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A TEXT BOOK OF PHYSIOLOGY.



in,iiinnrU!\TfO(.ocv



A TEXT BOOK



OF



PHYSIOLOGY



BY



JOHN GEAY M'KENDEICK, M.D., LL.D., F.E.S.

PROFESSOR OF THE INSTITUTES OF MEDICINE IN THE UNIVERSITY OF GLASGOW,
FELLOW OF THE ROYAL COLLEGE OF PHYSICIANS OF EDINBURGH.



INCLUDING

HISTOLOGY BY PHILIPP STOHE, M.D.,

OF THE UNIVERSITY OF WiJRTZBURG.



IN TWO VOLUMES.

VOL. I.— OENEEAL PHYSIOLOGY.



NEW YOEK:
MACMILLAN AND CO.

1888.






I/. I



^



B'C D Eh F G '




MoLc Muivn, deL acL Nax,



Ifanfwrb UJJv.



GENERAL PHYSIOLOGY:



INCLUDING



THE CHEMISTRY AND HISTOLOGY OF THE TISSUES
AND THE PHYSIOLOGY OF MUSCLE.



BY



JOHN GEAY M'KENDPtICK, M.D., LL.D., F.R.S.,

PROFESSOR OF THE INSTITUTES OF MEDICINE IN TUE UNIVERSITY OF GLASGOW,
FELLOW OF THE ROYAL COLLEGE OF PHYSICIANS OF EDINBURGH.



NEW YORK:
MACMILLAN AND CO.

1888.



V. t



PRINTED AT THE U^fIVERS^TY PRESS, QLASOOW,
BV ROBERT MACLEHOSE.



VIRO • ADMIRABILI

ACERRIMO . IN . NATVRA • INVESTIGANDA

IN . INTERPRETANDA • SAGACISSIMO

lOANNI • BVRDON ■ SANDERSON

QVI . LARGIS • INGENII ■ LVMINIBVS

QVODCVNQVE • ATTIGIT ■ ILLVSTRAVIT

SANO . IDEM . IVDICIO • ATQVE ■ AEQVO

SOLIDA • SEMPER • VANIS • PRAETVLIT ■ VTILIBVS • HONESTA

ATQVE . OMNIA . POSTHABVIT • VERITATI

HOC • MVNVS • QVALECVNQVE

D. D.
lOANNES • G • M'KENDRICK



PREFACE,



Although this work has been modelled to some extent on my " Out-
lines of Physiology," now out of print, it is essentially a new book, and
as it aims at giving a more detailed account of the subject, the title of
"Outlines of Physiology" has been abandoned for that of a "Text
Book of Physiology." I have not endeavoured to give anything like an
encyclopaedic account of the modern researches that have built up the
physiological science of the present time, but rather to weave into a con-
secutive narrative the main facts and principles of physiology, as these
present themselves to my mind, after nearly eighteen years of experi-
ence as a teacher of the science. Probably no science is advancing with
greater rapidity, if one may form an opinion from the large number of
original papers that are announced, month by month, and from the large
number of investigators in the physiological laboratories of the Con-
tinent, of America, and of our own country. Thus it is year by year
becoming more difficult for a teacher to keep abreast of the wave of
progress, and still more difficult to assimilate the new facts and to
incorporate them with his previous knowledge. Each teacher will also
view the subject more especially according to his predilections for the
morphological, the chemical, or the physical side of the science, and
it is not easy in the preparation of such a work as this to be im-
partial and to present the science in all its completeness to the
reader.

An explanation may be expected as to my object in introducing
another large work on physiology when in recent years several excellent
and voluminous treatises have issued from the press and have met with



X PREFACE.

great acceptance. My chief reason is to have it in my power to place
in the hands of my students a text book appropriate to the course of
instruction in physiology, which it is my duty to give annually in the
University of Glasgow. The present work contains most of the
theoretical teaching given to the student, and I hope that with it in his
hands, it will be possible for me to do more and more in the way of
demonstration and of practical instruction. At the same time, I am not
without the hope that the nature of the liook may be such as to com-
mend it to a wider circle of readers, more especially to members of the
medical profession who desire to become acquainted not only with the
facts but also with the methods of physiology.

