The idea for this book grew out of regular discussions between the authors when we were both on the staff of the Department of Physiology, Royal Free Hospital School of Medicine in London icon

The idea for this book grew out of regular discussions between the authors when we were both on the staff of the Department of Physiology, Royal Free Hospital School of Medicine in London




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Preface

The idea for this book grew out of regular discussions between the authors when we were both on the staff of the Department of Physiology, Royal Free Hospital School of Medicine in London. We felt that there was a need for a modern, concise textbook of physiology which covered all aspects of the preclinical course in physiology. The text is written primarily for students of medicine and related subjects, so the clinical implications of the subject are deliberately emphasized. Nevertheless, we hope that the book will also prove useful as core material for first- and second-year science students. We have assumed a knowledge of chemistry and biology similar to that expected from British students with 'AS' levels in these subjects. Our intention has been to provide clear explanations of the basic principles that govern the physiological processes of the human body and to show how these principles can be applied to the understanding of disease processes.

The book begins with cell physiology (including some ele­mentary biochemistry), and proceeds to consider how cells inter­act both by direct contact and by longer-distance signaling. The nervous system and endocrine system are dealt with at this point. The physiology of the main body systems is then discussed. These extensive chapters are followed by a series of shorter chapters describing integrated physiological responses, including the con­trol of growth, the regulation of body temperature, the physio­logy of exercise, and the regulation of body fluid volume. The final chapters are mainly concerned with the clinical applications of physiology, including acid—base balance, heart failure, hyper­tension, liver failure, and renal failure. This structure is not a reflection of the organization of a particular course but is intend­ed to show how, by understanding the way in which cells work, and how their activity is integrated, one can arrive at a satisfying explanation of body function.

In providing straightforward accounts of specific topics, it has occasionally been necessary to omit some details or alternative explanations. Although this approach occasionally presents a pic­ture that is more clear-cut than the evidence warrants, we believe that this is justified in the interests of clarity. Key points are illus­trated by simple line drawings as we have found that they are a useful aid to students in understanding and remembering import­ant concepts. We have not included extensive accounts of the experimental techniques of physiology but have tried to make clear the importance of experimental evidence in elucidating

underlying mechanisms. As far as possible, normal values have been given throughout the text in SI units but important physio­logical variables have also been given in traditional units (e.g. mmHg for pressure measurements).

Each chapter is organized in the same way. In answer to the frequently heard plea 'what do I need to know?' we have set out the key learning objectives for each chapter. This is followed, where appropriate, by a brief account of the physical and chemical principles required to understand the physiological processes under discussion. The essential anatomy and histology are then discussed, as a proper appreciation of any physiological process must be grounded on a knowledge of the anatomical features of the organs involved. Detailed discussion of the main physio­logical topics then follows.

To aid student learning, short, numbered summaries are given after each major section. From time to time we have set out important biological questions or major statements as section headings. We hope that this will help students to identify more clearly why a particular topic is being discussed. The reading material given at the end of each chapter is intended both to pro­vide links with other subjects commonly studied as part of the medical curriculum and to provide sources from which more detailed information can be obtained. Self-testing is encouraged by the provision of multiple choice questions or quantitative problems (or both) at the end of chapters. Annotated answers to the questions follow. Some numerical problems have also been given which are intended to familiarize students with the key formulae and to encourage them to think in quantitative terms.

We are deeply indebted to Professor Michael de Burgh Daly and Dr Ted Debnam, who not only advised us on their specialist topics but also read through and constructively criticized the entire manuscript. Any remaining obscurities or errors are entire­ly our responsibility. Professor de Burgh Daly has also kindly provided a foreword to this book. Finally, we wish to thank the staff of Oxford University Press for their belief in the project, their forbearance when writing was slow and their help in the realizarion of rhe final product.

G.P. C.D.R.

