A manual of physiology, with practical exercises

Front matter

New York

State College of Agriculture

At Cornell University

Preface To The Seventh Edition

In the present edition the book has been extensively revised. The rapid progress of biochemistry has rendered it necessary to enlarge greatly and practically to rewrite the chapter on Metabolism. Many changes and additions have also been made in the chapters on Circulation, Respiration, Digestion, Absorption, and Internal Secretion. The blood-gases are considered in much greater detail than in the last edition, and more space is devoted to the general phenomena of the action of enzymes. The newer work on the relation of the heat production and the chemical changes in muscle to the contraction has been taken account of. The chapters on the Nervous System have been brought up to date. The arrangement of the book has been improved, it is hoped, by breaking the longer chapters up into sections, and increasing the number of chapters. Many new illustrations have been added, and many of the old ones redrawn.

G. N. STEWART.

Cleveland,

August, 1914.

EXTRACT FROM THE PREFACE TO

THE FIRST Edition

In this book an attempt has been made to interweave formal exposition with practical work, according to a programme which I have followed for some time past in teaching Physiology to medical students on the other side of the Atlantic, and which has, it is believed, proved to be well adapted to their needs and opportunities. It ought, however, to be explained that, for various reasons, a somewhat wider range of experiment is open to the student in America than in this country. But as nobody will use this book except in a regular laboratory and under responsible guidance, it has not been thought necessary to mark in any special manner the parts of the exercises which the Enghsh student must do by proxy (that is, learn from demonstrations), and the parts he ought to perform for himself.

An arrangement of the exercises with reference to the systematic course has this advantage — that by a little care it is possible to secure that practical work on a given subject shall actually be going on at the time it is being expounded in the lectures. Cross-reference from lecture-room to laboratory, and from laboratory to lecture-room, from the detailed discussion of the relations of a phenomenon to the living fact itself, is thus rendered easy, natural, and fruitful.

As some teachers may wish to know how a course such as that described in the Practical Exercises may be conducted for a fairly large class, a few words on the method, we have followed may not be out of place. It is obvious that many of the exercises require more than one person for their performance; and it may be said

Viii EXTRACT FROM THE PREFACE TO THE FIRST EDITION

that, except in the case of the simpler experiments and the chemical work as a whole, which each student does for himself, it has been found convenient to divide the class into groups of four, each group remaining together throughout the session. It is possible that some may find a group of four too large a unit, and it is certain that three, or perhaps even two, would be better; but in a large school so minute a subdivision is hardly possible, without entailing excessive labour on the teachers.

The systematic portion of the book is so arranged that it can equally well be used independently of the practical work, and aims at being in itself a complete exposition of the subject, adapted to the requirements of the student of medicine.

As to the matter of the text, it is hardly necessary to say that this book does not aspire to the dubious distinction of originality ; and it is literally impossible to acknowledge all the sources from which information has been derived. In many cases names have been quoted, but names no less worthy of mention have often been of necessity omitted.

G. N. STEWART.

Cambridge,

September, 1895.

