y \AQA f\ ^^- '-

— From an engraving-.
WILLIAM HARVEY, 1578-1657.

^

PATHFINDERS OF
PHYSIOLOGY

— BY—

J. H. DEMPSTER, A. B., M. D.

EDITOR OF THE DETROIT MEDICAL JOURNAL; FORMERLY LECTURER IN

PHYSIOLOGY, DETROIT COLLEGE OF MEDICINE; MEMBER OF THE

DETROIT MEDICAL CLUB; WAYNE COUNTY MEDICAL SOCIETY;

MICHIGAN STATE AND AMERICAN MEDICAL ASSOCIATIONS

Science

" * * * / taught them how the stars do rise
And set in mystery, and devised for them
Number, the inducer of philosophy,
The synthesis of letters, and besides
The artificer of all things. Memory
That sweet Muse-Mother. "

'"Aeschylus

PUBLISHED BY THE
THE DETROIT MEDICAL JOURNAL COMPANY
19 14

QP

Ij i Published by

The Detroit Medical Journal Company,
1914

FOREWORD

The following pages are the result of the writer's indulgence in
biography as a recreation. The title Pathfinders is presumed to
describe the contents. The biographical essay, it is hoped, will be the
tribute of a stone to the cairn of those who have blazed the trail of
discovery in a domain that has meant so much to scientific medicine,
for ''Destiny reserves for man repose enough." The writer of bio-
graphy, in his self-appointed task fills a role similar to that of Old
Mortality in Scott's well-known novel who visited the graves
of the departed and renewed the moss-covered inscriptions on
their gravestones. The chapters which constitute this volume
have already appeared in the Detroit Medical Journal and are
reprinted here with slight alteration. There is no pretense towards
a complete history of physiology; far from it. Hence, while the
courteous reader will give the writer credit for having read, the criti-
cal reader will discover, perhaps, a great deal that he has either over-
looked or failed to read. The subject itself abounds with interest;
regarding the manner of its presentation, perhaps, not so much may
be said. An endeavor has been made to present as much of the hu-
man element as available data has permitted. Though the real life
of every great man lies in the story of his achievement, rather than
in the tale of how he passed his days, yet the human touches find re-
sponse in the mind of man. "I have remarked," said Carlyle, "that a
true delineation of the smallest man and his scene of pilgrimmage
through life is capable of interesting the greatest man; each man's
life is a strange emblem of every man's and human portraits faith-
fully drawn are of all pictures the welcomest on human walls."

The reader of the history of medicine cannot but be impressed by
the cosmopolitan nature of the science. National lines are unknown,
for thoughtful men of every clime have contributed to its progress.
Its beginnings are enveloped in the mazes of ancient superstition,
where here and there its fitful light gleamed forth to be succeeded
by long centuries of Cimmerian darkness. Owing to veneration for
the work of such men as Galen, to the sacredness with which the

lifeless body was viewed and to the slow development of its ancillary
sciences the progress of medicine up to the beginning of the nine-
teenth century was necessarily slow. Medicine, on the whole, how-
ever, has advanced during periods of great intellectual activity and
during times of intellectual torpor has remained in a quiescent
state. The rise and fall of systems and methods would dispose
one to wonder if the end is yet; if we have at last reached
the bedrock of fact in a scientific sense. The great advantage
of truth over error is that though at times crushed to earth,
it will rise again. Not until science and philosophy had freed them-
selves from the throes of ecclesiasticism, was any marked forward
movement possible, for, during the first fifteen centuries of the Chris-
tian era the most preposterous ideas of physiology obtained, being
founded upon the sacred writings and superstitions of the saints.
The growth of knowledge through observation was scarcely possible
until the priest was no longer physician. With this great event is
associated the name of Hippocrates who was the first to make deduc-
tions based upon experiment and observation. He lived during the
Golden Age when Pericles ruled with mild persuasion; when Phidias
made immortal the sculptured art of Greece and Herodotus recorded
the history of the illustrious people ; when Democritus proclaimed
the atomic theory of the universe and Socrates taught that the
greatest knowledge was to "Know thyself." Experiment, observation
and deduction have been aptly called the tripod of science. Though
much that Hippocrates taught has been discarded, yet in the field of
clinical observation many of his teachings prevail today. The "facies
Hippocrates" still designates the characteristic signs of impending
death. We have many accurate descriptions of disease made from
careful observations, but perhaps more than all else we owe to him
that lofty idealistic note which comes down to us in the Hippocratic
oath.

It was not until men disregarded authority and made direct appeal
to nature that medicine experienced its renaissance. Such was the
method of Harvey, Beaumont and of others whose contributions are of
permanent value. The sincere student of nature approaches his subject
with an open mind ; his is the quest for truth. He possesses "that en-
thusiasm for truth, that fanaticism for veracity, which is a greater
possession than much learning; a nobler gift than the power of in-
creasing knowledge." As Sir Michael Poster once said, "His nature
must be one which vibrates in unison with that of which he is in
search ; the seeker after truth must himself be truthful, truthful with
the truthfulness of nature, which is far more imperious, far more ex-
acting than that which man sometimes calls truthfulness." Such is
the religio medici.

Nor is the history of medicine without its martyrs. While scien-
tific inquiry has been the chief instrument in producing a higher and
better civilization, it has met at almost every step determined op-
position from the powers of ignorance and jealousy. There is great
satisfaction in giving to the world those things which all men see and
for which all men are grateful. The poet, the painter, the musician
and the architect vie with one another in their appeal to the esthetic
sense. Yet is there not something higher even than knowledge for
the sake of knowledge, or art for art's sake? Yes, there is honor
to him who chooses a less spectacular calling, to him who applies
scientific knowledge to the conquest of disease. Such men have bat-
tled with the enemy unencouraged by the blare of trumpets or the
throb of the war drum. They have pursued their work in hospital
ward or laboratory, or as "Weelum McLure," have braved the winter
storm on errands of mercy to the suffering.

"Speak History! Who are life's victors? Unroll thy long

annals and say;
Are they those whom the world calls victors who won the

success of the day?
The martyrs or Nero? The Spartans who fell at Ther-
mopylae's tryst,
Or the Persians and Xerxes? His Judges, or Socrates?
Pilate or Christ?"

J. H. D.

Among the works by which the writer has been assisted and to which his
grateful acknowledgments are due are the following: William Harvey, by D'Arcy
Power; Biology and it's Makers, by Locy; Lectures on the History of Physiology
and Claude Bernard, by Sir Michael Foster; Harvey's Work on the Circulation,
Sydenham Society Edition; Beaumont's Work on Digestion (original copy); Life
and Letters of William Beaumont, by Myer; Brain and Personality, by Thomp-
son; Recent Progress of Heredity, Variation and Evolution, by Locke; Heredity,
by Thompson; Gorton's History of Medicine; The Relation of Medicine to Philos-
ophy, Moon. — Alabama Student, by Osier.

CONTENTS

Frontispiece William Harvey, Portrait

CHAPTER I.

The Circulation of the Blood — William Harvey 1

The Renaissance — Anatomy and Physiology, Galen (1) — Vesalius (2) —
Harvey, birth and education (3) — Fabricius and Harvey, friends; Ana-
tomical Teaching previous to 1745 (4) — Harvey's personal characteristics
(5) — Harvey as Lecturer; His Lecture Precepts (7) — Harvey and Bacon;
Publication of the Work on the Circulation (8) — The Treatise on the Cir-
culation (10) — Capillary Circulation (12) — Asellius and the Lymphatic
Circulation (13) — Asellius opposed by Harvey (14) — The Lacteals, dem-
onstrated by Johannes Pecquet (15).

