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Physiology and Form of Fish Circulation (Satchell 1991)

Satchell, Geoffrey H.: Physiology and Form of Fish Circulation, Cambridge University Press, 1991




Preface xv

1 - Introduction 1

Some elementary haemodynamics 2
The Poiseuille equation 3
Viscosity 3
Total fluid energy 4
Resistance 5
Pressures on land and in water 1
Velocity and cross sectional area 9
2 - The heart 10

Introduction 10
The structure of cardiac muscle 11

The dimensions offish myocytes 12
Sarcoplasmic reticulum, caveolae and T tubules 13
Intercalated discs, fasciae adherentes and gap junctions 14
Specific granules 14
Pace-makers and pace-maker tissue 15
The atrioventricular and ventriculoconal delays 17
Fast conducting tracts 18

Electrical properties of fish cardiac muscle 19

The resting membrane potential and muscle action potential 19
The electrocardiogram (ECG) 20

The events of the cardiac cycle 21

The atrium 21
The ventricle 22
The pericardium and pericardial fluid 24
The bulbus arteriosus 27
The conus arteriosus 29
The sinus venosus 31

The myocardium and its blood supply 31

The compact layer 31
The coronary arteries; atherosclerosis and cardiac pathology 32
The spongy layer 34

The output of the heart 36

Starling's law of the heart 38
Homeometric regulation 39
3 - The peripheral circulation 41

Introduction 41

Arteries 41
Arterioles 43
Capillaries 44
Capillary beds, cross sectional area and velocity of flow 46
Interchanges across the capillary wall; the Starling principle 47
The two pathways through the capillary wall 48
The effect of plasma proteins on permeability 49
The fate of spilt plasma protein 50
Veins 51
Valves in veins 52

Blood pressure in fish 53
Reactive hyperaemia 53
The principal blood vessels of the fish circulatory system 55
4 - The blood 58

Haemoglobin 59

Blood oxygen capacity 59
The oxygenation of haemoglobin 61
Oxygen affinity, P50 and environmental PO2 62
The Bohr effect 64
The Root effect 65
The effect of cell haemoglobin concentration 66
The effect of temperature 66
The effect of nucleoside triphosphates (NTP) 67

Acclimation and adaptation in haemoglobins 68

Multiple haemoglobins 69

The carriage of CO2 in the blood 70
The plasma proteins 71

Fibrinogen and the blood clotting factors 73

Blood viscosity 75

The relation of η to temperature 75

The relation of η to haematocrit 76
The relation of η to shear rate 76

The leucocytes 77
5 - Haemopoiesis and phagocytosis - the mononuclear phagocytic system 80

The phagocytic cells of the MPS system 80

Macrophages 81

Haemopoietic and phagocytic organs and tissues 82

The thymus 82
The spleen 83
The gut-associated tissue 84
The kidney 85
The organ of Ley dig, the epigonal organ and the meninges 85
The cavernous bodies 86
6 - Circulation through special regions 89

I. The microcirculation of the gill 89

Introduction 89
The arterioarterial pathway 90
Blood pressure and blood flow through the secondary lamellae 91
The arteriovenous pathway 92

II. The secondary blood system 96
III. The renal portal circulation 99

The urophysis 100
The bladder veins 100
Connections with the posterior intestine and rectum 101
Afferent and efferent flows to the kidney 102

IV. Red and white muscle 103
7 - Retial counter-current systems: flow-diffusion-concentration 106

Introduction 106
Retia that concentrate oxygen and nitrogen 107

The retia of the swimbladder 107
The removal of gas from the bladder 110
The choroidal rete of the eye 112

Retia that concentrate heat 114

The responses of the circulatory system to temperature change in fish lacking retia 114
Acclimation to temperature 114
The muscle retia of tuna 116
Muscle retia in the mackerel sharks 120
Retia that concentrate heat in the digestive organs 121
Retia that concentrate heat in the brain and eye 123
8 - Venous return and venous pumps 126

Venous pressure 126
The haemal arch pump 126
The branchial pump 130
The 'cardinal heart' of the myxinoids 130
The caudal pump 131
The caudal heart of the carpet shark 133
The caudal heart of the eel 135
The caudal heart of the hagfish 137
The portal heart of the hagfish 139
9 - The autonomic nervous system 141

Introduction 141
The parasympathetic and sympatho-adrenal systems 142

Adrenergic and cholinergic receptors 145
Antagonism and alliance in the autonomic system 147

Atrial natriuretic peptide (ANP) 149
Vascular receptors 150

Pressure or baroreceptors 150
Hypoxia receptors 152
Gill nocicep tors 15 3
10 - The response to exercise 158

Introduction 158
The response of the heart 159

The extent of the increase in cardiac output 159
The intrinsic mechanism: the Starling response 160
The role of the cardiac vagus 160
The role of circulating catecholamines 161
The role of cardioaccelerator nerves 162

The response of the gill vasculature 163

Intrinsic dilation of gill vessels 163
Parasympathetic nerve fibres to the gills 163
The response of the gill vessels to branchial efferent adrenergic fibre stimulation 164
The response of the gill vessels to centrally liberated catecholamines 165

The response of the peripheral vessels 166

Intrinsic responses 166
The response to circulating catecholamines 167
The role of the autonomic vasomotor nerves 167
In summary 168
11 - The response to hypoxia 170

Introduction 170
I. Reflex responses (A) The heart 171

The bradycardia of hypoxia 171
Cardiorespiratory synchrony 172
Changes in coronary blood flow 173

I. Reflex responses (B) The peripheral circulation 173
II. The intrinsic responses of the gill vessels 173
III. The central liberation of catecholamines 175

The defence of the acidotic myocardium 175
Catecholamines and the increase in the oxygen diffusive conductance of the gill 176
The role of catecholamines in regulating oxygen uptake by erythrocytes 177
Effects on the efflux of H+ from the gill 179
Effects on carbohydrate metabolism 180
In summary 181

12 - Myxine, a speculative conclusion 183

Introduction 183
The heart 185

The performance of the myxinoid heart 185
The gross anatomy of the myxinoid heart 185
Some features of the myxinoid cardiac cycle 186

The blood 188
The peripheral circulation 189

The arterial system 189
The venous system 190
The sinus system and the subcutaneous sinus 190
Pathways into the sinus system 191
Pathways out of the subcutaneous sinus 191
The haematocrit and volume of sinus blood 192
What is the role of the subcutaneous sinus? 193
Can the subcutaneous sinus be related to other subdermal vascular spaces? 194

Some final speculations 195

References 198
Appendix of popular and scientific names 227
Index 233




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