London February 1999

Contents

Foreword by M. de Burgh Daly page xi Acknowledgements xiii Abbreviations xv A note to the reader xvii

  1. What is physiology? 1

  2. The chemical constitution of the body 7

  3. Introducing cells 21

  4. The transport functions of the plasma membrane 33

  5. Principles of cell signaling 51

  6. Nerve cells and their connections 65
    1
    Muscle 81




  1. Sensory systems 105

  2. The physiology of motor systems 147




  1. The autonomic nervous system 171

  2. Some aspects of higher nervous function 181

  3. The hormonal regulation of the body 197

  4. The properties of blood 233

  5. Defense against infection: inflammation and immunity 259

  6. The heart and circulation 275

  7. The respiratory system 323

  8. The kidney and the regulation of the internal environment 359

  9. The gut and nutrition 391

  10. The physiology of the male and female reproductive systems 435

  11. Fertilization and pregnancy 45 7

  12. Fetal and neonatal physiology 473

  13. Lactation 491

  14. The control of growth 501

  15. Energy balance and the control of metabolic rate 517

  16. The physiology of exercise 525

  17. The regulation of body temperature 53 7

  18. The regulation of plasma glucose 547

  19. The regulation of body fluid volume 559

  20. Acid—base balance 5 73

  21. The physiology of high altitude and diving 591

  22. Clinical physiology 599
    Appendix 611

Index 613

Foreword

It is a pleasure to have been invited to write a foreword to the first edition of this textbook which describes the fundamental prin­ciples of human physiology. It is written by two experienced teachers in the field and it is only through the cognizance gained over the years that the needs of students are appreciated. They have borne in mind the recommendation of the General Medical Council that the factual load burdened on students should be significantly reduced to provide an opportunity for the provision of subjects which engage critical, philosophical, and creative fac­ulties. Thus, the authors have provided basic information about the subject, and a text that is easy to read and to follow, that is neither too didactic nor too controversial. At the same time, it largely avoids the complexities of mathematical equations which are apt to deter all but those few who have a propensity for approaching problems in this way.

It has long been appreciated that a good understanding of physiology requires a sound background knowledge of functional anatomy. This was fully appreciated many years ago by Britain's most distinguished physician, William Harvey (1578-1657), who is well known for his discovery of the circulation of the blood and is considered to have been the first experimental physiologist. It was the combination of experimental evidence and the applica­tion of anatomical knowledge that enabled him to make this important discovery, which was to have such an impact on clini­cal medicine. From his experimental evidence he correctly deduced that there existed small blood vessels connecting arteries and veins through which blood flowed after entering a limb by the arteries before returning via veins to the right side of the heart. Even so, because no microscope was in existence at that time, he was unable to confirm his deduction. The subsequent discovery of the capillaries is a good example of the way in which histological and, later, electron microscopical studies, have con­tributed so much to the advance of physiological knowledge. Certain aspects of biochemistry and pharmacology have also made valuable contributions. These subjects cannot, in my opinion,

be as divorced from physiology as some might wish, and their sprinkling, where appropriate, in different parts of the present text substantiates this view.

This textbook follows the general format of others in dealing with individual aspects of physiology as major 'systems'. Past experience has shown that this is one of the simplest ways of presenting the subject, but students have to appreciate that the functioning of the body as a whole in health is more complex than this; it is determined by the integration of the functioning of two or more systems simultaneously. This is underlined in the present text by numerous cross-references to relevant sections.

The reader will find examples of the applications of physiology to practical problems and to clinical medicine attractive, par­ticularly as there is a trend, which is gathering momentum, of putting more clinical material into the preclinical course. How­ever, this textbook cannot and was never intended to be a definitive treatise in clinical physiology as well as basic physio­logy; that subject in its entirety warrants separate treatment. Thus the authors have had to achieve, successfully I believe, what I regard as a difficult balancing act of providing the student with a sound scientific background in physiology while at the same time covering some of its applications to problems encountered in applied physiological situations and in clinical medicine. As pre­clinical students always come to realize, the practice of medicine relies heavily on a thorough grounding in physiology!

Most courses in physiology consist of a series of formal lectures (handouts sometimes being provided), tutorials, and practical work. These provide students with a good insight to physiology as an experimental discipline. But these parts of the course serve another important function: they act as a guide to students' reading about the subject, for which this textbook will be found indispensable.

^ Royal Free Hospital School of Medicine, London M. de Burgh Daly 1998

Acknowledgments

We wish to acknowledge many colleagues who have helped us to clarify out thinking on a wide vatiety of topics. Those to whom especial thanks ate due fof detailed criticisms on particular chapters are:

Professor J. F. Ashmore FRS, Department of Physiology, University College London, UK

Professor S. ^ Bevan, The Novartis Institute of Medical Sciences, 5 Gower Place, London, UK

Dr T. V. P. Bliss FRS, Division of Neurophysiology, The National Institute for Medical Research, Mill Hill, London, UK

Dr E. S. Debnam, Department of Physiology, Royal Free and University College Medical School, University College London, UK

^ Professor M. de Burgh Daly, Department of Physiology, Royal Free and University College Medical School, University College London, UK

