Preface to the first edition ................................. xiii
Preface to the second edition .................................. xv
Preface to the third edition ................................. xvii
Authors ....................................................... xix
Contributors .................................................. xxi
1 Physiology of excitable cells ................................ 1
Introduction ................................................. 1
Membrane potential ........................................... 1
Resting membrane potential ................................... 2
Action potentials ............................................ 7
Propagated action potential .................................. 9
Ionic basis of the cardiac action potential ................. 11
Action potential in muscle .................................. 13
Voltage-gated ion channels .................................. 13
Sodium channels ............................................. 14
Potassium channels .......................................... 16
Calcium channels ............................................ 16
Neurotransmitters and receptors: Ion channels, G proteins
and second messengers ....................................... 16
G proteins .................................................. 17
Second messengers ........................................... 18
Acetylcholine ............................................... 18
Catecholamines .............................................. 19
Amino acid transmitters ..................................... 20
Neuropeptide transmitters ................................... 20
Opioids ..................................................... 20
Neurotransmitter release in sympathetic ganglia ............. 20
Neuromuscular transmission .................................. 21
Overview of neuromuscular transmission ...................... 21
Muscle ...................................................... 24
Cardiac muscle .............................................. 28
Smooth muscle ............................................... 29
Muscle spindles, golgi tendon organs and spinal reflexes .... 30
Muscle spindles ............................................. 30
Golgi tendon organs ......................................... 32
Spinal reflexes ............................................. 32
Initiation of skeletal muscle contraction ................... 33
Sensory receptors: Classification ........................... 33
Mechanisms of receptor activation ........................... 33
Sensation ................................................... 34
Pain receptors .............................................. 34
Reflections ................................................. 35
2 Physiology of the nervous system ............................ 39
Neurons ..................................................... 40
Synaptic transmission ....................................... 42
Neurotransmitters ........................................... 45
Central and peripheral nervous systems ...................... 46
Cerebrospinal fluid ......................................... 47
Cerebral blood flow and oxygenation ......................... 50
Intracranial pressure ....................................... 50
Brain metabolism ............................................ 52
Classification of sensorimotor neurons ...................... 52
Sensory system .............................................. 52
Motor function and its control .............................. 57
Muscle tone ................................................. 63
Control of posture .......................................... 63
Electroencephalography ...................................... 63
Evoked potentials ........................................... 64
Consciousness ............................................... 64
Sleep ....................................................... 65
Autonomic nervous system .................................... 66
Hypothalamus ................................................ 66
Visceral afferent system .................................... 67
Autonomic ganglia ........................................... 68
Sympathetic nervous system .................................. 68
Parasympathetic nervous system .............................. 69
Neurotransmitters ........................................... 70
Receptors ................................................... 71
Reflections ................................................. 71
3 Respiratory physiology ...................................... 75
Functions of the respiratory system ......................... 76
Functional anatomy of airways, alveoli and pulmonary
capillaries ................................................. 76
Muscles of ventilation ...................................... 77
Mechanics of lung ventilation ............................... 77
Pressures and flow during the breathing cycle ............... 78
Elastic recoil and expansion of the lung .................... 78
Non-elastic forces and expansion of the lung ................ 80
Laminar, transitional and turbulent gas flows ............... 80
Lung volumes ................................................ 81
Pressure-volume relationships of the respiratory system ..... 82
Lung compliance ............................................. 84
Airway resistance ........................................... 85
Work of ventilation ......................................... 86
Gas exchange in the lungs ................................... 87
Rate of transfer of oxygen and carbon dioxide in arterial
and venous blood ............................................ 87
Diffusion of oxygen and carbon dioxide across the alveoli
and pulmonary capillaries ................................... 88
Alveolar ventilation and dead space ......................... 88
Anatomical, alveolar and physiological dead spaces .......... 88
Oxygen and carbon dioxide composition of alveolar gas ....... 90
Venous admixture (shunt) .................................... 93
Ventilation-perfusion ratio ................................. 94
Carriage of oxygen and carbon dioxide in the blood .......... 95
Oxygen carriage in the blood ................................ 97
Carbon dioxide carriage in the blood ........................ 99
Pulmonary circulation ...................................... 101
Control of ventilation ..................................... 103
Respiratory centre in the medulla .......................... 103
Central chemoreceptors ..................................... 104
Peripheral chemoreceptors .................................. 105
Other factors involved in control of ventilation ........... 106