. CHAPTER II.

Physiology of Digestion in the Seventeenth and

A

^ Eighteenth Centuries 16

Stahl and Boerhaave attacked the Chemical Problems of Physiology (16)
— Peyer and Brunner; Mechanical and Chemical Views of Digestion (17)
— Borelli and Sylvius (18) — ^Haller's Elementa Physiologia appeared
1757 (19) — Reaumur and His Methods; Experiments with Gastric Juice;
Spallanzani (21) — Work of Reaumur and Spallanzani confirmed by Stevens
of Edinburgh (22). ^

V

CHAPTER III.

Physiology of Digestion — William Beaumont 23

The Investigations of Beaumont on the gastric juice of St. Martin freely
quoted in Medical Literature; Beaumont, His Early Life (23) — The Rou-
tine of a Medical Apprentice at the beginning of last century (24) — As-
sistant Army Surgeon (25) — Beaumont's Diary (25) — St. Martin's Acci-
dent (26) — Beaumont conceives the idea of experimenting on St. Martin
(27) — Beaumont Honored by the Michigan Medical Society (28) — Seeks
Assistance of two Leading Scientists (29) — St. Martin Attains Fame
Through His Stomach (30) — Beaumont resigns from the Army (31) —
His Death (31)— His -Work (32).

'\

CHAPTER IV.

Glycogenic Function of the Liver — Vaso Motor Nerves
— Claude Bernard 35

Bernard's Early Life and Education (35) — His Productive Period (36) —
Work on Gastric Digestion (37) — Glycogenic Function of the Liver (37)
— Vaso-Motor Nerves (39) — The Action of Carbon Monoxide Gas (39) —
A Friend of Pasteur (40) — Domestic Troubles (40) — Made One of the "Im-
mortals" (40) — His Dexterity as Experimenter (41) — ^Death in 1878,
State B\ineral (41).

^

CHAPTER V.

Respiration 42

Views of the Ancients on Respiration (42) — Mechanics of Respiration (42)
— Boyle and His Work (43) — Robert Hooke (43) — Mayow and His Re-
searches (44) — Respiration Prior to the Eighteenth Century (45) — The
Eighteenth Century School (46) — Priestly and His Dephlogisticated Air
(47) — Priestly and Benjamin Franklin (48) — Priestly Comes to America
(48) — The Phlogiston Theory (49) — Lavoisier and His Work (49) — The
First to Use the Word Oxygen (49).

CHAPTER VI.

The Nervous System 50

The Heart the Seat of the Soul thought the Ancient Hebrews (50) — ^The
Alexandrian School (50) — ^Galen Proclaimed the "Brain to be the Seat of
Thought and Sensation" (51) — Thomas Willis (51) — Frances Glisson Dis-
covers Irritability of Muscle (52) — Goll and Phrenology (52) — Bell and
Magendie (53) — Broca and the "Speech Center" (54) — Pathologic States
of the Brain and Nervous System (55) — Tuke, Benjamin Rush, and Pinel
(55).

CHAPTER VII.

i

^

The Cell Theory 56

Anticipated in the Seventeenth Century (56) — Bichat and His Contribu-
tion to the Theory (56) — The Theory Announced in 1838 (57) — Schleiden
and Schwann (57) — Johann Muller (58) — Vitalism (59) — Virchow's Eu-
logy on Muller (60) — Years of Discovery (58) — ^Virchow and "Cellular"
Pathology (60) — The Discovery of Protoplasm by Dujardin (61) — Proto-
plasm Defined by Starling (62) — The Cell Theory at the Present Time
(62)— The Nucleus (63) — Illustration of the Cell and Cell Division (64)—
The Cell in Heredity (65). ,^

f V

"There is no knowledge so useful to man as knowledge of
himself. Health and happiness are promoted by it. Before
the advent of the modem scientific spirit, biologic knowledge
was required to conform to the dominant superstitions of the
time. The human body was regarded as a peculiar and awful
thing, and not amendable to the laws which govern the rest
of the universe. Then it was found that the mechanics of the
body are entirely reconsilable with the principles of physics.
Humanity's debt of gratitude is incalculably great to those
men who at the risk of their lives and fortunes made dissec-
tions of dead bodies of men and animals, and discovered the
mechanism of the muscular system which imparts motion to
the joints, the valvular and pump-like arrangement of the
heart, and the hydraulic principles of the tubes which convey
the blood through the body. Then came those students of the
secrets of nature who discovered that the same laws which
govern man govern the lower and the lowest of creatures ; that
between soil and mineral, fluids and gases, plants and animals,
there is no dividing line; that the lily is the daughter of the
pool, and the man is the brother of the ox. This knowledge
was gotten for us, not by the philosopher among his books,
but by the patient investigator who went to the heart of
nature and studied her secrets." — J. P. Warbasse.

CHAPTER I.

THE CIRCULATION OF THE BLOOD— WILLIAM HARVEY

"This man lived in an age when alchemy was more popular than science,
and the love of mystery stronger than the love of philosophy." — Gorton.

"History is simply the biography of the mind of man; and our interest in
history, and its educational value to us, is directly proportionate to the complete-
ness of our study of the individuals through whom this mind has been manifested.
To understand clearly our positions in any science today, we must go back to
its beginnings, and trace its gradual development, following out our laws, difficult
to interpret and often obscured in the brilliancy of achievements — laws which
everywhere illustrate this biography, this human endeavor, working through the
long ages; and particularly is this the case with that history of the organized
experience of the race which we call science." — Sir William Osier.

The Renaissance — The renaissance, that transitional movement
in Europe between the mediaeval and modern world, affected medicine
and the sciences at a much later date than art and letters. It began
with Petrarch and the humanists in the fourteenth century in Italy,
where it became manifest in painting and sculpture. The movement
was accelerated in the sixteenth century by the capture of Constanti-
nople by the Turks in 1509, and the dispersion of its Greek scholars
to the shores of Italy, which event opened anew the science and learn-
ing of the ancient world at an hour when the intellectual energy of
middle ages had reached its ebb. It is significant to note that Flor-
ence, so long the abode of intellectual freedom and art, welcomed with
extended arms the exiled Greek scholars. Her traders returned from
the East with ancient manuscripts as the most valuable portion of
their merchandise. But we are more immediately concerned with the
movement as it affected medicine and its allied studies. However
much the new learning promoted literature and art, its influence was
anything but favorable to the progress of science. Admiration for
the literature of ancient Greece while it engendered a love for poetry,
history and philosophy, had a similar effect in promoting a spirit of
veneration for the writings of Hippocrates, Ptolmey and Galen, so
that it became almost an act of impiety to question their teachingSi— ^
It was not until the sixteenth century, as we shall see, that the spell
of ancient authority was broken by the direct appeal to nature. It
was not until then that the anatomist determined at all cost to exam
ine the human body for himself and to be guided by his own obser
vations.