Professor D. A. Eisner, Department of Preclinical Veterinary Sciences, University of Liverpool. Liverpool, UK

Dr B. D. Higgs, Division of Anaesthesia, Royal Free Hospital, London, UK

Dr A. Ionnadis, Royal Free Hospital, London, UK

^ Dr. A. Mathie, Department of Pharmacology, University College London, UK

Dr D. A. Richards, Department of Physiology and Biophysics, University of Colorado Health Sciences Center, Denver, Colorado, USA

Professor I. C. A. F. Robinson, Laboratory of Endocrine Physiology, The National Institute for Medical Research, Mill Hill, London, UK

Dr A. H. Short, Clinical Skills Laboratory, Medical School, Queen's Medical Centre Nottingham, UK

The authors wish to thank all those people who have granted permission to reproduce figures from books and original articles, either unmodified or in a modified form. They are:

Figures 3.1 & 14.11 are from Figures 1.6 and 8.8 of J. M. Austyn & K. J. Wood Principles of Cellular and Molecular Immunology

(1993) Oxford University Press, Oxford. Figure 3.5 is from Figure 2-18 of B. Alberts, D. Bray, J. Lewis, M. Raff, K. Roberts & J. D. Watson Molecular Biology of the Cell. 3rd Edition, (1989) Garland, New York. Figures 3.6 & 3.7 are from Figures 8.7 & 8.13 of W. H. Elliott, & D. С Elliott, Biochemistry and Cell Biology. (1997) Oxford University Press, Oxford. Figure 4.3 is based on Figure 13 of P. С Caldwell, A. L. Hodgkm, R. D. Keynes & T. I. Shaw, Journal of Physiology (I960) vol. 152 pp. 561—590. Figure 4.7 is from an original figure of Dr E. S. Debnam. Figure 4.10 is from an original figure of Dr P. Charles worth. Figures 5.3 & 15.35 are from Figures 2.3 & 13-6 of H. P. Rang, M. M. Dale & J. M. Ritter, Pharmacology. 3rd Edition (1995), Churchill-Livingstone, Edinburgh. Figures 6.3, 6.5, 8.3, 8.9, 8.10, 8.27, 8.40, 9.10, 9.15, 9-18 9.19, 9.20, 15.39 & 15.40 are based on Figures 2.6, 2.7, 2.28, 4.1, 14.13, 14.14, 13.24, 13.26, 13.12, 9.1, 11.2, 11.13, 10.2, 10.3, 1.15, 2.38 & 2.39 of P. Brodal, The Central Nervous System. Structure and function. (1992) Oxford University Press, Oxford. Figure 6.9 is based on Figure 4 of A. L. Hodgkin & B. Katz, Journal of Physiology (1949) vol. 108 pp. 37-77. Figure 6.10 is based on Figure 17 of A. L. Hodgkin & A. F. Huxley, Journal of Physiology (1952) vol. 117 pp. 500-540. Figure 6.19 is based on Figure 1 of P. Fatt & B. Katz, Journal of Physiology (1952) vol. 117 pp. 109-128. Figure 7.1 is based on Figure 11-19 of W. Bloom & D. W. Fawcett, Textbook, of Histology (1975). W. B. Saunders & Co. Figure 7.11 is based on Figures 12 & 14 of A. M. Gordon, A. F. Huxley & F. J. Julian, Journal of Physiology (1966) vol. 184 pp. 170-192. Figures 8.5, 8.7, 8.39, 8.41 & 8.42 are based on Figures 16.3, 16.6, 16.10, 17.2 & 17.6 of H. B. Barlow & J. D. Mollon, (Eds) (1982) The Senses. Cambridge University Press. Figure 8.8 is based on Figure 30.6 of D. Ottoson, The Physiology of the Nervous System (1984) Macmillan Press, London. Figures 8.15, 8.21 & 11.2 are based on Figures 7.3, 7.18 & 13.4 of R. H. S. Carpenrer, Neurophysiology, 3rd Edition (1996). Edward Arnold, London. Figure 8.17 is based on Figure 28-6 of E. R. Kandel, J. H. Schwartz & Т. М. Jessell (Eds) Principles of Neural Science, 3rd Edition (1991). Elsevier Science, New York. Figure 8.18 is from Figure 4.5 of R. F. Schmidt (Ed.), Fundamentals of Sensory Physiology 3rd Edition, (1986) Springer-Verlag, Berlin. Figure 8.25 is from P. H. Schiller, Trends in Neurosciences (1992) vol. 15 p. 87 with permission from Elsevier Science. Figure 8.30 is based on Figures 2.4 and 2.5 of J. O. Pickles, An Introduction to the Physiology of Hearing (1982)

