 2.2.2. The Optimal P50 35
  - 2.3. Erythrocyte Function 40
    - 2.3.1. Allosteric Modulators of Hemoglobin P50 40
    - 2.3.2. H+ as Allosteric Effectors 41
    - 2.3.3. Physiology of Carbonic Anhydrase 43
    - 2.3.4. The Jacobs-Stewart Cycle 44
    - 2.3.5. Steady-State Erythrocyte pH 44
    - 2.3.6. Active Regulation of Erythrocyte pH 46
  - 3. Regulation of Blood O2 Capacitance In Vivo 48
    - 3.1. Hypoxia 49
      - 3.1.1. Capacity Response 49
      - 3.1.2. Affinity Response 52
    - 3.2. Exercise 54
      - 3.2.1. Capacity Response 55
      - 3.2.2. Affinity Response 56
    - 3.3. Temperature 58
      - 3.3.1. Capacity Response 59
      - 3.3.2. Affinity Response 61
    - 4. Interaction Between O2 and CO2 Transport 63
      - 4.1. The Potential to Enhance Blood O2 Capacitance via the Bohr-Effect 63
      - 4.2. The Optimal Bohr-Coefficient 68
    - 5. A Novel Mechanism of Enhanced Hb-O2 Unloading in Teleosts 69
      - 5.1. Short-Circuiting pH Disequilibria 71
        
        5.1.2. Short-Circuiting pH Disequilibria in the Plasma 71
        
        5.1.1. Short-Circuiting pH Disequilibria Across the Erythrocyte Membrane 71
        
        5.1.3. Evidence for RBC Î²-NHE Short-Circuiting 73
      - 5.2. The Distribution of Plasma-Accessible Carbonic Anhydrase in the Circulation of Fishes 77
        
        5.2.1. Plasma-Accessible Carbonic Anhydrase at the Gill 79
        
        5.2.1.1. Biochemical Studies 79
        
        5.2.1.2. Inhibitor Studies 82
        
        5.2.1.3. Imaging and Molecular Studies 83
        
        5.2.2. Plasma-Accessible Carbonic Anhydrase at the Tissue 85
        
        5.2.3. Plasma Carbonic Anhydrase Inhibitors 87
        
        5.2.4. The Evolution of Differential Plasma-Accessible Carbonic Anhydrase Distribution in Fishes 89
      - References 97
    - Chapter 2: Cardiovascular Development in Embryonic and Larval Fishes 133
      - 1. Introduction: Development in a Challenging Environment 134
        