Anatomy and Physiology — As anatomy precedes physiology, in
order to adequately appreciate the work of Harvey, a brief account
of the progress in anatomy is necessary. The great anatomist of an-
tiquity, who surpassed all others, was^Saleri (1^0-200 A. D.). He
lived for a time at Pergamos and for five^ars^t Rome. He was a
man of talent both as observer and writer. His writings embody all
the important anatomical discoveries of his predecessors, enriched and
much enlarged by the results of his own originality. His observations,
however, were made upon the lower animals on the faith of which he

0

2 PATHFINDERS OP PHYSIOLOGY

expounded the human subject. Huxley declares that "No one can
read Galen's works without being impressed with the marvelous ex-
tent and diversity of his knowledge and by his clear grasp of those
experimental methods by which alone physiology can be advanced."
Romewaathe field of his greatest triumph as physician. So great
was ^tSierih influence that for more than a thousand years his works
heldSittdieputed sway over anatomigaljteaching until a greater name
arose in the person of Vesalius. ('Ves^it^s, born in Brussels the last
day of 1514, inherited from an ancestr/'of learned men a keen appe-
tite for scientific learning. His wasjbha^nd^ependen^^
niind which has characterized ?ns"countrymehr~beforE~and since his
faay. The great importance of his work lies in the fact that he over-
\ threw adherence to authority as a means of arriving at truth and
\ employed instead, observation and reason. Slavish obedience to author-
Sty characterized the thought and methods of the Dark Ages. This
was in accord with the ecclesiastical influence dominant during this
long period. It was the method of the theologian, which had, un-
fortunately, survived almost to our own day. Darwin was perhaps
'he most recent object of theological invective. As the Scriptures
were an infallable guide to spiritual truth, so the works of Galen were
unfailing guides to scientific truth. Vesalius was bitterly
opposed not only by the ecclesiastic forces, but by medical men
af his time. The theologians opposed him because, among other
things, he differed from the widely accepted dogma that man should
have one less rib on one side because according to Scripture Eve was
formed from one of Adam's ribs. He was also at variance with them
on the subject of the Resurrection bone. Vesalius was willing, how-
ever, to leave the matter with the theologians, since it did not appear
to him to be an anatomical question. Sir Michael Foster writes that
Vesalius "Tried to do what others had done before him — he tried to
believe Galen rather than his own eyes, but his eyes were too strong
for him ; and he cast Galen aside and taught only what he could see
and what he could make his students see, too. Thus he brought into
anatomy the new spirit of the time, and especially the young men of
the time answered with a new voice." It is said that students flocked
to his lectures, his audience amounting to some five hundred. The
history of anatomy precedes that of physiology as a logical sequence.
The work of Vesalius placed the structure of the human body in a

^new light.

j William Harvey was the first man to study and proclaim the f unc-

I tion of structures which Vesalius had in such a masterly manner

^^demonstrated.

"The work of Harvey," says Locy, "Was complemental to that of
Vesalius and we may safely say that, taken together, the work of
these two men laid the foundations of the modern method of inves-
tigating nature. * * * in what sense the observations of the two men
were complimental will be better understood when we remember that
there are two aspects in which living organisms should always be
considered in biological studies; the first, the structure, and then
the use that the structures subserve."

The new learning spread over Europe in a westerly and northerly
direction. England was the last to partake of its benign blessing.
England had but two universities — Oxford and Cambridge; France

WILLIAM HARVEY 3

had six; Germany eight; Italy sixteen. Medicine was a prominent
department in all of them. Compared with the reception accorded
literature and philosophy, science lagged in England. Green sums
up the situation (1645) : "Bacon had already called men with a
trumpet voice to such studies. But in England, at least. Bacon stood
before his age. The beginnings of physical science were more slow
and timid there than in any country of Europe. Only two discoveries .^
of any real value came from English research before the Restoration
— the first, Gilbert's discovery of terrestial magnetism, in the close of
Elizabeth's reign; the next, the great discovery of the circulation of
the blood which was taught by Harvey in the reign of James. Apart
from these illustrious names England took little share in the scien-
tific movement of the continent ; and her whole energies seemed to I
be whirled into the vortex of theology and politics by the Civil War:^^-''''^

Birth and Education — William Harvey was born in Folkstone,
England, April 1st, 1578. Very little is known of his early life. His
preliminary education was obtained at his native town, where he
made his first acquaintance with Latin. He proceeded to
the King's School, Cambridge, where he remained five years, and
afterward, at 16 years of age, entered Caius College, Cambridge, in
1593. Harvey even early in his school life possessed habits of minute
observation. His fondness for dissections and his love for compara-
tive anatomy had shown his mental bias from his earliest years. To
Caius, the founder of the College at Cambridge, is accredited the in-
troduction into England of the study of practical anatomy. He ob-
tained for his college a charter which allowed the authorities of the
institution to take annually the bodies of two criminals condemned to
death and executed at Cambridge, free of all charges, for the purposes
of dissection, with the view to increase the knowledge of medicine
and to benefit the health of her majesty's heges, without interfer-
ence on the part of any of her officers. To what extent the college
availed itself of the privilege is not known. In all probability Har-
vey pursued the course of study which consisted of a sound knowledge
of Greek and Latin ordinarily followed until he obtained his B. A.
degree in 1597. A year after graduation, at the age of twenty, we
find him traveling on the continent where he studied the scientific
branches tributary to medicine, as well as medicine itself. As has
been said, the universities of northern Italy were the first to welcome
the new learning as it emanated from the east in the minds of Greek
scholars, as well as rescued manuscripts. The universities of north-
ern Italy, namely, Bologna, Padua, Pisa and Pavia, were at the time
at the height of their renown as centers of mathematics, law and
medicine. Harvey studied more particularly at Padua, renowned for
its anatomical school, and rendered famous by the work of such men
as Vesalius, the first of modern anatomists, and his successor, Fabri-
cius. The tolerance shown towards Protestants in Padua, the univer-
sity town of Venice, the great commercial republic, attracted many ^
law and medical students from England and other Protestant coun- vC^
tries of Europe.

It is interesting to recall that each entry in the university (Pa-
dua) register was accompanied by a note describing some physical' .
pecularity of the student, as a means of his identification. Thus j
Johannes Cookaeus, Anglus cum cicatrice in articulo medii di^iti die

if

4 PATHFINDERS OF PHYSIOLOGY

dicta. John Cook, an Englishman, with a scar over the joint of his
middle finger. (Matriculated) on the same day, and so on. Harvey
evidently did not enter Padua University as a regular matriculant,
as no such record occurs on the university register regarding him.

Fabricius and Harvey Friends — The fame of some of its medical
teachers undoubtedly attracted Harvey to Padua. While there he
was instructed in anatomy and physiology by Fabricius, one of the
most learned scholars of Italy. The fame as anatomist and surgeon
"j^ of Fabricus ab Aquapendente (from the name of his birthplace)
^^ phad spread well over Europe. During Harvey's sojourn in Padua he
^^Vand Fabricius became fast friends. At that particular time Fabricius
'' >was engaged in perfecting his knowledge of the valves of the veins.

His idea was that these valves prevented over-distention of the ves-
sels when the blood passed from the large to the smaller veins, while
they were not required in the arteries because the blood was always
in a state of ebb and flow. Harvey, however, pointed out their true
importance as anatomical proof of the circulation of the blood. It
was not so much what Harvey learned from Fabricius, as the stim-
ulus of his friendship that proved of such great assistance to him,
for we can see even in the instance quoted his view of the purpose of
the valves of the veins was entirely incorrect.

In 1602, Harvey was graduated M. D. from Padua. His diploma
conferred upon him the degree of Doctor of Physic, with leave to
practice and teach arts and medicine in every land and seat of learn-
ing. It further stated, that "he had conducted himself so wonder-
fully well in the examination and had shown such skill, memory and
learning that he had far surpassed even the greatest hopes which
his examiners had formed of him. They decided, therefore, that he
was skillful, expert and most efficiently qualified both in arts and
medicine, and to this they put their hands unanimously, willingly
and with complete agreement and unhesitatingly." The University
of Cambridge conferred the degree of M. D. on him the same year.

