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TEXT-BOOK OF PHYSIOLOGY.



CONTRIBUTORS.

J. S. EDKINS, M.B., Lecturer on Practical Physiology in St. Barthol-
omew's Hospital Medical School, London.

ARTHUR GAMGEE, M.D., F.R.S., Emeritus Professor of Physiology
in Owens College, Manchester.

W. H. GASKELL, M.D., LL.D., F.R.S., Lecturer on Physiology in the
University of Cambridge.

FRANCIS GOTCH, B.Sc, F.R.S., Waynflete Professor of Physiology
in the University of Oxford.

ALBERT A. GRAY, M.D., University of Glasgow.

W. D. HALLIBURTON, M.D., F.R.S., Professor of Physiology in King's
College, London.

J. BERRY HAYCRAFT, D.Sc, M.D., Professor of Physiology in
University College, Cai-diff.

LEONARD HILL, M.B., Lecturer on Physiology in the London Hospital
Medical School, London.

F. GOWLAND HOPKINS, B.Sc, M.B., Demonstrator of Chemical
Physiology in Guy's Hospital Medical School, London.

J. N. LANGLEY, D.Sc, F.R.S., Lecturer on Physiology in the University
of Cambridge.

J. G. M'KENDRICK, M.D., LL.D., F.R.S., Professor of Physiology in
the University of Glasgow.

B. MOORE, M.A., Assistant Professor of Physiology in University

College, London.

D. NOEL PATON, M.D., Lecturer on Physiology in the School of

Medicine, Edinburgh.

M. S. PEMBREY, M.D., Lecturer on Physiology in Charing Cross
Hospital Medical School, London.

E. WAYMOUTH REID, M.B., Professor of Physiology in University

College, Dundee.

W. H. R. RIVERS, M.D., Lecturer on Physiological Psychology in the
University of Cambridge and in University College, London.

J. BURDON SANDERSON, M.D., D.C.L., F.R.S., Regius Professor
of Medicine in the University of Oxford.

E. A. SCHAFER, LL.D., F.R.S., Jodrell Professor of Physiology in
University College, London.

C. S. SHERRINGTON, M.D., F.R.S., Holt Professor of Physiology in

University College, Liverpool.

E. H. STARLING, M.D., Joint-Lecturer on Physiology in Guy's
Hospital Medical School, London.



TEXT-BOOK



OP



PHYSIOLOGY



EDITED BY



E. A. SCHAFER, LL.D., F.E.S.,

JODRELL PROFESSOR OF PHYSIOLOGY, UNIVERSITY COLLEGE, LONDON.



VOLUME FIEST.



EDINBURGH & LONDON:
YOUNG J. PENTLAND.

1898.



EDIXBURGII : PRI^'TED FOR TOrXG .T. TEXTLASD, 11 TEVIOT PLACE, ASD
3S WEST SMITHFIELD, LONDON, E.C., BY MORRISON AXD GIBE LIMITED.



(A/l rights rescned. )



PREFACE.



The want of a text-book in the English language to which students
could turn for information beyond that contained in the ordinary
manuals has long been felt by teachers of physiology in this country.
The most extensive of the existing text-books do not aim at giving
the full and precise information nor the references to original
authorities which are required by the advanced student. It has
hitherto been necessary for those who seek such information to consult
original articles — an operation which frequently involves a familiar
acquaintance with foreign languages and an expenditure of time rarely
at the disposal of the student. The present work is not intended
altogether to supersede this consultation of original papers, but will,
it is hoped, reduce the need of it to more reasonable limits, and will,
moreover, by the references to literature which throughout form an
important feature of each article, facilitate such study where it is still
necessary.

A book of this character, from the enormous amount of literary
labour which is involved in its production, and from the progressive
character of the science with which it deals, could hardly be undertaken
by one person. The editor has been fortunate enough to secure the
co-operation of many of the leading physiologists in this country, each
of whom deals with some branch of the subject to which he has given
special attention. Accordingly the reader will find in each article, in
addition to information as to the present state of knowledge as com-
plete as it has been possible to make it, many original observations
upon the matter to which it relates.