Section II. — General Physical and Chemical Properties of THE Blood 23

Viscosity of blood 23

Reaction of blood 24

Specific gravity of blood 26

Electrical conductivity of blood 26

Relative volume of corpuscles and plasma 27

Haemolysis 28

Agglutination - 30

Precipitins - 31

Anaphylaxis 32

Coagulation of blood t 33

Vaso-constrictor property of shed blood - 45

Section II. — ^The Beat of the Heart in its Physical or Mechanical Relations 85

Events in the cardiac cycle 85

The sounds of the heart - 88

The cardiac impulse 9°

Endocardiac pressure 92

The ventricular pressure-curve 94

The auricular and venous pressure-curve - - 98

Section HI. — Physical or Mechanical Phenomena of the

Circulation in the Bloodvessels - 10 1

The arterial pulse loi

Arterial blood-pressure 109

Measurement of the blood -pressure in man 113

Velocity of the blood - 117

Measurement of velocity of blood 120

The volume-pulse 127

The circulation in the capillaries 129

The circulation in the veins ' 132

The circulation-time 135

Work and output of heart 139

Section IV.— The Heart-Beat in its Physiological Rela^ TIONS

Intrinsic nerves of the heart - Cause of the heart-beat

Conduction and co-ordination in heart 146

Auriculo-ventricular bundle 147

Fibrillary contractions 15 1

Chemical conditions of heart-beat - 152

Resuscitation of the heart 153

Refractory period of heart 155

140 141 141

CONTENTS xi

PAGE

Section V. — ^The' Nervous Regulation of the Heart (Extrinsic Venous Mechanism of the Heart) 156 Action of poisons on the heart 164 Normal excitation of cardiac nervous mechanism - i65

Section I. — ^Preliminary Anatomical Data - 221 Physiological anatomy of the respiratory apparatus 222

Blood-supply of the lungs 222

Section II. — ^Mechanical Phenomena of External Respiration - - 224 T3rpes of respiration - 228 Artificial respiration 229 Respiratory sounds - 230 Frequency of respiration 233 Vital capacity - 235 Intrathoracic pressure 235 Respiratory pressure 237

Section VI. — Relation of Respiration to the Nervous System 268

The respiratory centre and its connections 268

Regulation of respiration through the vagus - 270

Action of other afferent fibres on the respiration 274

The chemical regulation of the respiration 275

Apnoea 277

Automaticity of the respiratory centre 278

Special modifications of the respiratory movements - 281

Section III. — Chemistry of the Digestive Juices — Ferments 330 Saliva 338

Gastric juice - 343

Antiseptic functions of gastric juice - - 350

CONTENTS xiii

PAGE

Section III. (continued) —

Pancreatic juice - 352

Bile 357

S11CCUS entericus - 364

Section IV. — Secretion of the Digestive Juices — ^Microscopical Changes in the Gland Cells- 368 Changes in pancreas and parotid during secretion 369 Changes in gastric glands during secretion 371 Changes in mucous glands during secretion 375 Mode of formation of the digestive juices 377 Why the tissues of digestion are not affected by the digestive fennents - 382

Section V. — Influence of the Nervous System on the Digestive Glands - 385

Influence of nervous system on salivary glands 385

Influence of nervous system on gastric glands 395

Influence of nervous system on the pancreas - 399

Secretin - 401

Influence of nervous system on secretion of bile 405 Influence of nervous system on the secretion of intestinal

juice - 408

Secretion of the digestive juices (summary) 409

Section III. — ^Metabolism of Proteins - - 563 Living and dead proteins - - - 566

Formation of amino-acids from tissue-proteins - - 569

Formation of hippuric acid - . - - ^^i

CONTENTS XV

Section III. (contimted) — page

Formation of urea 573

Formation of uric acid - 579

; Metabolism of nucleic acids and purin bases - 582 Kreatin and kreatinin - : 587

Autolysis 588

Section IV. — Statistics of Nutrition — Income and Expenditure OF THE Body in Terms of Matter 590 The nitrogen balance-sheet — ^nitrogenous equilibrium 591 Relation of nitrogenous metabolism to muscular work 599 Relative value of different proteins in nutrition 601 The carbon balance-sheet 606 The oxygen deficit 608 Inorganic salts in nutrition G08

Section III. — Physical and Mechanical Phenomena of the

Muscular Contraction 716 Optical phenomena — structure of muscle 716 Mechanical phenomena 719 Muscular fatigue 723 Electrical tetanus 730 Voluntary contraction . 734 Thermal phenomena and transformation of energy in muscular contraction - 736 Relation between mechanical energy and heat-production in active muscle 739

Section IV. — Chemical Phenomena of Muscular Contraction 741 Formation of lactic acid 743