Harvey married in 1604, the daughter of Dr. Browne, who was
physician to Queen Elizabeth and to James I.
nV)***U^ Harvey, as we shall see, excjgUed as lecturer. His lectures
^^ — - showed an intimate acquaintance with the anat?nnical structure of
more than sixty kinds of animals, as well as a thorough knowledge of
human anatomy, which must have taken years of study to acquire.
He was elected fellow of the College of Physicians in 1607. An im-
portant position which Harvey held was Physician to St. Bartholo-
mew's Hospital in 1609. "The charge of the Physician of St. Bar-
tholomew's Hospital" required the incumbent to devote at least one
day a week throughout the year to charity. He was further enjoined,
"not for favour, lucre, or gain, to appoint or write anything for the
poor but such good and wholesome things as he shall think with his
best advice will do the poor good, without any aff:ection or respect to
be had to the apothecary. And he shall take no gift or reward of
any of the poor of this house for his ;counsel." This "charge" Har-
vey is said to have faithfully observed.

Anatomical Teaching Previous to 1745 — During Harvey's day
and until 1745, the teaching of Anatomy in England was vested in a
few corporate bodies. Private teaching was discouraged by fine and

WILLIAM HARVEY 5

imprisonment. The College of Physicians and Barber Surgeons had a
monopoly in London. The value of Anatomy as a foundation to medi-
cine was fully recognized at the time . The subjects for dissection
were the bodies of executed criminals. Those were the times of
public executions, witnessed by immense crowds whose opposition
and sympathy for the felon and his friends often interfered with the
procuring of the body for dissection.

The method of anatomical instruction is of interest. The sub-
ject was taught practically by a series of demonstrations on the body.
The absence of means of preservation of cadavers precluded instruc-
tion in detail. A single body was dissected to show the muscles ; an-
other to demonstrate the bones, and a third to exhibit the viscera.
Attendance on anatomical lectures and demonstrations was com-
pulsory; violation meant the forfeiture of a fine. Some were ex-
empted from the penalty, as one entry shows that a Robert Mudsley
"has licence to be absent from all lectures without payment of any
fine, because he has given over the art of surgery, and doth occupy
only a silk shop and shave."

The anatomical demonstrations were open to the public. The
following note appears in Pepy's Diary:* "Up and to my office.

. . . Commissioner Pett and I walked to Chyrurgeon's Hall (we
being all invited thither, and promised to dine there) , where we were
led into the Theater; and by and by comes the reader, Dr. Teame,
with the master and company in a very handsome manner; and all
being settled, he began his lecture, this being the second upon the
ureters and kidneys, which was very fine; and his discourse being
ended, we walked into the hall, and there being a great store of com-
pany, we had a fine dinner and good learned company, many Doctors
of Physique, and we used with extraordinary great respect . . . After
dinner Dr. Scpfeorott^:M;ook some of his friends, and I went along
with them to\see the bod^ alone, which we did, which was a lusty
fellow, a seaman that was hanged for a robbery. I did touch the
dead body with my bare hand; it felt cold, but methought it was a
very unpleasant sight. . . . Thence we went into a private
room where I perceive they prepare the bodies, and there were the
kidneys and ureters, etc., upon which he read today, and the doctor,
upon my desire and the company's, did show very clearly the man-
ner of the disease of the stone and the cutting and all other questions
that I could think of." Pepy's interest in the operation of cutting
for stone is said to be due to the fact that he had undergone the
ordea>hiTn5eif.-'*'Tk^Dr. Scarborough mentioned in Pepy's note was
a f rifend and pupil oiHarvey.

IPersonal Characteristics — Harvey is described as a man of the
**lowest^^«tature, round facejd, with a complexion like the wainscot;
his eyes smaliriuund, verSMblack and full of spirit, his hair black as
a raven and curling ; rapid in his utterance, chivalric even to gesture,
and used when in discourse with anyone to play unconsciously with

♦Samuel Pepys (1632-1703), was a famous diarist. His Diary, which
extends from 1660 to 1669, was written in shorthand, and was deciphered
by Lord Braybrooke in 1825. This delightful book of gossip is one of the
most interesting memorials of the domestic life of the time.

6 PATHFINDERS OF PHYSIOLOGY

the small dagger he wore by his side." His individuality was marked,
as was evidenced by the strong impression he made upon those with
whom he came in contact. His intellectual power and independence
of character were unusual. His interests were wider than his scien-
tific studies. According to an anonymous biographer* of the eight-
eenth century, "He was well read in ancient and modem history;
and when he was wearied with too close attention to the study of
nature, he would relax his mind by discoursing to his friends on
political subjects and the state of public affairs. He took great
pleasure in reading from the ancient poets, and especially Virgil,
with whose work he was exceedingly delighted. He was laboriously
studious, regular and virtuous in his life and had a strong sense of
reli^^on^ In his familiar conversation there was a mixture of^-gi'avity
and cheerfulness; he expressed himself with great perspicuity, and
with much grace and dignity; and was eminent for his great candor
and moderation. He never endeavored to detract from the merit of
other men; but appeared always to think that the virtues of others
were to be imitated and not envied."

In spite of his choleric and hasty disposition he had the faculty
of making close friendships. His replies to his critics showed great
moderation. Harvey's true character is probably best seen in that
period of his life which was beset with opposition and reproach, im-
mediately following the publication of his great work on the circu-
lation. TaJais_traducer^-h^ of the divine
Master, "To return evil speaking with evil speaking I hold to be un-
worthy of a philosopher and searcher after truth. I believe I shall
do better and more advisedly if I meet so many indications of ill-
breeding with the light of faithful and conclusive observation." His
attitude also resembles that of Darwin who, on the publication of his
Origin of the Species, was met with a storm of abuse from clerical
ignorance. It is said that the great evolutionist not only observed a
tranquility impassionate and unique but even condescended to reply at
length with courtesy to the rantings of those who vilified without
even reading his work or comprehending the object of their denunci-
ations,

Harvey was not a religious man in the narrow sense of the term
despite the fact that he lived in an age of warring creeds. His views
were broad as befitted a student of the design and workmanship of
the Great Architect of the universe. Aceor4ingjto Sir Russell Rey-
noM«,J^ devout and reverential recognition of God"' permeated his
work, "not only as the great primal ever-acting force, defined outside
and before all the works of nature; but as the Being, 'the Almighty
and Eternal God' to whom he says in his last will and testament, *I
do most humbly render my soul to Him who gave it; and to my
blessed Lord and Saviour Jesus Christ.' "

Harvey's knowledge of Latin was so thorough that he could con-
verse with facility equal to his native tongue. He was accustomed
to employ both English and Latin even in the same sentence, for ex-
ample, speaking of the eyes and their function: "Oculi eodem loco,
viz, nobilissimi supra et ante ad processus eminentes instar capitis
in a lobster snayles comubus tactu pro visu utuntur unde oculi as a
centinell to the army locis editis anterioribus."

•British Biographies, Vol. IV., London, 1768.

WILLIAM HARVEY 7

Harvey as Lecturer — Harvey's lectures were partly read and
partly oral. The cadaver lay on the table with the dissecting instru-
ments close to it. An assistant dissected or demonstrated while the
lecturer read his remarks. The anatomical lecturer of the sixteenth
century was a personage of importance. The greatest consideration
was exercised for his personal comfort. The stewards were instruct-
ed, *'to see and to provide that there be a mat about the hearth in the
hall that the Doctor be made not to take cold upon his feet. * * *
And further, that there be two fine white rods appointed for the
Doctor to touch the body where it shall please him ; and a wax candle
to look into the body, and that there be always for the Doctor two
aprons to be from the shoulder downward and two pair of sleeves
for his whole arm. . . . and not to occupy one apron and one
pair of sleeves every day, which is unseemly." Harvey laid down the rf
following precepts for his own guidance as lecture precepts which (
the modem anatomical lecturer might observe with propriety: j

(1) To show as much as may be at a glance, the whole belly
for instance, and afterwards to subdivide the parts according to their
position and relations.