The subjects of generation and reproduction have been omitted in
this text-book, because, although strictly speaking appertaining to
physiology, they are studied almost entirely by morphological methods,
and are more conveniently treated in connection with morphology. It
has therefore been decided that it would be better not to swell the bulk'
of these volumes, which have already grown beyond the limits originally
intended, by the introduction of subjects such as these, which possess
an enormous recent literature, and are exhaustively dealt with in
special works accessible to every student. The same remark will apply
to the general physiology of the cell, a branch of biology which has



X PREFACE.

of late attained so great an extent and importance as to necessitate
text-books devoted to itself alone, and which it is usual to study rather
as an introduction to, than as a part of, animal physiology.

Of the two volumes of which it is intended this book shall consist,
the articles in the first volume deal mainly with the chemical constitu-
tion and the chemical processes of the animal body, and with those
physical and chemical phenomena which are connected with the pro-
duction and elaboration of the secretions and other fluids of the
body. The articles in the second volume include the mechanics of the
circulation and respiration, and of special muscular movements ; the
general physiology of muscle and nerve ; the special senses ; and the
functions of the central nervous system.

It is nearly twenty years since the publication in six volumes of the
important " Handbuch der Physiologic," under the editorship of Pro-
fessor L. Hermann. The articles in that book, as in this, were under-
taken by physiologists who were specially conversant with the particular
branches of the science with which they severally dealt ; and since most
of the articles in it are prefaced by short historical introductions,
and interspersed with abundant references to the literature of the
subject, the whole work constitutes a storehouse of information,
which has proved of great value to teachers and investigators. But
the size of the work, and the fact that it is written in the German
language, have limited its utility to students in this country ; moreover,
in the course of the twenty years that have elapsed since its appearance,
rapid progress has been made in every branch of physiology, so that
several of the articles in it have been long out of date. Nevertheless
its publication served both to lay a firm foundation for the exposition of
the science in its modern aspect, and also to clear the ground for all
future publications of a similar character. It has thus been a marked
advantage, in preparing many of the articles for the present book, to have
had the work of Hermann and his coadjutors to refer to ; and although
due acknowledgment is made both of this and of other sources of infor-
mation in the articles themselves, it has seemed right specially to
mention the " Handbuch " in this preface.

University College, London,
Febi'uary 1898.



COKTEKTS OF VOLUME FIRST.



THE CHEMICAL CONSTITUENTS OF THE BODY
AND FOOD.

By W. D. HALLIBURTON.

The Carboliydrates — The Fats — Lecithin — Cholesterin — The Proteids — Decom-
position Products of Proteids — Synthesis of Proteids — Theories of Proteid
Constitution — General Properties and Reactions of Proteids — Classification
of Proteids — -Yegetable Proteids — Poisonous Proteids — Compound Proteids
— The Albuminoids — Inorganic Compounds page 1



THE CHEMISTEY OF THE TISSUES AND OEG-ANS.

By W. D. HALLIBURTON.

Cells and Protoplasm — Liver — Spleen — Thymus — Thyroid — Suprarenals — Pancreas
— Kidnevs — Testis — Muscle— Skeletal Tissues — Nervous Tissues — The Eve —
Milk . ' page 80

THE BLOOD.

By E. a. SCHAFER.

General Properties — Amount — Colour— Specific Gravity — Reaction — Coagulation-
Relative Amounts of Plasma and Corpuscles — Number of Corpuscles — General
Composition of Blood — Composition of Blood Corpuscles — Composition of
Plasma — Proteids of Plasma — Theories of Coagulation — Causes of Coagulation
— Lymph and Allied Fluids page 141

H.F:M0GL0BIN : ITS COMPOUNDS AND THE PEINCIPAL
PEODUCTS OF ITS DECOMPOSITION.

By ARTHUR GAMGEE.

Distribution in the Animal Kingdom — Relations to other Constituents of Red
Corpuscles (Arterin and Phlebin) — Oxyhsemoglobin — Methods of Obtaining
— Composition of — Crystalline Form — Action of Reagents on — Spectrum^ —
Spectrophotometry — Photographic Spectrum — HoemoglolDin — Preparation of —
Colour and Spectrum — Compounds with Gases — Derivatives and Products of
Decomposition *.........., page 185



xii CONTENTS.

A GENERAL ACCOUNT OF THE PEOCESSES OF DIFFUSION,
OSMOSIS, AND FILTPtATION.