The general plan of the book is to give an account both of methods
and of results, and to introduce illustrations as far as possible |from the
best sources. No expense has been spared to obtain good illustrations,
not merely diagrams, but illustrations that will give the reader a fair
idea of the appearance of the thing represented. The reader is recom-
mended carefully to peruse the descriptions of the various figures,
as, in these, information is often given which is not contained in the
text.

The work is primarily divided into two parts. First, that relating to
the general physiology of the tissues, which forms the subject of the
first volume ; and, second, that relating to the special physiology of
organs, to be discussed in the second volume. This arrangement is
simple and comprehensive. After an introductory section dealing with
general notions as to living matter, more especially with reference to
the great doctrines regarding energy, which form the basis of modern
science, I proceed to discuss the nature and properties of the chemical
.substances found in the body and the nature of the chemical reactions
with which the phenomena of life are associated. In this section I have
introduced a chapter explaining to the physiological student the views
he should hold as to the true value of chemical formulae. This chapter,
in the preparation of Avliich I received valuable assistance from Professor
W. Dittmar, F.E..S., I introduced after much consideration, and because
I know that in the minds of most students there is not a little confusion
regarding this matter. No doubt such a chapter would be more
appropriate in a strictly chemical work, but my experience as a teacher



PREFACE. xi

assures me that it will not be without its value even in a physiological
text book, did it serve no other purpose than to shoAV how little we yet
know of the molecular structure of organic chemical substances. In
the preparation of this section I derived much assistance from Dr.
Arthur G-amgee's Physiological Chemistry and from Beaunis' Physiologie
Bumaine.

I venture to direct the attention of the reader to the chapter on pig-
ments in which these interesting substances are more fully discussed
than has yet been attempted in any text book of physiology. The
writings of Dr. C. A. MacMunn of Wolverhampton have largely assisted
me in writing this chapter. Dr. MacMunn, as is well known, has made
this subject his own special field of research, and from his stores of
knowledge he gave me the valuable measurements of wave-lengths.
He also kindly prepared the chart of spectra which forms the frontis-
piece of this volume, and he added further to my indebtedness by read-
ing the proof sheets of the chapter on pigments and by furnishing me
with, many valuable suggestions. I consider myself fortunate in having
secured Dr. MacMunn's co-operation, and I offer him my most cordial
thanks.

The next section deals with the physiology of the tissues. Here I
was met by a great difiiculty. In my previous work scarcely any
description of the microscopical structure of the tissues or organs, or
Avhat is termed histology, was introduced, and the omission was
regarded by critics as diminishing the value of the book. My own
feeling was in favour of omitting the discussion of histology from a
physiological text book, but after consulting several physiologists, in
whose judgment I place great confidence, and after taking into account
the wishes of my students, I found that the balance of opinion was in
favour of introducing histology. At the same time, I am bound to admit
that several physiologists gave good reasons for omitting histology.
Having resolved to introduce this aspect of physiological science, the
next difiiculty was whether I was prepared to face the trouble and cost
of preparing a new set of histological woodcuts. After some consider-
ation, I solved the problem by purchasing from Mr. Gustav Fischer, the
well-known publisher in Jena, a set of electrotypes of the woodcuts in
Professor Philipp Stohr's Lehrbuch der Histologie, published last year, and
also the right of translating the work, and of ineori:)orating it with my



3fii PREFACE.

book. This arrangement was effected Avith the cordial consent of the
author, Dr. Stohr, whose name I therefore place on the title page. Dr.
Stohr's work commended itself to me on account of the excellence of its
descriptions, the fidelity of its illustrations, — which are not diagrams but
drawings of real preparations, — and the value of its practical directions
I have therefore made free iise of it, introducing special descriptions and
illustrations, where these seemed to be needful, and I have done this the
more readily as the directions given l)y Dr. Stcihr are substantially those
followed from year to year in the practical classes held during the
summer session in the University of Glasgow. Having Dr. Stohr's
work at my command has also led me to introduce more in the way of
practical directions as to histological methods than I might otherwise
have done, so that the present volume will be a guide to the student
during the practical work of summer, and at the same time serve as a
systematic text book. One unique feature of Dr. Stohr's work is that
he gives an account of the method by Avhich each preparation figured
was prepared. These descriptions of methods I have relegated to an
appendix and numbered consecutively so as to admit of ready
reference.