(2) To point out what is peculiar to the actual body being dis-
sected.

(3) To supply only by speech what cannot be shown on your
own credit and authority.

(4) To cut up as much as may be in the sight of the audience.

(5) To enforce the right opinion by remarks down from far
and near and to illustrate more by the structure of animals accord-
ing to the Socratic rule.

(6) Not to praise or dispraise other anatomists, for all did well
and there was some excuse even for those who are in error.

(7) Not to dispute with others.

(8) To state things briefly and plainly.

(9) Not to speak of anything which can be explained without
the body or can be read at home.

Here we have a combination of orthodox medical ethics and
sound pedagogy. Harvey's particular role as *Lumlian lecturer in-
cluded the position of lecturer upon the viscera. Discussing the tho-
racic viscera he ennunciated the remarkable discovery with which
his name is inseparably associated, initialing the notes to indicate
that the ideas were peculiarly his own.

constat per fabricam cordis sanguinem.
per pulmones in Aortam perpetuo.
Transf erri, as by two clacks of a
water bellows to rayse water,
constat per ligaturam transitum saguinis
ab arteriis ad venas
unde perpetuum sanguinis motum
in circulo fieri pulsu cordis,
W. H.

♦The Lumlian lecture was a surgical lecture established at a cost of
£40 a year, which sum accrued from the rental of lands of Lord Lumley, of
Essex, England.

8 PATHFINDERS OF PHYSIOLOGY

"It is plain from the structure of the heart that the blood is
passed continuously through the lungs to the aorta as by the two
clacks of a water bellows to raise water.

"It is shown by the application of a ligature that the passage of
the blood is from the arteries into the veins.
( "Whence it follows that the movement of the blood is constantly

t< ^ in a circle and is brought about by the beat of the heart." It was
^ / not until twelve years after this important announcement that he
S(^ proclaimed it to a wider audience.

' Harvey's literary style was somewhat figurative. He loved to

indulge in metaphors — witness:

An cerebrum rex, whether the brain is king.
Nervi majistratus, the nerves his ministers.
Musculi cives populus, the muscles the, citizens or the people.
He also draws a similtude liking the brain to a military com-
mander, the leader of an orchestra, an architect, and he speaks of the
muscles and nerves as subordinate officers.

Year by year Harvey delivered the Lumlian lectures to the Col-
lege of Physicians. His private practice grew so as to be fairly lucra-
tive.

Harvey and Bacon — In 1618 he was appointed physician to
James I. In 1631 he was appointed physician in ordinary to King
James' son, Charles I. Not only gained he an entrance to the house-
hold of the king but he, was employed in .the homes of the most dis-
tinguished nobles. Among others he attended Sir Francis Bacon,
who was always a weak and ailing man with a disposition to be hypo-
chondriac. "In William Harvey and Francis Bacon," says Gorton,
may be observed two men like planets in conjunction; bom in the
same generation, each illustrious in the annals of history, the one in
philosophy, the other in science but in striking contrast to each
other. The one was a thinker, the other was an actor; one con-
ceived methods, the other put methods into operation; one was an
academic philosopher, the other a man of science and discovery; one
immortalized himself by his profundity of thought, the other by his
contribution to science. Both were stars in the firmament of great
men, but long after one has become dim or gone out, the other will
continue to shine with splendor."

Though honored by England's Lord Chancellor as the custodian
of his health, Harvey evidently failed to be impressed with Bacon's
greatness even as philosopher, for speaking of him, Harvey refers to
him as "writing philosophy like a Lord Chancellor."

p Publication of His Work on the Circulation — In 1628, the crown-

^ ing event of his life took place when he published his well considered
/and matured account of the circulation of the blood. He had demon-
strated his ideas of the circulation for twelve years before publishing
them, which event occurred in the fiftieth year of his life. This
monumental work of the great physiologist was accomplished while yet
in his thirties. Why Harvey should allow so much time to elapse be-
tween the event of his epochal discovery and its publication is not
clear. Evidently the passion to rush into print was not so great as
it is with the investigator of to day. It is interesting to note, how-

WILLIAM HARVEY 9

ever, that among the greatest thinkers and investigators Harvey is
not unique in this respect. Copernicus is said to have detained his
"Treatise of Revolutions" thirty years before permitting its publi-
cation; Bacon kept his Novum Organum by him for twelve years;
Isaac Newton "brooded in silence over the motion of the spheres"
for more than twenty years before publishing his Principia ; between
the first draft and the publication of the Origin of the Species seven-
teen years were permitted to intervene. Perhaps it was Harvey's
reluctance toward "quitting the peaceful haven," that constrained
him for so long a time, for elsewhere he tells us that his practice
fell off or, to use his own words, he "fell mighty in practice." Re-
garding him a contemporary wrote, "though all of his profession
would allow him to be an excellent anatomist, I never heard of any
who admired his therapeutic way. I knew several practitioners in
this town that would not have given three pence for his bills (pre-
scriptions) as a man can hardly tell by his bills what he did aim at."
Harvey is said to have been the first to be persecuted by the medical
profession for making discoveries at variance with the drift of public
thought and opinion. The story of all discoveries of the first rank
has borne out Locke's aphorism that "Truth scarce ever yet carried
by vote at its first appearance." The greatest obstacle to the accept-
ance of truth seems to be our present knowledge. Men are by nature
conservative; they resent innovations. Bagehot tells us that the
"pain of a new idea is one of the greatest pains to human nature."
Socrates somewhere likens himself to a midwife but his peculiar
function in life was to assist in that mental labor which gave birth
to ideas, a similitude which is suggestive of pain. The man who ex-
presses a new idea is apt to be abused, perhaps stoned. Whatever
may be said of the twentieth century the scientific world can be ac-
cused no longer of tardiness in the acceptance of new truth, but it
reserves the right to "prove all things and to hold fast to that which
is good." While Harvey's practice may have fallen off, his discovery (^
did not by any means consign him to obscurity. He still found favor T
with King Charles I, whose personal physician he was. His constant V
attendance at court greatly interfered with his duties at St .Bar-
tholomew's Hospital and resulted in the appointment of an assistant, 7
but with no diminution in Harvey's stipend. A contemporary of Har-*j
vey states as follows: "I have heard him say that after his Booke
of Circulation of the Blood came out he fell mightily in practice, and
'twas believed by the vulgar that he was crack-brained, and all the
physicians were against him, with much adoe at last in about twenty
or thirty years' time it was received in all the universities of the
world, and as Dr. Hobbs says in his book 'De Corpore,' he is the only
man perhaps that ever lived to see his own doctrine established in his
lifetime ;" sreritas est magna et prevalebit I

And yet, after the discovery has beejijrje^ognized as one of mo-
mentous import, the scientist has his defector|^ Harvey was no ex-
ception. There were those who sought ta'7i!siJfove^Ql£L.03PiginaU^^^ of
his work. Some attributed the merTf*"ornST§c6vering the circula-
tion to Servetus, some to Realdus Columbus, others to Caesalpinus.
True, Servetus, a Spaniard, bom in 1511 and burned at the stake in
Geneva, 1533, at the bidding of Calvin, in a copy of his Restitutio,
which was saved when an edition of 1,000 copies met the fate of the
author, rejected the contention that the blood passed through the