By E. WAYMOUTH KEID.
Diffusion — Osmosis — Filtration . . . . . . ." . page 261

THE PRODUCTION AND ABSORPTION OF LYMPH.

By ERNEST H. STARLING.

The Production of Lynipli — The Physical Forces concerned in the j\Iovement of
Lymph — The Absorption of Lymph from the Connective Tissues — On the
Functions of the Lymph in the Nutrition of the Tissues . . . page 285

CHEMISTRY OF THE DIGESTIVE PROCESSES.

By B. MOORE.

Digestive Ferments — Chemical Composition of Digestive Juices — SaliA'a — Gastric
Juice — Pancreatic Juice — Intestinal Juice — Bile — Digestion of Carbohydrates
— -Digestion of Proteids — Absorj)tion of Carbohydrates and Proteids —
Digestion and Absorption of Fats — Bacterial Digestion — Composition of
Fffices page 312

THE SALIVARY GLANDS.

By J. N. LANGLEY.

Anatomical Characters — Histological Characters — Origin and Course of Nerves —
Changes during Secretion — Reflex Secretion — The Dyspnceic Secretion —
Stimulation of the Cranial Nerve — Stimulation of the Sympathetic Nerve —
The Augmented Secretion — Effect of Protracted Stimulation on the Amount
and Percentage Composition of Saliva — Relation of the Rate of Secretion to
the Percentage Composition of Saliva — Some General Characters of Saliva —
Substances secreted in Saliva — Effects of the Cranial and Sympathetic Nerves
upon the Blood Flow — Mutual Effects of the Cranial and Sympathetic Nerves
upon Secretion — Effect of Variations in the Amount and Quality of the Blood
sujjjDlied to a Gland — Relation of Secretion to the Flow of Lymph — The
Secretory Pressure — Reflex Inhibition of Saliva — The Action of Alkaloids —
Formation of Heat — Electrical Changes — Section of Glandular Nerves — The
Paralytic Secretion — Secretion due to Reflex Action of Peripheral Ganglia
— Direct Irritability of Gland Cells — Extirpation of the Glands — Injection
into the Blood of Saliva and of Gland Extracts — General Considerations —
Theories of the Mode of Action of Secretory Nerves . . . page 475

MECHANISM OF SECRETION OF GASTRIC, PANCREATIC,
AND INTESTINAL JUICES.

By J. S. EDKINS.

Histological Ap]!)earances accompanying Secretory Conditions of Stomach — Functions
of the Cells and Regions of the Stomach — Methods of obtaining Gastric Juice —
Influence of the Nervous System on Gastric Secretion — Conditions which
provoke Secretion — Formation of the Ferments of Gastric Juice — Formation
of Rennin — Variations in Gastric Juice during Digestion — Histological Appear-
ances of the Secretory Conditions of the Pancreas — Influence of the Nervous
System upon Pancreatic Secretion — Conditions which provoke tlie Flow of
Pancreatic Juice — Ferments of the Pancreatic Juice and their Antecedents —
Variations in Pancreatic Juice during Digestion — Evidence of Secretion in the
Intestine ' . . page 531



CONTENTS. xiii

MECHANISM OF BILE SECEETIOX.

By D. XOEL PATOX.

Mode of Formation of Bile Constituents — Water — Inorganic Salts — Xucleo-
Proteid — Bile Acids — Bile Pigments — Cliolesterin — Lecithin, etc. — Influence
of A''arious Factors on the Secretion of Bile — Flow of Blood — Food —
Pressure of other Organs — Nerves — Chemical Substances — General Con-
clusions .....'....... page bb^



THE CHEMISTEY OF THE URIXE.

By F. CtOWLAXD HOPKIXS.

Introductory — Quantitative Composition of Urine — Variations in its Auiount and
Specific Gravity — Its Chemical Reaction — The Xitrogenous Constituents :
Total Nitrogen ; Urea ; Ammonia ; Uric Acid ; Xanthin Bases ; Creatinin :
Hippuric Acid ; Amido-Acids — Proteids — The Aromatic Substances — Th&
Carbohydrates — Glycuronic Acid and its Conjugated Compounds — Oxalic Acid
— Acids and Oxy acids of the Fatty Series — Colour of the Urine and the
Chemistry of its Pigments : The Preformed Pigments of Xormal Urine ;
Chromogenic Substances ; The Pigmentation of Pathological Urine — The In-
organic Constituents — General Characteristics of the Organic Urinary Com-
pounds — Comparative Chemistry of the Urine .... page 570



THE MECHANISM OF THE SECEETION OF UEINE.