The substances metabolized in muscular contraction 745

Chapter XIV. Nerve

Section I. — The Nerve-Impulse or Propagated Disturbance: its Initiation and Conduction 755 Stimulation of nerve 757 Excitability of nerve 758 Electrotonus - 759 Conduction in nerve - - 764 Velocity of the nerve-impulse 767

Section II. — Chemistry, Degeneration, and Regeneration

OF Nerve 768

Chemistry of nerve 768

Degeneration of nerve - 769

Regeneration of nerve 772

Trophic nerves - - 778

Classification of nerves 780

CONTENTS- xvii

Chapter XV. Electro-physiology

PAGE

Currents of rest and action - - 796

Relation between action current and functional activity • - 797

Polarization of muscle and nerve - - 802

Electrotonic currents 803

Heart-currents 806

Human electro-cardiogram 808

Glandular currents 810

Eye-currents 811

Electric fishes 812

Section VIII. — Functions of the Central Nervous System — THE Spinal Cord - 859

Decussation of the sensory paths - - 866

Reflex action - 869

Principle of the common path 871

Role of the receptor in reflex action - 872

Characteristic properties of the reflex arc 874

Irradiation of reflex action 877

Co-ordination of reflexes 880

Influence of the brain on spinal reflexes - - 882

Automatism, of the spinal cord " 888 xviii CONTENTS

PAGE

Section X. — Functions of the Brain - - 902 Functions of the cerebellum - - 904

Equilibration and orientation - - • 9^7

Forced movements - 912

Functions of the cerebral cortex 9^4

Motor areas 918

Histological differentiation of the cortex 924

Sensory areas - - 931

Aphasia - 934

Localization of function. in central nervous system 940

Reaction time - - 947

Section I. — Vision 968 Physical introduction 968

Structure of the eye 974

Chemistry of the refractive media 976

Refraction in the eye - - 977

Accommodation - 980

Functions of the iris - 986

Defects of the eye - - 987

Ophthalmoscope . - . ggi

Skiascopy - - ' 994

Diplopia - - - 997

Steroscopic vision . - . ggg

Visual judgments and illusions - . - 1000

Purkinje's figures - ... 1002

Blind spot - 1004

Rods and cones in vision - - 1005

Talbot's law ... iqio

Colour vision ... ion

Contrast - - . . 1016

Perimetry - - - - -1018

CONTENTS XIX

Section I. {continued) — page

Colour-blindness 3^o^9

Movements of the eyes ^ '^"^^

Chapter XIX. Reproduction

Regeneration of tissues 1074

Reproduction in the higher animals • io75

Menstruation - - ^°7^

Development of the ovum - 1078

Parthenogenesis 1080

Formation of the embryo - 1081

Development of the connections lO^S

Exchange of materials in the placenta - 1085

Metabolism of the embryo - - '^"^^

Parturition - *. ^°93

Milk - - i°95

Cultivation of tissues outside of the body- - io97

Transplantation of tissues - - ^°9^

Parabiosis ^■'"■^

APPENDIX - "°3

PRACTICAL EXERCISES.

CHAPTER I. General reactions of proteins - - - - 7

Colour reactions of proteins - - °

Precipitation reactions of proteins ^

Special reactions of groups of proteins 9

Reactions of derivatives of proteins - 9

Carbo-hydrates - . - Fats ------

Scheme for testing for proteins and carbo-hydrates - 13

10 II

CONTENTS

Chapter II

1. Reaction of blood - 62

2. Specific gravity of blood - 62

3. Coagulation of blood - ' 62

4. Preparation of fibrin-ferment 65 5; Preparation of extracts containing thrombokinasc 65 61 Serum 65 7. Action of serum on artery rings 66 8: Comparison of action of serum and epinephrin on artery rings 66 9. Comparison of action of serum and plasma on artery rings 66