10 PATHFINDERS OF PHYSIOLOGY

cardiac septum. He had grasped the true features of the pulmonary
circulation — the passage of the blood from the right side to the
lungs, thence to the left side or ventricle. Realdus Columbus, bom
at Cremona, 1516, a presumptuous personage, speaks of the blood
carried, "by the artery-like vein to the lung and being there made
thin is brought back thence together with air by the vein-like artery
to the left ventricle of the heart." Then he goes on to press his claim
by declaring that, hitherto, no one had made this observation or re-
corded it in writing. Andreas Caesalpinus Was bom at Arezzo in
1519. He held for many years the professorship of medicine at Pisa.
Learned in all the lore of the ancients, he was noted among other
things for his determined opposition to Galen; Caesalpinus appears
to have grasped one important truth, namely, that the heart at systole
discharges its contents into the aorta and pulmonary artery, and at
its diastole receives blood from the vena cava and pulmonary vein.
Let all this be granted, yet the great work of Harvey is not a
whit less meritorious. The steam engine was in existence before the
day of James Watt, yef his name is inseparably associated with the
invention which transformed a mere toy into a gigantic factor which
has revolutionized human industry. No person, not even the genius is
independent of his time ; he is the heir of all the ages, and his great-
ness does not depend so much in presenting something unprecedented
as it does in seeing something clearly and tening_in^.jL.simple way
what he has seen. ^.-^'^"'^^^^^^^ ^^^

Treatise on the Circulation. — Harvey's greatest work was un-
doubtedly his Exercitatio Anatomica de Motu Cordis et Sanguinis in
Animalibus, an anatomical treatise on the movement of the heart
and blood in animals, published in Frankfort, Germany, in 1628. The
book was a small quarto volume of 72 pages. It opens with a dedi-
cation to "The Most Illustrious and Indomitable Prince, Charles, King
of Great Britain, France, and Ireland, Defender of the Faith," etc.
The dedication proceeds : "The heart of animals is the foundation of
their life, the sovereign of everything within them, the sun of their
microcosm, that upon which all growth depends, from which all
power proceeds. The king in like manner, is the foundation of his
kingdom, the sun of the world around him, the heart of the republic,
the fountain whence all power, all grace doth flow." Whatever may be
said regarding Charles I, who was the victim of public execution, he
certainly befriended Harvey. Then to the president of the Royal Col-
lege of Physicians and to other learned physicians the author ad-
dresses himself in a dedication which he concludes : *****! profess
both to learn and to teach anatomy not from books but from dissec-
^tions; not from the positions of philosphers but from the fabric of
nature. * * * I avow myself the partisan of truth alone: and I can
indeed say that I have used all my endeavors, bestowed all my pains
on an attempt to produce something that should be agreeable to the
good, profitable to the learned, and useful to letters." Harvey's
method here ennunciated is the method of every scientist since his
day, whose contribution has possessed real meri1> — that is, reasoning
based upon experiment and observation. ^^--^ ^n

The work on the circulation comprises seventeen short chapter's.
It is an interesting account, lucid and connected, of the heart's' action
and the circulation of the blood. Harvey had no means of knowing

WILLIAM HARVEY 11

the connection between the smallest arteries and the smallest veins,
for the microscope was not in such a stage of perfection as to
permit of much fine work in minute anatomy. It was not until the
invention of the compound microscope in 1675 that Leeuwenhoek
described blood corpuscles and the capillary circulation. In the first
chapter the author reviews some of the fantastic theories regarding
the functioning of heart and lungs. The heart was held to be the
great heat center of the body. The blood was sucked into it during
diastole and expelled from it during systole. The arteries cooled the
blood; the lungs fanned and cooled the heart. The term "spirits''^
meant a great deal to Harvey's predecessors, but not to him. ^Xhe^
word blood has nothing of grandiloquence about it, for ir^Igmfies a
substance which we have before our eyes and can touch; but before
such titles as spirit and calidum innatum (inherent heat) we stand
agape." , ^^

Chapter I, he continues: ^' .^^^

"When I first gave my mind to vivisections, as a means of discovering the
motions and uses of the heart, and sought to discover these from actual inspec-
tion, and not from the writings of others, I found the task so truly arduous, so
full of difficulties, that I was almost tempted to think, with Fracastorius, that
the motion of the heart was only to be comprehended by God. For I could
neither rightly perceive at first the systole and when the diastole took place, nor
when and where dilatation and contraction occurred, by reason of the rapidity
of the motion, which in many animals is accomplished in the twinkling of an eye,
coming and going like a flash of lightning; so that the systole presented itself to
me now from this point, now from that; the diastole the same; and then every-
thing was reversed, the motions occurring, as it seemed, variously and confusedly
together. * * *

"At length, and by using greater and daily diligence, having frequent re-
course to vivisections, employing a variety of animals for the purpose, and collat-
ing numerous observations, I thought that I had attained to the truth, that I
should extricate myself and escape from this labyrinth, and that I had discovered
what I so much desired, both the motion and the use of the heart and arteries;
since which time I have not hesitated to expose my views upon these subjects,
not only in private to my friends but also in public, in my anatomical lectures
after the manner of the academy of old."

He goes on to tell how his views pleased some, displeased others.

He finds it advantageous to study the movement of the heart in
the cold-blooded animals — frogs, snakes and fishes. He ascertained
that the heart was a muscular organ, that its systole was the result
of muscular contraction. The contraction of the heart was more im-
portant than its dilitation. "During its contraction the heart becomes
erect, hard and diminished in size, so that the ventricles become
smaller and are so made more apt to expel their charge of blood. In-
deed, if the ventricle be pierced the blood will be projected forcibly
outward at each pulsation when the heart is tense." Harvey -shQwM__
that the pulsation of the arteries depended upon the contraction of
the left ventricle. The contraction of the right ventricle propelled

*The extracts which follow illustrate Harvey's style. The Motion of the
Heart and Blood, by William Harvey, can be procured in convenient form
in the Everyman's Library Series (E. P. Button «& Co., New York). This is
a reprint from the Sydenham Society's edition of 1847.

12 PATHFINDERS OF PHYSIOLOGY

the blood into the pulmonary arteries, the pulsations of which were
simultaneous with the other arteries of the body. He demonstrated
that the two ventricles contracted simultaneously and that the two

,/ auricles contracted at the same time.

iJ

0 AT Motion, Action and Office of the Heart. — In the fifth chapter

^V^^;^arvey deals with the motion and function of the heart. It j:fiad*j-

j^y somewhat like a modernjwprkjn physiology.

/ "First of all, the auricle contracts, and in the course of its contraction

throws the blood (which it contains in ample quantity as the head of the veins,
the storehouse, and cistern of the blood), into the ventricle, which, being filled,
the heart raises itself straightway, makes all, its fibres tense, contracts the ven-
tricles, and performs a beat, by which beat it immediately sends the blood sup-
plied to it by the auricle into the arteries; the right ventricle sending its charge
into the lungs by the vessel which is called vena-arteriosa, but which, in structure
and function, and all things else, is an artery; the left ventricle sending its
charge into the aorta, and through this by the arteries to the body at large.
These two motions, one of the ventricles, another of the auricles, take plpce con-
secutively, but in such a manner that there is a kind of harmony or rhythm pre-
served between them, the two concurring in such wise that but one motion is
apparent, especially' in the warmer blooded animals, in which the movements in
question are rapid."