By ERXEST H. STARLIXG.

Theories of Urinary Secretion — Theor}- of Bowman — Theory of Ludwig — Secretion
of Water — Methods — The Concentration of the Urine — Heidenhain's Criticism
of the Theory of Ludwig — Experiments of Xussbaum — Experiments of Ribbert
— Ex23eriments of Bradford — The Influence of the Xervous System on the:
Secretion of Urine .......... page 639-



THE MECHANISM OF THE SECRETION OF MILK.

By E. a. SCHAFER.

General Considerations — Influence of the Xervous System — Action of Pilocarpine-
and Atropine — Influence of Diet — Place of Formation of the Organic Con-
stituents — Manner in which the Secreted Materials pass out of the Cells —
Mechanism of the Discharge of Milk ...... page 662.



SECEETION AND ABSORPTION BY THE SKIN.

By E. WAYMOUTH EEID.

Chemical Xature of Skin Secretions — The Secretion of Sweat — Electro-Motive-
Phenomeua in Skin Glands — Absorption by the Skin of ]\Ian— Of Lower
Mammals— Of the Frog page 66'J.'



xiv CONTENTS.

CHEMISTRY OF EESPIRATIOK

By M. S. PEMBREY.

Historical — Respiratory Clianges in Air — Methods — Conditions affecting Respiratory
Exchange — Cold- Blooded Animals — Fishes — Warm - Blooded Animals — In-
fluence of External Temperature — Of Muscular Activity— Of Food — Of
Size of Animal — Of Time of Day — Of Age — Resj)iration by Skin
in Amphibia — In Mammals — Effects of Varnishing Skin — Respiration in
Alimentary Canal — Respiration of Fojtus — Of Embryo — The Respiration of
Different Gases— The Respiration of Vitiated Air — Asphyxia — Exchange of
Gases between Blood and Air — Frequency of Respiration in Man — In Animals

Changes in Composition of Air — Effect of Respiration on Blood — Gases of

Blood — Methods— Arterial and Venous Blood — Condition of Gases in Blood —
Causes of Gaseous Exchange between Blood and Air— Exchange of Gases
between Blood and Tissues — Causes of such Exchange . . . page 692

ANIMAL HEAT.

By M. S. PEMBREY.

Thermometry — Warm and Cold Blooded Animals — Temperature of Man and other
Warm-Blooded Animals — Hibernation — Influence of Various Conditions upon
Temperature — Time of Day — Age — Muscular Work — Mental Work — Food —
Sleep — Seasons — Race — Menstruation and Pregnancy — Individual Peculiarities

Temperature of Surroundings — Extreme Heat and Cold — Baths — Drugs —

Temperature of Different Parts of Body — Of Arterial and Venous Blood — Of
the Skin — Regulation of Temperature — Heat Production — Historical — Relation
to Chemical Changes — Specific Heat of Body — Seats of Heat Production-
Measurement of Heat Production — Calorimetry — Respiratory Exchange as
Measure of Heat Production — Heat Production in Cold-Blooded Animals

Regulation of Heat Loss — Influence of Size of Body — Influence of Nervous

System — Development of Power of Regulation — Temperature of Body after
Death page 785

METABOLISM.

By E. a. SCHAFER.

Introductory — Balance of Nutrition — Composition of Foodstuft's — Heat Value of
Foodstuffs — Necessary Amount of Proteid — Special Constituents of Diet —
Their Effect on Metabolism — Gelatin — Carbohydrates — Fats— Inorganic Sub-
stances — Metabolism in Inanition^ — With purely Proteid Diet — Relative Meta-
bolic Activity of Tissues— Nitrogenous Metabolism — In Muscle — In the Liver
Effect of Muscular Activity on Proteid Metabolism — Metabolism of Carbo-
hydrates— Glycogen Formation — Phloridzin Diabetes — Glycogenesis— Puncture
Diabetes — Influence of Pancreas on Carbohydrate Metabolism — Metabolism of
Fat— Source and Formation of Fat — Action of Liver on Metabolism of
Fat page 868

THE INFLUENCE OF THE DUCTLESS GLANDS UPON
METABOLISM— INTERNAL SECRETIONS.