Before taking up the physiology of the tissues, I have discussed their
origin in the light of recent investigations regarding the phenomena of
fecundation and the minute structure of cells and of nuclei. This is a
novel arrangement, so far as recent systematic works are concerned, but
it has commended itself to me as one leading to a philosophical view of
the whole subject. The cpestions as to the origin of the tissues and the
mysterious Avay by which they are impressed with hereditary characters
are of profound importance and have a bearing not only on physiological
but on pathological theories. The portion dealing A^ath the theories of
heredity has been read by Professor E. Eay Lankester, F.R.S., who, Avhile
not agreeing with me in m}' general conclusion, has giA^en me A^aluable
suggestions and criticisms. The Avhole of this section has also been read
by Mr. J. H. Fiillarton, M.A., B.Sc.

Section III. deals Avith the contractile tissues, the studj' of Avhich
requires physical appliances. I still hold that AAdthout burdening a
student by requiring of him a knoAA'ledge of complicated apparatus, it
is possible to giA^e him, shortly, such information regarding methods as
AAdll enable him to take an intelligent view of results. Instead, hoAvever,



PREFACE. xiii

of introducing a description of apparatus in the parts describing results,
I have discussed apparatus and methods as far as possible in separate
chapters, so that the reader may pass over these if he chooses. The
importance of the uses of electricity in practical medicine and surgery
appears to me to justify the account given of electrical apparatus, for
assistance in the preparation of which I am indebted to Mr. Thomas
Gray, of the Physical Laboratory of this University, and an accomplished
electrician ; and the immense value of the graphic method in all sciences
dealing with movements also warranted me in giving a brief account of
its chief instruments. As to the introduction of physical questions into
a text book of physiology, a good deal can be said on both sides. No
doubt if students of medicine and practitioners read good text books on
general physics, or the excellent works on physiological physics that
have ajDpeared in recent years, there would be little or no necessity for
introducing these matters into a text book of phj^siology, except in so far
as all physiological questions ultimately resolve themselves into physical
problems. But the fact is that the teacher has usually to deal with
students who know little or nothing of physics. The examination in
mechanics required for the registration of a medical student is of no use,
but is just sufficient to worry him and exhaust his energies without con-
ferring any real benefit in the shape of a knowledge of the principles of
physical science. Had he a course of instruction in general physics before
beginning the study of medicine, matters would be on a different footing,
but that is not at present available. Consequently the teacher need not
lecture on the appearance of a muscle as seen with polarized light without,
in the first instance, stating the more elementary facts regarding polarized
light, and without explaining the construction of a polarizing apparatus.
Again, before beginning the discussion of muscle in which the physio-
logical teacher must make use of electricity, he is obliged to give some
explanation of the nature of voltaic cells, of induction coils, and of the
general facts regarding resistance, and of the detection and measurement
of currents. For these reasons I have introduced certain details as to
physics just as I am obliged to introduce them in my lectures. At the
same time, I admit that this is so far a provisional arrangement, and if
the time arrives when the student, beginning the study of physiology,
has already been well grounded in even elementary physics, it may be
abandoned. I have in the meantime chosen the mode of teaching



xiv PREFACE.

which experience has shown mc to lie the most useful under the
circumstances.

The chapter on the electrical fishes is longer than might be exi)ectcd
in a general treatise on jihysiology, but I have thought it right to enter
into details regarding these remarkable animals, as their physiology has
an important bearing on questions as to the functions of nerve and
muscle. I am much indebted to Professor Burdon-Sanderson for aiding
me with a description and with drawings of the electric organ of the
skate, in the investigation of which he and Mr. Francis Gotch, M.A.,
have been recently engaged and the results of Avhich have not been
published. I have also to thank Mr. Gotch for looking over the proof
sheets of this chapter and for valuable suggestions.