So far as Harvey's reasoning is based upon his observations his
conclusions are in the main correct, as proved by more recent re-
search ; where he indulges in speculation we get the following :

"In the larger and more perfect animals of mature age Nature
has rather chosen to make the blood percolate the parenchyma of the
lungs. * * * It must be because the larger and more perfect animals
are warmer, and when adult their heat greater, ignited I may say
and requiring to be damped or mitigated, that the blood is sent
through the lungs, in order that it may be tempered by the air that
is inspired and prevented from boiling up and so becoming extin-
guished or something else of the sort," or, to modernize it, the lungs
serve as radiator and the heart the gasoline or internal^ combustion
engine. /^"^^^

Capillary Circulation. — Since Harvey's time Malpighi, in 1661,
hinted at the capillary circulation, which was still further investi-
gated by Leuwenhoek in 1674, who studied it with his microscope in
the web of a frog's foot and in other transparent membranes. In
1676, Blankaart, and in 1697 Cowper, studied the arrangement of the
capillaries by means of injected specimens. A long interval elapsed
between the histological study of the circulation before chemistry
was sufficiently advanced to afford definite knowledge in regard to
oxidation of the blood and the explanation of the true function of
the lungs. The work of Priestly in 1775 was a notable contribution
to the physiology of respiration. The nineteenth century, through
the work of Ludwig in Germany, Chauveau in France, and Foster in
England, has seen the physics of the heart and circulation reduced
almost to an exact science. - )

Any account of the works of Harvey would be incomplete were
no mention made of his work in embryology. Harvey discussed the
nature of development and exhibited extraordinary powers as re-

WILLIAM HARVEY 13

gai:ds accuracy of reasoning. He may be considered as having made
the lSrs1ria4epejideiit-M^ in the subject. That he did not ac-
compHsh more was due to laclc oFinstruments of precision, and to
the fact that he had to build on the general level of the science of
the time. His work on embryology was published in 1651. It was
entitled "Exercitationes de Generatione AnimaHum." In it is an
account of not only the development of the chick, but of deer and
other mammals as well.

All honor to him who blazes the trail. The refinements, what-
ever they may be, can never merit for the investigator the honor
which is due the ^ioneerJ As was said by Haller, one of the best \
informed minds of the eighteenth century, "It is not to Caesalpinus, 1 1
because of some words of doubtful meaning, but to Harvey, the able / [
writer, the laborious contriver of so many experiments, the staid y
propounder of all the arguments available in his day, that the im- I
mortal glory of having discovered the circulation of the blood is to J
be assigned."

One of his last acts was to set aside a certain sum derived from
his estate for the delivery of an oration in commemoration of the
benefactors of the College of Physicians. This oration, the Har-
veian Oration, is still delivered each year by some distinguished mem-
ber of the medical profession. Even in his declining years his
thoughts were turned to the future. The Harveian Lecture is in-
tended to further the progress of science, especially a knowledge of
the body in health and disease. "Much of the nobility of the profes-
sion," says Osier, Harveian lecturer, 1906, "depends upon the great
cloud of witnesses* who pass into the silent land — ^pass and leave no
sign, becoming as though they had never been bom. And it was the
pathos of this fate not less prophetic because common to all but the
few, that wrung from the poet that sadly true comparison of the race ~y
of man to the race of the leaves." Harvey was one of the "few" to <^
have achieved that immortality which places him with "The divine J
men of old time."

He died June 3rd, 1657, in the eightieth year of his age.
Asellius and the Lymphatic Circulation.

Corollary to the circulation of the blood is the lymphatic circula-
tion. The discovery of the lymphatics was almost synchronous with
that with which Harvey achieved an immortal name. While the
memory of Harvey has been fittingly honored in various ways, that
of Asellius or Aselli has not been sufficiently recognized. The data
referring to Aselli's life are extremely meagre. He was bom in 1581,
at Cremona, Italy, the descendant of a patrician family. He studied
at the University of Pavia, where he became laureate in medicine,
surgery and philosophy, after which he located in Milan, where he
taught anatomy privately and engaged in the practice of surgery. It
was while in Milan that he made his discovery, in 1622, of the lym-
phatic vessels which he called venae lactae. His discovery was rec-
ognized by his election, two years later, to the chair of anatomy and
surgery in his alma mater, a position he was destined not long to
hold, for he died in 1626 at the age of forty-five. His book De Lac-
tibus was published a year after his death. William Harvey was

14 PATHFINDERS OF PHYSIOLOGY

forty-four years old at the time of Aselli's discovery. Aselli's dis-
covery of the lacteals is related by himself as follows :

"On the 23rd of July in that] year (1622) I had taken a dog in good condi-
tion and well fed, for a vivisection at the request of some friends, who very
much wished to see the recurrent nerves. When I had finished this demonstra-
tion of the nerves, it seemed good to watch the movements of the diaphragm in
the same dog, at the same operation. While I was attempting this, and for that
purpose had opened the abdomen and was pulling down with my hand the intes-
tines and stomach gathered together into a mass, I suddenly beheld a great
number of cords, as it were, exceedingly thin and beautifully white, scattered
all over the whole of the mesentery and the intestine, and starting from almost
innumerable beginnings. At first I did not delay, thinking them to be nerves.
But presently I saw I was mistaken in this, since I noticed the nerves belonging
to the intestine were distinct from these cords and wholly unlike them, and,
besides, were distributed quite separately from them. Wherefore struck by the
novelty of the thing, I stood for some time silent while there came to my mind
the various disputes, rich in personal quarrels no less than in words, taking place
among anatomists concerning the mesariac veins and their function. And by
chance it happened that a few days before I had looked into a little book by
Johannes Costaeus written about this very matter. When I gathered my wits
together for the sake of the experiment, having laid hold of a very sharp scalpel,
I pricked one of those cords, and indeed one of the largest of them. I had
scarcely touched it, when I saw a white liquid like milk or cream forthwish gush
out. Seeing this, I could hardly restrain my delight, and turning to those who
were standing by, to Alexander Tadinus, and more particularly to Senator Sep-
talius, who was both a member of the great college of the Order of Physicians
and, while I am writing this, the medical officer of health, 'Eureka,' I exclaimed
with Archimedes, and at the same time invited them to the interesting spectacle
of such an unusual phenomenon. And they indeed were much struck with the
novelty of the thing."

Aselli noted the presence of valves in the lymphatic vessels and
recognized their function, namely, to prevent the backward flow of
the lymph. He recognized also that the lacteals were vessels for con-
veying chyle away from the intestine. He went wrong, however, in
fregard to the ultimate course taken by the newly-discovered vessels,
[ for he thought he could trace them to the liver. Aselli was heavily
handicapped by his previous learning, which consisted of a Careful
study of as well as veneration for the teachings of the ancienti^
Galen had, in fact, taught that all nutritive material from digestive
processes passed through the liver. Aselli speaks in his book of a
group of lymphatic glands lying in the mesentery, as the pancreas —
hence the name pancreas Aselli. The force which caused the move-
ment of the fluid in the lacteal vessels was believed by him to be two-
fold, a vis a tergo and a vis a f rente ; the latter derived from supposed
suction of the liver and the former supplied by the movements of the
intestines.