XIV

Acknowledgments


Academic Press, London. Figure 8.33 is part based on Figure 5 of I. M. Russell & P. M. Sellick, Journal of Physiology vol. 284 pp. 261-290. Figure 8.34 is part based on Figure 1 of J. O. Pickles & D. P. Corey, Trends in Neurosciences vol. 15 p. 255, with permission of Elsevier Science. Figure 8.35 is based on data of E. F. Evans. Figure 8.38 is adapted from Lindeman Ergebnisse der Anatomie vol. 42 pp. 1—113 (1969)- Figure 10.2 is adapted from Figure 18.2 of G. M. Shepperd, Neurobiology, Oxord University Press, Oxford after Heimer. Figure 11.3 is from Figure 6.3 of S. P. Springer & G. Deutsch Left Brain, Right Brain, 3rd Edition (1989). W. H. Freeman & Co. New York. Figure 11.11 is from original data of Dr D. A. Richards. Figures 12.9, 12.12, 12.16 & 12.19 are from Figures 3.4, 5.2, 4.1 & 4.12 in С Brook and N. Marshall, Essential Endocrinology (1996). Blackwell Science, Oxford. Figure 13.4 is from an original figure of Dr E. S. Debnam. Figures 14.1, 14.7 & 14.10 are based on Figures 9-1, 9.2, 13.1 & 13.2 of J. H. Playfair, Infection & Immunity, (1995) Oxford University Press, Oxford. Figures 15.1, 15.17, 15.26 & 15.37 are based on Figures 1.4, 1.6, 9.2 & 13.6 of J. R. Levick An Introduction to Cardiovascular Physiology, 2nd Edition (1995), Butterworth-Heinemann, Oxford. Figures 15.4, 15.10, 15.34, 16.1, 16.5 & 19.7 are from Figures 5.4.10, 5.4.6, 5.4.7, 5.3.3, 5.3.4 & 6.10.3 of P. С. В. MacKinnon & J. F. Morris (1988) Oxford Textbook of Functional Anatomy. Volume 2. Thorax and Abdomen, Oxford University Press, Oxford. Figure 15.15 is based on Figures 4 & 10 from O. E Hutter & W. Trauwein (1956) Journal of General Physiology vol. 39, pp. 715—733 by copyright per­mission of The Rockefeller University Press. Figures 15.21 is based on Figure 19-14 of A. C. Guyton, Textbook of Medical Physiology, 7th Edition (1986), W. B. Saunders Co. Figure 15.22 is based on Figure 7 of A. E. Pollack & E. H. Wood {19^9) Journal of Applied Physiology vol. 1, pp. 649-662 by permission of the American Physiological Society. Figure 15.25 is based on Figure 5 of L. H. Smaje, В. Н. Zweifach & M. Intaglietta, (1970) Microvascular Research, vol. 2 pp. 96—110. Figure 15.31 is based on Figure 5—9 of R. F. Rushmer, Cardiovascular Dynamics (1976), W. B. Saunders. Figures 16.2, 16.11, 16.18, 16.26 are based on Figures 1.5, 2.4, 5.4 & 8.5 of J. Widdicombe & A. Davies, Respiratory Physiology (1991), Edward Arnold, London. Figure