Reflex ventilatory responses ............................... 107
Exercise and the control of ventilation .................... 109
Anaesthetic agents and the control of ventilation .......... 109
Breath holding ............................................. 109
Reflections ................................................ 110
4 Cardiovascular physiology .................................. 113
Functions and layout of the cardiovascular system .......... 114
Pumps and circuitry ........................................ 115
Distribution of blood volume in the cardiovascular system .. 116
Heart ...................................................... 117
Cardiac action potentials .................................. 119
Ionic basis of the cardiac action potential ................ 120
Conduction of the cardiac action potential ................. 123
Relationship between cardiac action potential, muscle
contraction and refractory periods ......................... 124
Excitation-contraction coupling in cardiac muscle cells .... 124
Electrocardiography ........................................ 125
Mechanical events of the cardiac cycle ..................... 127
Biophysical determinants of cardiac muscle contraction ..... 129
Studies of isolated cardiac muscle preparations ............ 130
Studies of mechanical performance of the whole heart ....... 133
Ventricular pressure-volume relationships .................. 134
Physical factors governing blood flow through vessels ...... 139
Observed physiological deviations from Poiseuille's
equation ................................................... 141
Pressure and flow .......................................... 143
Vessels of the systemic circulation ....................... 144
Arteries and arterial blood pressure ....................... 146
Arterioles ................................................. 148
Capillaries ................................................ 151
Lymphatics ................................................. 154
Veins and venous return .................................... 154
Relationship between venous return and cardiac output ...... 157
Control of the cardiovascular system ....................... 161
Functional organization of cardiovascular sympathetic and
parasympathetic nerves in the medulla ...................... 161
Central nervous system control and integration of the
cardiovascular system ...................................... 163
Efferent pathways and effectors ............................ 166
Sensors and measured variables ............................. 166
Short-term and long-term regulation of arterial blood
pressure ................................................... 169
Control of special circulations ............................ 169
Integrated cardiovascular responses ........................ 175
Valsalva manoeuvre ......................................... 178
Exercise ................................................... 179
Reflections ................................................ 182
5 Gastrointestinal physiology ................................ 187
Oral cavity ................................................ 187
Pharynx and oesophagus ..................................... 188
Stomach .................................................... 190
Small intestine ............................................ 197
Large intestine ............................................ 202
Reflections ................................................ 203
6 Liver physiology ........................................... 207
Anatomical aspects ......................................... 207
Functions of the liver ..................................... 209
Carbohydrate metabolism .................................... 209
Lipid metabolism ........................................... 210
Bile production ............................................ 211
Bilirubin metabolism ....................................... 211
Protein metabolism ......................................... 212
Phagocytic functions ....................................... 213
Storage functions .......................................... 213
Drug metabolism ............................................ 213
Liver blood flow ........................................... 214
Reflections ................................................ 217
7 Renal physiology ........................................... 219
Functions of the kidneys ................................... 220
Fluid and electrolyte balance and dietary requirements ..... 221
Functional anatomy of the kidneys .......................... 222
Glomerular filtration ...................................... 224
Control of renal blood flow ................................ 227
Tubular reabsorption and secretion ......................... 229
Renal clearance ............................................ 233
Loop of Henle and production of concentrated urine ......... 234
Summary of tubular handling of the glomerular filtrate ..... 237
Hormonal control of tubular function ....................... 239
Control of renal sodium, water and potassium excretion ..... 241
Renal control of acid-base balance ......................... 245
Mechanisms of action of diuretic drugs ..................... 248
Reflections ................................................ 249
Acid-base physiology ....................................... 251
Definitions ................................................ 251
pH system .................................................. 252
Buffers .................................................... 252
Hydrogen ion balance ....................................... 252
8 Acid-base homeostasis ...................................... 253
Whole-body (in vivo) titration curves ...................... 257
Compensatory mechanisms .................................... 257
Clinical effects of acid-base changes ...................... 260
Temperature and acid-base control .......................... 261
Clinical aspects of acid-base control ...................... 263
Reflections ................................................ 267
9 Physiology of blood ........................................ 269
Haemopoiesis and its control ............................... 269
Red blood cells ............................................ 270
White blood cells .......................................... 278
Platelets .................................................. 280
Coagulation ................................................ 284
Cell-based theory of coagulation ........................... 285
Blood transfusion .......................................... 287
Plasma ..................................................... 294