        1.1. Modes of Reproduction and Development in Fishes 135
        
        1.2. Physiological Consequences of Reproduction and Development Mode 136
        
        1.3. Developmental Rate and Timing 138
      - 2. A Framework for Understanding Cardiovascular Development 139
        
        2.1. The Changing Roles of the Embryonic and Larval Heart and Circulation 139
        
        2.2. Cutaneous Materials Exchange 140
        
        2.3. The Cutaneous-Branchial Transition and Role of Circulatory Bulk Flow 141
        
        2.4. Early Functions of the Beating Heart 141
      - 3. Cardiac Development 142
        
        3.1. Formation and Development of the Heart 143
        
        3.1.1. Heart Tube Formation 143
        
        3.1.2. Cardiac Chamber Formation 144
        
        3.1.3. Cardiac Valve Formation 147
        
        3.1.4. Myocardial Differentiation 151
        
        3.1.5. Development of the Cardiac Conduction System 152
        
        3.2. Regulation of Heart Formation 153
        
        3.2.1. Mechanical Factors 154
        
        3.2.2. Molecular and Genetic Factors 156
        
        4. Vascular Development 159
        
        4.1. Vascular Structure and Pattern 159
        
        4.1.2. The Larva 159
        
        4.1.1. The Embryo 159
        
        4.2. Vasculogenesis/Angiogenesis 161
        
        4.2.1. Formation of the Circulatory Network 161
        
        4.2.2. Vascular Remodeling During Development 165
        
        4.3. The Secondary Circulation 165
        
        4.4. Lymphatic System 166
        
        5. Ontogeny of Cardiovascular Regulation 166
        
        5.1. Introduction 166
        
        5.2. Ontogeny of Heart Function 167
        
        5.2.1. Conduction System of the Heart 167
        
        5.2.2. Heart Rate 170
        
        5.2.2.1. Control of Heart Rate 170
        
        5.2.2.2. Emerging Techniques for Electrocardiology in Fish Larvae 172
        
        5.2.3. Blood Pressure 174
        
        5.2.4. Stroke Volume and Cardiac Output 176
        
        5.2.5. Ontogeny of Vascular Regulation 178
        
        5.3. Cardiac Regulation of Whole Body Metabolism 180
        
        5.4. Environment and Cardiac Function 182
        
        5.4.1. Temperature 183
        
        5.4.2. Water Chemistry 185
        
        5.4.2.1. Hypoxia 185
        
        5.4.2.2. Toxicants 186
        
        6. Conclusion and Suggested Directions for Future Research 189
        
        6.1. Key Importance of the Cardiovascular System 189
        
        6.2. Assessment of Fitness 190
        
        6.3. Vulnerability to Environment 190
        
        6.5. Need for "Vertical Integration" From Molecules to Physiology, Behavior, and Ecology 191
        
        6.4. Technological Advances 191
        
        6.6. Epigenetic Influences 192
        
        References 192
        
        Chapter 3: Cardiac Preconditioning, Remodeling and Regeneration 211
        
        1. Introduction 212
        
        2. Cardiac Response to Hypoxia, Anoxia and Anemia 213
        
        2.1. Hypoxia Preconditioning 213
        
        2.2. Cardiac Remodeling in Response to Hypoxia 216
        
        2.3. Cardiac Remodeling in Response to Anemia 220
        
        3. Response of the Heart to Acute and Chronic Changes in Temperature 221
        
        3.1. The Initial Response to a Temperature Challenge 221
        
        3.2. Response of the Heart to a Chronic Temperature Stressor 222
        
        3.3. Remodeling of Connective Tissue With Thermal Acclimation 227
        
        3.4. Initiation of Cardiac Remodeling With Thermal Acclimation 229
        
        4. Endocrine Regulation of Cardiac Remodeling in Fish 232
        
        4.1. Role of Angiotensin II in Cardiac Remodeling 232
        
        4.2. Role of 11-Ketotestosterone in Cardiac Remodeling 233
        
        5. The Zebrafish Heart as a Model for Cardiac Regeneration 234
        
        5.1. Summary of the Regenerative Process 234
        
        5.2. Role of the Myocardium in Regeneration 236
        
        5.3. Role of the Epicardium in Regeneration 241
        
        5.4. Role of the Endocardium in Regeneration 242
        
        5.5. Cardiac Fibrosis Pathways 243
        
        5.6. Role of microRNAs in Cardiac Regeneration in Zebrafish 244
        
        5.7. Heart Regeneration Across Species 246
        
        5.8. Heart vs Limb Regeneration 246
        
        6. Perspectives 248
        
        References 249
        
        Chapter 4: Temperature and the Cardiovascular System 261
        
        1. Temperature Effects on Cardiovascular Function 262
        
        1.1. Morphology 262
        
        1.2. Cardiac Output 268
        
        1.2.1. Acute Responses 268
        
        1.2.2. Acclimation Responses 270
        
        1.3. Heart Rate 273
        
        1.3.1. Acute Responses 273
        
        1.3.2. Acclimation Responses 284
        
        1.4. Stroke Volume 286
        
        1.4.1. Acute Responses 286
        
        1.4.2. Acclimation Responses 288
        
        1.5. In Vitro and In Situ Cardiac Performance 289
        
        1.6. Cardiac β-Adrenoceptors 290
        
        1.7. Energy Metabolism and Mitochondrial Capacity 295
        
        1.8. Regional Blood Flow 297
        
        1.9. Blood Parameters 300
        
        1.9.1. Hematological Variables 300
        
        1.9.2. Arterial Blood Oxygenation 301
        
        1.9.3. Venous Blood Oxygenation 303
        
        2. Ecological Implications 304
        
        2.1. Role of the Cardiovascular System in Supporting Whole Animal Performance 304
        
        2.2. Cardiac Collapse at High Temperatures 306
        
        2.3. Ecological, Evolutionary, and Conservation Implications 308
        
        References 310
        
        Chapter 5: Cardiovascular Responses to Limiting Oxygen Levels 325
        
        1. Introduction 326
        
        1.1. Oxygen: The Molecule of Life 326
        
        1.2. Hypoxia and Anoxia in the Aquatic Environment 328
        
        1.3. Focus of the Chapter 330
        
        2. In Vivo Cardiovascular Responses to Oxygen Limitation in Severe Hypoxia/Anoxia-Intolerant Species 332
        
        2.1. Brief Overview of Fish Cardiovascular Physiology 332
        
        2.2. Water-Breathers 336
        
        2.3. Air-Breathers 349
        
        3. Strategies of Survival, Cardiovascular Responses and Cardiovascular Control in Severe Hypoxia/Anoxia-Tolerant Species ... 355
        