By E. a. SCHAFER.

Introductory— The Thyroid Gland— The Pituitary Body— The Suprarenal Bodies
—Influence of the Spleen on Metabolism . ' page 937



Index of Subjects page 963

Index of Authors page 999



LIST OF ILLUSTRATIONS.



I. PLATES.



[I. — Spectra of lipemoglobin, its compounds and derivatives (modified from

Preyer, " Die Blutkrystalle ").
[I. — Spectra of various colouring matters (from MacMunn, " The Spectroscope

in Medicine ").

1. Solar spectrum, with. Frauenhofer's lines and millimetre scale.

2. Fresh human l)ile.

3. Alcoholic extract of human bile.

4. Diluted solution of human bile treated with hydrochloric acid.

5. Bile treated with nitric acid, and the precipitate dissolved in boiling

absolute alcohol.

6. Pig's bile.

7. Ox or sheep bile.

8. Ox or sheep bile treated with nitric acid, and the precipitate dissolved

in boiling alcohol.

9. Ox bile treated with hydrochloric acid, and the precipitate dissolved in

boiling alcohol.

10. Guinea-jjig's bile.

11. Rabbit's bile.

12. Mouse's bile.

13. Crow's bile.

14. Pettenkofer's test on human bile salts.

15. „ „ pig's bile salts.

16. Band of urobilin in normal human urine.

17. Bands in urine of rheumatic fever ; the urine treated with nitric acid.

18. „ ,, „ with albuminuria ; the urine treated with nitric acid.

19. Urine of same case treated with caustic potash.

20. Bands of hsematin from ovarian cyst.

21. The same treated with a reducing agent.

22. Spectrum of 02, after twenty-four hours.

23. „ ,, lutein, from peritoneal fluid.

24. ,, ,, ,, from serum of dog's blood.



LIST OF ILL USTRA TIOXS.