In the preparation of the entire volume, I have gratefully to acknow-
ledge the valuable services of my assistants. Dr. J. M'Gregor-Robertson,
M. A., and of Dr. William Snodgrass, M.A. Dr. Snodgrass has read all
the proof sheets twice over, and Dr. M'Gregot'-Robertson has read the
final revise. Both have given me great help and excellent suggestions.
Mr. J. T. Bottomley, M.A., F.E.S., and Mr. Magnus Maclean, M.A., have
also aided me in the discussion of various physical questions.

I have also to thank Professor Rutherford, F.R.S., Professor Marey,
the Cambridge Scientific Instrument Company, Messrs. Churchill,
Blackie & Son, A. & C. Black, Cassell & Co., Collins <fc Sons,
Yieweg & Son of Brunswick, Fischer of Jena, Hahn of Hanover, Carl
Ricker of St. Petersburg, Vogel of Leipsig, and Masson of Paris, who
have favoured me -with electrotypes of illustrations for this work. The
soiu:'ces of all the illustrations are given in the list of woodcuts.

I am indebted for the index to my pupil, Mr. Alexander Far-
quharson.

The second volume is ready for the press and will be published with-
out delay.

With these introductory remarks, I submit the work to the reader,
in the hope that he may find it a guide to the study of physiology and
possibly a stimulus to further research.

JOHN G. M'KENDRICK.

University of Glasgow,
May, 188S.



CONTENTS.



SECTION I.— GENERAL INTRODUCTION.

PAGE

Chap. I. — Nature and Objects of Physiology, . . . . 1

Chap. II. — Matter and Energy, ...... 4

Matter ; Measurements; Eneryy.
Chap. III. — General Principles of Biology, . . . . .1.3

Physical Structure ; Chemical Composition ; Organic Form and Mode of .
Growth; Dynamical Characters; Evolutional History of Living
Beings; Theories of Life.

SECTION II.— THE CHEMISTRY OF THE BODY.

Chap. I. — The Inorganic Constituents, ..... 34

Water ; Mineral Matters,
Chap. II. — The Chemical Constitution of the Organic Constituents, . 43

Classification of Organic Compounds.
Chap. III.— The Proteids or Albuminoids, . . . . .57

Chemical Characters ; Physical Characters ; Polarized Light and
Polariscopes ; Physiological Characters.
Chap. IV. — The Characters of the Special Proteids, . . , . 72

Tr^ie Alhumiiis ; Albuminous Derivatives.
Chap. V. — The Nitrogenous Proximate Principles other than the Proteids, 81

Fatty Nitroge7ious Principles ; the Amides,
Chap. VI. — The Nitrogenous Proximate Principles, . . .89

The Amides, or Amido-Acidn.
Chap. VII. — The Nitrogenous Acids, ..... 97

Bile Acids.
CH.iP. VIII. — The Nitrogenous Bodies containing no Oxygen, . . 109

Chap. IX.— The Pigments, . . . . . . .111

Principles of Spectroscopy ; Pigment of the Blood and its Derivatives;
Pigments of the Bile ; Pigments of the Urine ; Pigments of the Faeces ;
Pigments of the Tissues ; Luteins or Lipochromes ; Chromophanes ;
Black Pigment; other Animal Pigments; Conclusions regarding the
Pigments,



xvi CONTENTS.



PAGK



Chap. X. — The Non-Nitrogenous Matters, ..... 146
The Alcohols; the Fats; the Carbohydrates; the Non-Nitrogenous
Orrfanic Acids.

Chap. XL— The Gases, ....... 168

Chap. XII. — The Chemical Keactious in the Living Organism, . .170

Oxidations ; Reduction ; Decom'position ; Synthesis ; Fermentation.

Chap. XIII. — Fermentation, . . . . . . .175

The Soluble Ferments ; the Orrjanized Ferments ; Chemical Classification
of Ferments ; Nature of. the Organized Ferments ; Modes ofCultivatini/
Schizomycetes.



SECTION III.— THE PHYSIOLOGY OF THE TISSUES.

Chap. I. — Historical Introduction, . . . . . . . 201

Chap. II.— The Origin of the Tissues, . . . . . .214

Spermatozoa; Ova; Influence of Spermatozoid on Ovum ; Polar Bodies ;
Fecundation.