10. Enumeration of the blood-corpuscles - 67

11. Haematocrite 68

12. Electrical conductivity of blood 68

13. Opacity of blood 70

14. Laking of blood 70

15. Haemolysis and agglutination - 71 16! Osmotic resistance of coloured corpuscles 73 17. Blood-pigment - 73

(i) Preparation of haemoglobin crystals 73

(2) Spectroscopic examination of haemoglobin and its

derivatives 74

(3) Guaiacum test for blood 76

(4) Quantitative estimation of haemoglobin 76

(5) Haemin test for blood-pigment 78

Chapter III

1. Microscopic examination of the circulating blood 191

2. Anatomy of the frog's heart - 191

3. Beat of the heart 192

4. Apex of the heart 192

5. Heart tracings - 192

6. Dissection of vagus and cardiac sympathetic in frog 194

7. Stimulation of the vagus in the frog - 196

8. Stimulation of the junction of the sinus and auricles 196

9. Action of muscarine and atropia on the heart 197

10. Stannius' experiment - . - . - igy

11. Stimulation of cardiac sympathetic in frof; 197

12. Action of inorganic salts on heart-muscle 198

13. Action of the mammalian heart 199

14. Perfusion of the isolated mammalian heart 203

15. Action of the valves of the heart -- - 204 i6. Sounds of the heart .. - -. - 205 17. Cardiogram . . - . . 205

CONTENTS xxi

PAGE

i8. Sphygmographic tracings - 206

19. Venous pulse tracing from jugular 207

20. 'Polygraph tracings - 207

21. Plethysmographic tracings - 208

22. Pulse-rate - - 208 23.. Blood-pressure tracing - - 208

24. Estimation of arterial pressure in man 211

25. Influence cf position of the body on blood-pressure' 211

26. Effects of haemorrhage and transfusion on blood-pressure 212

27. Influence of proteoses on blood -pressure 213

28. Effect of suprarenal extract on blood-pressure 214

29. Action of epinephrin on artery rings - - 214

30. Section and stimulation of cervical sympathetic in rabbit 215

31. Determination of the circulation-time 2I5

32. Measurement of the blood-flow in the hands - 218

33. Vaso-motor reflexes 220

Chapter IV

1. Traicing of the respiratory movements in man 293

2. Production of apnoea and periodic breathing in man 294

3. Tracing of the respiratory movements in animals 294

4. Heat dyspnoea - - - - 296

5. Measurement of volume of air inspired and expired 297

6. Cardio-pneumatic movements 297

7. Auscultation of the lungs - 298

8. Measurement of the respiratory pressure 298

9. Estimation of carbon dioxide and water given off by an animal 299

10. Muscular contraction in the absence of free oxygen 300

11. Oxidizing ferments - 300

CHAPTERS VI. AND VII.

1. Contraction of isolated intestines in Ringer's solution 446

2. Effect of serum on the contractions of intestinal segments 447

3. Action of epinephrin on intestinal segments - - - 447

4. Quantitative estimation of ferment action - 447

5. Chemistry and digestive action of saliva ^148

6. Stimulation of the chorda tympani - - - 450

7. Effect of drugs on the secretion of saliva • - 451

8. Digestive action of gastric juice - - 152

9. To obtain chyme and gastric juice - 453

10. Digestive action of pancreatic juice - - 454

11. Chemistry of bile 456

12. Microscopical examination of faeces - - - 457 xxu CONTENTS

PAGE

13. Absorption of fat 457

14. Time required for digestion, and absorption of food sub-

stances - - 458

15. Quantity of cane-sugar inverted and absorbed in a given

16. Auto-digestion of the stomach 459

CHAPTER IX.

1. Specific gravity of urine 508

2. Reaction of urine 508 3. , Chlorides in urine 508

4. Phosphates in urine - 509

5. Sulphates in urine 510

6. Indoxyl in urine 510

8. Ammonia in urine - - 513

9. Total nitrogen in urine - - - 514

11. Kreatinin - 515

12. Hippuric acid 516

13. Proteins in urine 516

14. Sugar in urine - - - 517

15. Pentoses in urine - 520

16. Acetone in urine 521

17. Determination of the freezing-point of urine 521

18. Examination of urine - 523

19. Urinary sediments 523

CHAPTERS X., XL, AND XII.