II. FIGITES IX TEXT.

FIG. PAGE

1. Crystals of ptenvlglncosazone ...... 8

2. „ ., phenylinalto.sazone . . . . . .11

3. Lactose crystals (Frey) . . . . . . .12

4. Crystals of pheiiyllactosazone . . . . . .13

5. Inosite crystals (Trey) . . . . . . .17

6. Chole-sterin crystals (Frey) ....... 23

7. Leucine cry.stals (Kilhne) ....... 28

8. Tyro-sine crystals (Frey) ....... 28

9. Crystals of egg albumin ....... 44

10. Proteid crystals from human urine (BrauiArell and Noel Paton) . . 45

11. Crystallised vitellin of the oat kernel (Osljome) . . . .53

12. Charcot's crystals ........ 94

13. Creatine crystals (Klilme) . . . . . . .100

14. Creatinine crystal? (Kiilme) . . . . . . .100

15. Creatine-zinc cUoride crystals (Kiihne) ..... 101

16. Spherical compound of mercury and creatine (G. S. Johnson) . . 102

17. Compounds of xanthine and hypoxanthine, by means of which these sub-

stances may be isolated and identified (Kiihne) .... 103

18. Zinc sarcolactate (Kiihne) . . . . . . .107

19. Calcium sarcolactate (Kiihne) . . . . . .107

20. Absorption spectra of retinal pigments (Kiihne) . . . .123

21. Olirers apparatus for estimating the numlier of blood corpuscles . . 149
a. ,, hsemoglobinometer . . . . . . .151

23. The hjematinometer . . . . . . . .211

24. The hsemato.?cope .. . . . . . .211

25. Graphic representation of the .spectrum of oxyhemoglobin and htemo-

globin (KoUett) ........ 212

26. Double slit employed in Tierordt's method of spectrophotometry . 216

27. Glass troughs for containing the liquids to be examined by the methods

of spectrophotometry (Krii,s.s) . . . . . .217

28. Trough mounted on stand, as used in spectrophotometry (Krii.?.s) . . 217

29. Section of glass trough (Kriiss) . - . . . . 217

30. Spectrophotometer with absorfition trough and lamp . . . 218

31. Hiifner's spectrophotometer ....... 221

32. Schematic representation of the path followed by the rays of light before

entering the slit of the collimator of Hiifners spectrophotometer
(Kriis.s) 222

33. The photographic spectrum of liEemoglobin and oxy haemoglobin . . 227

34. Graj»hic representation of the spectra of oxyhajmogloblin and haemoglobin

(Ptollett) 233

35. The photographic spectrum of oxyhaemoglobin and of CO-haemoglobin . 240

36. .. „ „ oxyhaemoglobin and methaemoglobin . 247

37. .. „ ,, haemin ..... 254

38. ,, „ ,, oxygenised h«mochromogen and of haemo-
chromogen ........ 256

39. The photographic spectrum of haematoporphyrin .... 259

40. Diagram to show the dilution of the blood produced in dogs by the

injection of dextrose (Leathe.?) ...... 294

41. Diagram to show the influence of the intravenous injection of dextro.se

on the blood pressure in the abdominal viscera .... 294

42. Diagram to show the effect of injecting dextrose after a previous

bleeding ......... 295



LIST OF ILLUSTRATIONS.



FIG.

43.

44.
45.

46.
47.
48.
49.
50.
51.
52.
53.
54.



Diagram to show effects of the injection of a lymphagogue of the first

class on the blood pressures in the abdominal organs
Diagram to indicate variations in pepsin after food (Griitzner)
Chart showing acidity of gastric juice after feeding with mixed

food ........

Chart of the course of secretion of pancreatic juice

Chart of the percentage composition of the flow of pancreatic juice

Showing influence of various foodstuft's upon the secretion of bile

Urea nitrate and oxalate crystals

Uric acid crystals



Ammonium and sodium urate
Creatinin and hippuric acid
Leucine and tyrosine

55. Cystine

56. Calcium oxalate

57. Chart of spectra of urinary pigments ....

58. Stellar phosphates ; triple phosphates ....

59. Eoy's oncometer .......

60. Diagrammatic section through Eoy's oncometer

61. Eoy's oncograph .......

62. Eegnault and Eeiset's respiration apparatus

63. Toit's respiration apparatus .....

64. The respiration apparatus in the Physiological Laboratory, Oxford

65. Haldane's respiration apparatus .....

66. Lowy's „ „ .....

67. Frederick's curve of daily variation in the absorption of oxygen .

68. Hutchinson's spirometer ......

69. Pfliiger's blood-pump ....-•

70. Leonard Hill's blood-pump ....-•

71. Pfliiger's lung catheter ...•••

72. Curves of dissociation of oxyhsemoglobin , . . .

73. Pfliiger's aerotonometer ......

74. Frederick's ,, ......

75. Bohr's htemataerometer ....••

76. Chart showing daily variation in temperature observed by Einger and

Stewart ......••

77. Chart showing daily variations in temperature observed by Ogle

Clifford Allbutt, Casey and Eattray, and Crombie

78. Chart showing daily variation in temperature observed by Jiirgensen and

Liebermeister .....••

79. Curve of daily variation in the temperature of the urine .

80. Chart sho^-ing daily variations in temperature observed during U, Mosso'

experiments . . . • • • •

81. Diagram of ice calorimeter ....••

82. Diagram of Dulong's water calorimeter ....

83. Diagram of air calorimeter (Haldane, Hale White, and Washbourn)

84. Monkey deprived of thyroid (Horsley) ....

85. Eftect upon the blood pressure in the dog of the intravenous injection

of decoction of thyroid ....••

86. Tracing showing effect of pituitary extract upon heart-beats and blood

pressure in the dog ......

87. Tracing showing effect of suprarenal extract upon muscle contraction in

the frog . . . . . .

I



297

544

545
553
553
566

582

587

589

591

600

602

603

615

624

633

641

642

643

694

696

697

697

698

721

752

758

759

774

775

776

777

777

800

800

800
801

802
844
845
845
941

944

947

951



LIST OF ILLUSTRATIONS.



88. Tracing sliowing effect of suprarenal extract upon lieart-lDeats and blood



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