        3.1. Overview of Strategies to Balance ATP Supply and Demand During Oxygen Deprivation 355
        
        3.2. Cardiovascular Responses of Severe Hypoxia/Anoxia-Tolerant Species to Oxygen Deprivation 358
        
        3.3. Cardiovascular Control in Severe Hypoxia/Anoxia-Tolerant Species During Prolonged Oxygen Deprivation 365
        
        4. Additional Insights Into the Mechanisms of Anoxic Cardiac Survival in Crucian Carp 371
        
        4.1. Cold Temperature Prepares the Heart for Anoxia Exposure 371
        
        4.2. Absence of "Channel Arrest" in the Anoxic Crucian Carp Heart 373
        
        4.3. Management of Metabolic Wastes 374
        
        5. Concluding Remarks 377
        
        References 379
        
        Chapter 6: Environmental Pollution and the Fish Heart 399
        
        1. Introduction 400
        
        1.1. The Significance of the Fish Heart as a Target Organ for Widespread Environmental Pollution 400
        
        1.2. Major Sources of Toxic Chemicals in Fish Habitats: Past, Present and Future 401
        
        1.3. Routes of Chemical Uptake and Transport to the Fish Cardiovascular System 406
        
        2. Developmental Cardiotoxicity 408
        
        2.1. Factors Contributing to the Particular Vulnerability of Fish Early Life Stages to Cardiotoxic Agents 408
        
        2.2. Brief Summary of Heart Development in Embryos and Subsequent Growth in Larvae and Juveniles (Also See Chapter 2, Vol ... 408
        
        2.3. Brief Summary Relating Cardiac Anatomy to Swimming Performance (Also See Chapter 4, Volume 36A: Farrell and Smith, 2017) 411
        
        2.4. Case Example: Dioxins and PCBs 412
        
        2.4.1. The Collapse of Lake Trout Populations in the Great Lakes 412
        
        2.4.2. Cellular Pathways and Mechanisms of Organochlorine Cardiotoxicity 414
        
        2.4.3. Legacy Pollution and the Evolution of Chemical Resistance 416
        
        2.5. Case Example: Oil Spill-Derived PAHs 418
        
        2.5.1. Broad Insights Gained From the 1989 Exxon Valdez Disaster 418
        
        2.5.2. Linking Field and Laboratory Findings 419
        
        2.5.3. Sorting Out the Causes of a Complex Heart Failure Syndrome 422
        
        2.5.4. Mechanisms of Acute Toxicity to the Embryonic Fish Heart From Crude Oil Exposure 427
        
        2.5.4.1. Roles of the AHR 428
        
        2.5.4.2. AHR-Independent Effects on Intracellular Ion Handling and Cardiomyocyte Physiology 429
        
        2.5.4.3. Potential for Interactions and Synergistic Cardiotoxicity 432
        
        2.5.5. Long-Term Impacts to Cardiac Anatomy and Function in Fish That Survive Embryonic Oil Exposures 433
        
        2.5.6. Adverse Impacts on Cardiac Function and Development Through Effects on Gene Expression 437
        
        2.5.6.1. Effects on Genes Involved in Action Potential Generation and E-C Coupling 437
        
        2.5.6.2. Effects on Genes Related to Terminal Differentiation of Cardiomyocytes and Cardiac Form 439
        
        3. Focal Areas for Ongoing and Future Research 442
        
        3.1. An Improved Mechanistic Understanding of Environmental Cardiotoxicity 442
        
        3.2. Next-Generation Molecular Indicators to Assess Fish Health in Polluted Habitats 443
        
        3.3. Biological Scaling: Application to Fisheries Management 444
        
        3.4. Fish Models for Genetics: Crossover Applications for Human and Fish Cardiovascular Health 445
        
        References 447
        
        Chapter 7: Cardiovascular Effects of Disease: Parasites and Pathogens 461
        
        1. Introduction 462
        
        2. Non-infectious Diseases and Disorders 463
        
        2.1. Morphological Anomalies 463
        
        2.2. Coronary Arteriosclerosis 464
        
        2.3. Nutritional Effects 465
        
        2.4. Anemia 467
        
        2.4.1. Types of Anemia 467
        
        2.4.2. Functional Responses to Anemia 468
        
        3. Infectious Diseases and Disorders 470
        
        3.1. Immune Cell Populations in the Heart and Immune Responses 470
        
        3.2. Viral Diseases and Cardiac Inflammation 472
        
        3.2.1. Infectious Salmon Anemia 472
        
        3.2.2. Cardiomyopathy Syndrome 473
        
        3.2.3. Heart and Skeletal Muscle Inflammation 474
        
        3.2.4. Pancreas Disease 477
        
        3.3. Parasitic Infection and Cardiovascular Performance 479
        
        3.3.1. Cardiac Microsporidiosis in Atlantic Cod 479
        
        3.3.2. Ichthyophoniasis 480
        
        3.3.3. Amoebic Gill Disease 482
        
        3.3.4. Lernaeocera branchialis 483
        
        4. Concluding Remarks 485
        
        References 486
        
        Index 497
        
        Other volumes in the fish physiology series 511
        
        Back Cover 515

Development, Plasticity and Physiological Responses

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