1. Glycogen - - 689

2. Catheterism - 690

3. Experimental glycosuria - - 690

(i) Injection of sugar into the blood 690

(2) Phlorhizin glycosuria - 691

(3) Alimentary glycosuria - 691

8. Excretion of urea (and total nitrogen) and proteins in food - 693

9. Action of epinephrin - - 6g3 10. Measurement of the heat given off in respiration 694

CONTENTS

CHAPTERS XIII. AND XIV.

1. Difference of make and break induction shocks 780

2. Stimulation by the voltaic current 783

3. Ciliary raotion 784

4. Direct excitability of muscle — curara - 784

5. Graphic record of ' twitch ' - 784

6. Influence of temperature on the muscle-curve . 786

7. Influence of load on the muscle-curve - 786

8. Influence of fatigue on the muscle-curve 786

9. Seat of exhaustion in fatigue of the muscle-nerve preparation 786

10. Influence of veratrine on muscular contraction - 787

1 1 . Measurement of the latent period of muscular contraction 788

12. Summation of stimuli - - 789

13. Superposition of contractions - - 789

14. Composition of tetanus. 789

15. Contraction of smooth muscles - 790

16. Velocity of the nerve-impulse 791

17. Chemistry of muscle - - 792

18. Reaction of muscle in rest, activity, and rigor - 793

CHAPTER XV.

1. Galvani's experiment - - 814

2. Contraction without metals - - - 814

3. Secondary contraction - - 814

4. Demarcation and action currents with capillary electrometer 814

5. Action current of the heart - 816

6. Electrotonus - - 816

7. Paradoxical contraction - 816

8. Alterations in excitability and conductivity produced in nerve

by a voltaic current - 816

9. Formula of contraction 817

10. Formula of contraction for (human) nerves in situ 818

11. Ritter's tetanus - 818

CHAPTER XVI.

1. Section and stimulation of nerve-roots - - 957

2. Reflex action in the ' spinal ' frog - 958

3. Reflex time - - 958 4'. Inhibition of the reflexes 959

5. Spinal cord and muscular tonus - - - 959

6. Spinal cord and tonus of the bloodvessels - - 959

7. Action of strychnine - 959

CONTENTS

8. Mammalian spinal preparation - - 959

9. Reflexes in man ■ - . . . 961

10. Excision of cerebral hemispheres in the frog - •■ 961

11. Excision of cerebral hemispheres in the pigeon 961

12. Stimulation of the motor areas in the dog - • 962

CHAPTER XVIII.

1. Dissection of the eye - - - - 1058

2. Formation of inverted image on the retina 1059

3. Helmholtz's Phakoscope 1059

4. Scheiner's experiment - 1060

5. Kiihne's artificial eye 1061

6. Astigmatism f ophthalmometer) 1062

7. Spherical aberration - 1063

8. Chromatic aberration 1063

9. Measurement of the field of vision - 1063

10. Mapping the blind spot - - 1064

11. The yellow spot - - 1064

12. Ophthalmoscope - - 1065

13. Retinoscopy ' 1066

14. PupUlo-dUator and constrictor fibres - 1067

15. Colour-mixing - - 1068

16. After-images - 1068

17. Retinal fatigue 1068

18. Visual acuity 1068

19. Colour-blindness 1069

20. Talbot's law - 1670

21. Purkinje's figures 1670

22. Relation of pitch and vibration frequency 1070

24. Sympathetic vibration - 1070

25. Galton's whistle - - 1070

26. Cranial conduction of sound 1670

29. Touch and pressure - 1071

30. Temperature sensations 1072

CHAPTER XIX.

Contractions of isolated uterine rings - - *■ - iioi

A MANUAL OF PHYSIOLOGY