Chap. III. — Theories as to the Physiological Basis of Heredity, . . 234

Chap. IV. — Formation of the Blastodermic Layers, .... 244

Chap. V. — Tlie Microscope and the Methods of Microscopical Research, . 252

The Microscojye ; Accessory Appliances; Reagents; Preparation of
Tissues and Organs.

Chap. VI. — Microtomes and Serial Section Cutting, . . . .• 278

Chap. VIL— Protoplasm and Cells, . . . . . .288

Modes of Studying Protoplasm; Cells; Size, Nutrition, Movements,
Formation., Secretion, Growth, and Evolution of Cells.

Chap. VIIL— Structure of the Varieties of Cells and of Cellular Tissue, . 299
Leucocytes ; Coloured Blood Cells ; Ejnthelial Cells : Connective Tissue
Cells ; Fat Cells ; Aluscle Cells ; Examination of Muscle by Polarized
Light; Nerve Oells.

Chap. IX. — The Physiological Properties of Epithelium, . . . 317

Ciliary Motion.

Chap. X. — The Intercellular Substance, ..... 322

White Fibrous Tissues.

Chap. XL — The Structure of the Connective Tissues, . . . 324

The Connective Tissues Proper ; Cartilage; Chemical Composition of
Cartilage ; Bone; Microscopical Structure of Bone ; Marrow ; Develop-
ment of Bone or Ossification ; Chemical Composition of Bone.

Chap. XII. — The Physical and Vital Properties of the Connective Tissues, 343
Specific Weight; Consistence; Cohesion; Elasticity; Vital Properties.

Chap. XIII. — Phenomena of Filtration and of Osmosis in Relation to

Tissues, ........ 347



CONTENTS.



xvu



SECTION lY.— THE CONTEACTILE TISSUES.

PAGE

Chap. I. — The Special Structure of Muscles and their relations to Xerves, 356

Motor End- Plates.
Chap. II. — The Chemical Composition of Muscle, .... 360
Chap. III. — Electrical Apparatus employed in the Study of Muscle, . 363

General Statement and Definitions ; Voltaic Elements ; Induction Coils ;
Accessory Aijpliances ; Key; Metronomic; Kro7iecher\s Contact -
Brealcer ; PohVs Commutator ; Polarizable Electrodes ; Non-Polariz-
able Electrodes.

Chap. IV.— The Graphic Method, . . . . . .381

Chronognxphy ; Revolving Cylinder ; Direct recording of Movement;
Transmissio7i of Movement.
Chap. V. — The Physical Properties of Muscle, .... 397

Consistence ; Coliesion ; Elasticity.
Chap. VI. — Muscular Irritability, . . . . . . 400

Chap. VII. — General Phenomena of Muscular Contraction, . . 405

Phases of a Single Contraction or Twitch ; Propagation of a Wave of
Contraction.
Chap. VIII. — The Genesis of Tetanus, . . . . .414

Chap. IX. — Modes of Exciting Muscular Contraction, . . .418

Electrical Stimuli ; Mechanical Stimuli ; Thermal Stimuli ; Chemical
Stinmli.
Chap. X.— The Production of Heat by Muscle, . . . .421

Chap. XI.— The Work done by Muscle, . .424

Chap. XII.— The Muscle Sound, . . . . . . 427

Chap. XIII. — -The Phenomena of Muscular Fatigue, . . . 428

Chap. XIV.— The Nutritive Changes or Metabolism in Muscle, . 430

Chap. XV. — The Phenomena of Cadaveric Rigidity, . . . 432

Chap. XVI. —The Growth and Atrophy of Muscle, .... 433

Chap. XVII. — The Properties of Non-Striated Muscular Fibre, . . 435

Chap. XVIIL— The Electrical Phenomena of Muscle, . . .436

Chap. XIX. — Summary of the Phenomena of a Living Muscle, . . 460

Chap. XX. — The Phenomena of the Electric Fishes, . . .461

Appendix I. — Methods of Histological Eesearch, .... 487

Appendix II. — Chemistry of Muscle, ..... 501

Appendix III. — Latent Period of Muscle, ..... 503

Index, .......... 505

Explanation of Plate of Spectra,' page 144.



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75. Stages oE KazyakinBas is Nwfeas •£ CdUi of

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