16.4 is based on Figure 11.1 of E. R. Weibel The Pathway for Oxygen. Structure and Function of the Mammalian Respiratory System (1984) Harvard University Press. Figures 16.7, 16.8, 16.9, 16.11, 24.2 & 24.3 are based on Figures III.4, III.3, III.ll, III.12, III.80 & 111.81 of С A. Keele, E. Neil & N. Joels Samson Wright's Applied Physiology, 13th Edition (1982), Oxford University Press, Oxford. Figure 16.14 is adapted from Figure 3—8 of M. G. Levitzky, Pulmonary Physiology 3rd Edition (1991)- McGraw-Hill, New York. Figures 16.17 & 25.3 are from original figures of Professor M. de Burgh Daly. Figure 16.19 is from Figure 19 of J. B. West, WentilationlBlood flow and Gas Exchange, Blackwell Science, Oxford. Figure 16.24 is from Figure 39 of P. Dejours Respiration (1966), Oxford University Press, Oxford. Figures 17.2 and 17.5 are based on Figures 1 & 2 of W. Kritz & L. Bankir, American Journal of Physiology vol. 254 pp. F1-F8 (1988) by permission of the American Physiological Society. Figure 17.4 is based on Figure 2.3 of В. М. Koeppen & B. A. Stanton, Renal Physiology (1992) Mosby, St Louis. Figure 18.3 is based on Figure 21—13 of D. F. Moffett, S. B. Moffett & C. L. Schauf, Human Physiology (1993) McGraw-Hill. New York. Figure 18.14 is adapted from Figure 13 of J. H. Szurszewiski, Journal of Physiology, vol. 252 pp. 335-361. Figure 18.28 is from Figures 6.1 & 6.2 of P. A. Sanford, Digestive System Physiology 2nd Edition (1992) Edward Arnold, London. Figure 21.5 is adapted from Figure 13.1 (after Smith, 1963) of D. J. Begley, J. A. Firth & J. R. S. Hoult, Human Reproduction and Developmental Biology (1980) Macmillan Press, London. Figures 21.10 & 21.11 are from Figures 7.3 & 7.2 of N. E. Griffin & S. R. Ojeda, Textbook of Endocrine Physiology 2nd Edition (1995). Oxford University Press, Oxford. Figure 23.1 is from Figure 16 of J. M. Tanner, Foetus into Man. 2nd Edition (1989) Castlemead, London. Figures 25.1 & 25.2 are from Figures 7.3 and 7.10 of P-O. Astrand & K. Rodahl, Textbook of Work Physiology, Physiological bases of exercise. 3rd Edition (1986). McGraw-Hill. New York. Figure 26.3 is from Figure 1 of J. Werner Pflugers Archiv vol. 367 pp. 291-294 (1977). Figure 30.2 is from Figure 1 of N. Pace, B. Meyer & В. Е. Vaughan, Journal of Applied Physiology vol. 9 pp. 141-144 (1956) by per­mission of the American Physiological Society.

List of abbreviations





ACh

acetylchohne

epp

ACTH

adrenocorticotropic hormone (corticotropin)

epsp

ADH

antidiuretic hormone (vasopressin)

ER

ADP

adenosine diphosphate

ESV

AMP

adenosine monophosphate

FAD

ANP

atrial natriuretic peptide

FADH2

ATP

adenosine triphosphate

FEV,

ATPS

ambient temperature and pressure saturated with

FRC




water vapor (with reference to respiratory gas)

FSH

AV

atrioventricular

FVC

av

arteriovenous

GABA

b.p.m.

beats per minute

GDP

BER

basic electrical rhythm

GFR

BMR

basal metabolic rate

GH

BP

blood pressure

GHIH

BTPS

body temperature and pressure saturated (with

GHRH




water vapor)

GI

С

gas content of blood (e.g. CyO2)

GIP

CCK

cholecystokinin

GMP

CN

cranial nerve

GnRH

CNS

central nervous system

G-Protein

CO.

cardiac output

GTP

CoA

coenzyme A

Hb

CRH

corticotropin-releasing hormone

HbF

c.s.f.

cerebrospinal fluid

HbS

CVP

central venous pressure

hCG

DAG

diacylglycerol

HIV

dB

decibel

HLA

DIT

di-iodotyrosine

hPL

DNA

deoxyribonucleic acid

HPNS

2,3-DPG

2,3-diphosphoglycerate

HRT

ECF

extracellular fluid

HSL

ECG

electrocardiogram

ICF

ECV

effective circulating volume

ICSH

EDTA

ethylenediaminetetraacetic acid




EDV

end-diastolic volume

IgA

EEG

electroencephalogram

IGF-l.IGF-2

ENS

enteric nervous system




end-plate potential

excitatory postsynaptic potential

endoplasmic reticulum

end-systolic volume

flavine adenine dinucleotide

reduced flavine adenine dinucleotide

forced expiratory volume at 1 second

functional residual capacity

follicle-stimulating hormone

forced vital capacity

y-aminobutyric acid

guanosine diphosphate

glomerular filtration rate

growth hormone (somatotropin)

growth hormone-inhibiting hormone

growth hormone-releasing hormone

gastrointestinal

gastric inhibitory peptide

guanosine monophosphate

gonadotropin-releasing hormone

heterotrimeric GTP-binding protein

guanosine triphosphate

hemoglobin

fetal hemoglobin

sickle-cell hemoglobin

human chorionic gonadotropin

human immunodeficiency virus

human leukocyte antigen

human placental lactogen

high-pressure nervous syndrome

hormone replacement therapy

hormone-sensitive lipase

intracellular fluid

interstitial cell stimulating hormone (identical

to luteinizing hormone)