Reflections ................................................ 296
10 Physiology of the immune system ............................ 299
Innate immunity ............................................ 299
Acquired immunity .......................................... 307
Hypersensitivity ........................................... 317
Transplant immunology ...................................... 320
Assessment of immune function .............................. 320
Effects of anaesthesia on immune function .................. 323
Reflections ................................................ 324
11 Endocrine physiology ....................................... 325
Introduction ............................................... 325
Hormone production and secretion ........................... 326
Regulation of hormone secretion ............................ 326
Actions of hormones ........................................ 327
Hypothalamus ............................................... 328
Anterior pituitary ......................................... 330
Posterior pituitary ........................................ 333
Pancreatic islets .......................................... 334
Thyroid .................................................... 340
Calcium metabolism ......................................... 342
Adrenal cortex ............................................. 345
Adrenal medulla ............................................ 349
Erythropoietin ............................................. 352
Atrial natriuretic factor .................................. 353
Sex hormones ............................................... 354
Reflections ................................................ 354
12 Metabolism, nutrition, exercise and temperature
regulation ................................................. 357
Cellular respiration ....................................... 357
Energy compounds ........................................... 358
Adenosine triphosphate: The currency of cellular energy .... 359
Anaerobic or aerobic metabolism ............................ 359
Basic metabolic pathways ................................... 360
Metabolism ................................................. 363
Catabolic pathways ......................................... 363
Anabolic pathways .......................................... 369
Control of metabolic pathways .............................. 370
Nutrition .................................................. 370
Interrelation between fat and carbohydrate metabolism ...... 372
Dietary energy sources ..................................... 372
Basal metabolic rate ....................................... 373
Starvation ................................................. 374
Exercise ................................................... 375
Cardiovascular responses to exercise ....................... 380
Respiratory responses to exercise .......................... 381
Muscle and bone responses to exercise ...................... 381
Gastrointestinal and endocrine effects ..................... 381
Temperature regulation ..................................... 382
Afferent temperature sensors ............................... 382
Central regulation ......................................... 382
Efferent responses ......................................... 383
Cutaneous responses to heat ................................ 383
Effects of anaesthesia on thermoregulation ................. 385
Physiology of altered temperature and the thermoneutral
zone ....................................................... 385
Responses to hypothermia ................................... 385
Responses to high temperatures ............................. 387
Reflections ................................................ 387
13 Physiology of pain ......................................... 391
Introduction ............................................... 391
Peripheral mechanisms of pain .............................. 392
Reflections ................................................ 405
14 Maternal and neonatal physiology ........................... 409
Maternal physiology ........................................ 409
Demands of pregnancy ....................................... 409
Physiology of the placenta ................................. 417
Perinatal physiology ....................................... 421
Foetal circulation ......................................... 421
Foetal respiratory system .................................. 424
Haematology ................................................ 426
Acid-base status ........................................... 427
Renal function ............................................. 427
Liver function ............................................. 428
Metabolic balance .......................................... 428
Nervous system ............................................. 428
Thermoregulation ........................................... 428
Reflections ................................................ 430
15 Physiology of ageing ....................................... 433
Functional decline with ageing ............................. 433
Changes in the nervous system .............................. 434
Changes in the cardiovascular system ....................... 434
Changes in the respiratory system .......................... 435
Changes in body compartments ............................... 436
Changes in renal function .................................. 436
Changes in liver function .................................. 437
Endocrine changes .......................................... 437
Thermoregulation ........................................... 437
Reflections ................................................ 437
16 Special environments ....................................... 439
Physiology of diving ....................................... 439
Physical laws .............................................. 439
Direct effects of increased pressure ....................... 439
Effects of breathing hyperbaric gases ...................... 441
Physiological effects of altitude .......................... 442
Hypobaric environments ..................................... 442
Effects of rapid ascent to altitude ........................ 442
High-altitude residents .................................... 444
Physiology of space travel ................................. 445
Physiological effects of weightlessness .................... 446
Reflections ................................................ 446
Key equations and tables ...................................... 449
Further reading ............................................... 459
Index ......................................................... 461
|