immunoglobulin A

insulin-like growth factor 1, insulin-like growth

factor 2

^ List of abbreviations





IgG

immunoglobulin G

PTH

IgE

immunoglobulin E

RBF

IgM

immunoglobulin M

REM

IP,

inositol trisphosphate

Rh

ipsp

inhibitory postsynaptic potential

RNA

LH

luteinizing hormone

RPF

LTB4

leukotriene В,

RQ

LTP

long-term potentiation




MALT

mucosa-associated lymphatic tissue

RV

MAP

mean arterial pressure

SA

mepp

miniature end-plate potential

SPL

мне

major histocompatibility complex

STP

MIH

mullerian inhibiting hormone

STPD

MIT

mono-iodotyrosine

sv

MMC

migrating motility complex

sws

mph

miles per hour

г

Mr

relative molecular mass

T3

MSH

melanophore stimulating hormone

т4

MVV

maximum ventilatory volume

TH

NAD

nicotinamide adenine dinucleotide

тт

NADH

reduced nicotinamide adenine dinucleotide

TPA

NTS

nucleus of the tractus solitarius

TPR

P

pressure (see Box 16.1 for explanation of

TSH




subscripts)

V

p5o

pressure for half saturation




PAH

^-aminohippuric acid

V

PGE2

prostaglandin E2

vc

PGF

prostaglandin F

VIP

PIH

prolactin inhibitory hormone (dopamine)

V/Q

PRL

prolactin




parathyroid hormone

renal blood flow

rapid eye movement

rhesus factor (D antigen)

ribonucleic acid

renal plasma flow

respiratory quotient (also known as the

respiratory exchange ratio)

residual volume

sinoatrial

sound pressure level

standard temperature and pressure

standard temperature and pressure dry

stroke volume

slow-wave sleep

absolute temperature

tn-iodothyronine

thyroxine

thyroid hormone

transport maximum

tissue plasminogen activator

total peripheral resistance

thyroid stimulating hormone

volume (usually of gas, see Box 16.1 for

explanation of subscripts)

flow rate

vital capacity

vasoactive intestinal polypeptide

ventilation /perfusion ratio (in lungs)

^ A note to the reader

The chapters in this book cover the physiological material normally taught in the first and second years of the medical cur­riculum and degree courses in physiology. While each chapter can be read on its own, the book has been laid out as follows: Chapter 1 is a broad introduction to the subject. Chapters 2—4 present basic information on the properties of cells and how they com­municate. Chapter 5 deals with the mechanisms by which the body is able to coordinate and regulate the activities of its various parts. Chapters 6-18 include much of the core material of tra­ditional courses in physiology and discuss the functioning of the principal organ systems. Chapters 19—22 are concerned with the physiology of reproduction and that of the neonate. Chapters 23-29 are concerned with the interactions between different organ systems. In an effort to bridge the gap between the basic-science taught in the preclinical years and clinical practice, the final chapters are devoted to applied respiratory physiology and clinical applied physiology.

Each chapter begins with a list of learning objectives which set out the principal points we think you, the reader, should try to assimilate. We have assumed a basic knowledge of chemistry and biology, but important physical topics are discussed briefly where necessary. Key terms and definitions are given in italics where they first occur. The contents of each chapter are arranged in numbered

sections in the same order as the learning objectives, and each major section ends with a summary of the main points. We have tried to avoid repetition as far as possible by cross-referencing. Many chapters have boxes which contain more advanced material or deal with numerical examples. It is not necessary to read these boxes to understand the core material.

At the end of each chapter there is a reading list which is intended to link physiology with other subjects in the medical curriculum (particularly with anatomy, biochemistry, and phar­macology). These have been chosen for their clarity of exposition but many other good sources are available. Some more advanced texts are also included in the list for rhose who wish to study a particular topic in greater depth.

Many chapters end with a set of problems. These are mainly in the form of multiple-choice questions but some numerical problems are also included. Do make the effort to test your knowledge—it will help to lodge the key information in your mind. Answers (with explanations where appropriate) are given at the end of the chapters. When you find that a particular topic is difficult to understand, break it down into its components to identify where your difficulties lie. This is the first step towards resolving them. If, after further study, you still have difficulty, seek help from your tutor or lecturer.

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