Aselli Opposed by Harvey. Aselli in his modesty endeavored to
prove that the lacteals were known to the ancients, especially to
Herophilus and Erasistratus, founders of the Alexandrine school of
medicine. His discovery met the same opposition as did Harvey's, and
from the same men, among them Riolan and Primrose, and strange to
say Harvey himself failed to recognize the importance of the work

ASELLIUS 15

of his contemporary. In a private letter written in April, 1652, he
writes :

"With regard to the lacteal veins discovered by Aselli, and by the further
diligence ot! Pecquet, who discovered the receptacle or reservoir of the chyle,
and traced the canals thence to the sub-clavian veins, I shall tell you freely, since
you ask me, what I think of them. I had already, in the course of my dissections,
I venture to say even before Aselli had published his book, observed these white
canals. * * * But, for various reasons, and led by several experiments, I
could never be brought to believe that that milky fluid was chyle, conducted thither
from the intestines, and distributed to all parts of the body for their nourish-
ment; but that it was rather met with occasionally and by accident, and proceeded
from too ample supply of nourishment and a peculiar vigor of concoction ;" and
Harvey continues: "Why indeed, should we not as well believe that the chyle
(digested contents of the intestines) enters the mouth of the mesenteric veins
and in this way becomes immediately mingled with the blood, where it might
receive digestion and perfection. * * * And that the thing is so in fact, I
find an argument in the distribution of innumerable arteries and veins to the
intestines, more than to any other part of the body, in the same way as the
uterus abounds in blood vessels during the period of pregnancy."

Sir William Osier) (Harveian oration, 1906) refers to this inci-
dent in Harvey^s'career: **How eminent so ever a man may become
in science, he is very apt to carry with him errors which were in^
vogue when he was young — errors that darken his understanding^
and make him incapable of accepting^-eveii_tha-most--0bviaus truths.
It is a great consolation to know that Harvey came within the range
of this law — in the matter of the lymphatic system; it is the most
human touck in his career."

Th^ lacteals were demonstrated in man in 1628, the subject be-
ing an executed criminal examined shortly after execution. Twenty-
one years after Aselli's death the thoracic duct was discovered by
Johannes Pecquet, of Dieppe, France. He not only accurately de-
scribed these lymphatic structures, but showed that Aselli's lacteals
poured their contents into what he called the receptaculum chyli, but
that the thoracic duct — a continuation of the receptacle — poured its
contents into the venous system at the junction of the jugular and
subclavian veins. Pecquet was twenty-five years old when he made
this discovery, which he himself described as the gift of fortune
sporting with the ignorant. Munus est fortunae cum inscio ludentis.
Pecquet, however, did not follow up this solitary triumph. His ap-
petite for alcoholic beverages got the better of him and eventually
caused his death.

Harvey's work on the circulation appeared between the discov-]
ery of Aselli and that of Pecquet and so profoundly had it influenced^
the medical thought of the time, that the discovery of the thoracic)
duct and its function was accepted without question. y

CHAPTER II.

PHYSIOLOGY OF DIGESTION IN THE SEVENTEENTH
AND EIGHTEENTH CENTURIES

The circulation of the blood was worked out and proclaimed to
the world by one man, and his work was so complete that it has not
been rendered obsolete by subsequent knowledge. The history of the
physiology of digestion has been of gradual growth so that no one
man can claim credit for our present knowledge. Before the develop-
ment of chemistry, any marked progress in the physiology of alimen-
tation would not have been possible; the early workers in this par-
ticular field were chemists rather than physiologists. The history
of physiology during the seventeenth and eighteenth centuries in-
volves the lives and work of numerous investigators, each accomplish-
ing all that was possible considering the advancement of the general
scientific knowledge of the time.

Two names of the latter part of the seventeenth and early part
of the eighteenth century are prominent as exerting important infl-
ence in the way of solution of the chemical problems of physiology.
These were George Ernest Stahl and Hermann Boerhsa-^T'^^hf was
born at Anspach in 1660 ; he studied at Jena, and after graduating be-
came court physician at Weimer, and in 1694 professor of medicine at
Halle. He died in 1734 in Berlin, where he moved in 1716 on his ap-
pointment as physician to the King of Prussia. Stahl was an ac-
complished chemist of his day. His views on gastric digestion may be
summed up in the following sentence from his work: "Some people
suppose that gastric digestion results from the action of particular
and specific ferments, and indeed go so far as to regard the stomach
as not only the seat but also the origin of a particular ferment,
whereas in the whole construction of the stomach nothing particular
is observed which would render the elaboration of such a special
agent likely." He was a firm believer in the psyche of Aristotle and
introduced a principle which he termed anima. He was wholly out of
sympathy with those who tried to explain the physical and
psychical phenomena of life and mind on chemical and mechanical
principles. He could not think of himself as a chemical retort subject
to ferments. The soul was to him the living force of the body; "It
was susceptible of being played upon by a thousand different influ-
ences, such as joy, sorrow and grief, love and friendship, the beauti-
ful, the true, the reverent, the sublime. * * * Can these things be the
product of chemical acids and alkalies and the mechanical devices of
the mason and builder ?" Sir Michael Foster sums up the teaching of
Stahl thus : "Learn as much as you can of chemical and physical pro-
cesses, and in so far as the phenomena of the living body exactly re-
semble chemical and physical events appearing in non-living bodies,
you may explain them by chemical and physical laws. But do not
conclude that that which you see taking place in a non-living bodj"
will take place in a living body, for the chemical and physical phe-
nomena of the latter are modified by the soul. The events of the bod^
may be rough hewn by chemical and physical forces, but the soul will

PATHFINDERS OF PHYSIOLOGY 17

/ shape them to its own end and will do that by its own instrument, mo-
tion." Stahl, it will be seen, belonged to the^'Vitalists/' which par-
ticular type of physiologist has only within recent years become ex-^ ^ ,
tinct. His fundamental position was, between living and non-living^' v^^.
things there is a great gulf fixed. Living things so long as they are]
alive are actuated by the sensitive soul; non-living things are not.
The rational soul of man governed his whole body. The healing
power of nature, vis medicatrix naturae, has been recognized from
the most ancient to the present time. Stahl's system was founded
upon the supposition that the vis naturae existed entirely in the ra-
tional soul. In consequence of Stahl's doctrine, he and his followers
proposed the art of curing by expectation, medicina expectans, which ^
practice led to the prescribing of inert remedies, placebos.

Payer and Brunner — In the catalogue of workers in physiology of f^ ^^^
the seveneteenth century are the names of Jean Conrad Peyer and >^
Brunner. Peyer was bom in Switzerland in 1653. He studied at
Basel and Paris and returned to his native town, Schaffhausen, to
practise, where he died in 1712. In 1677 he published a brochure in
which he described certain new glands scattered" over the intestine;
these glands are familiar to every student of physiology or histology
as 'fPeyer's patches." He was the first to give a full description of
these glnds are familiar to every student of physiology or histology
lower part of the small intestine and in the ileum, making a distinction
between the single or solitary and the patches of agminated glands.
"His discovery harmonized with that of Brunner a few years later.

Brunner was born at Dieffenhausen in 1653. He studied at
Strassburg and was eventually called to the chair of medicine at Hei-
delberg, shortly after entering upon his position he published his
Dissertatio Inauguralis de Glandulis Duodeni, in which he describes
the glands which have since borne his name, Brunner's glands. He
attributed to these glands a function similar to the pancreas and
spoke of them as a "pancreas secondarium." Brunner had made num-
erous experiments by removing the pancreas from dogs. He con-
cluded that the animals thus operated suffered in no wise from ill
health, consequently the digestive powers of pancreatic juice were
practically nothing. These gropings of the seventeenth century are
curiously interesting viewed in the light of the twentieth. The work
of Peyer and Brunner served to deprive of its glory that of Sylvius
and DeGraaf, who had attributed important digestive powers to the
pancreatic juice. The attention of physiologists was again centered
on the older view that the stomach was the chief seat of digestion.

Mechanical and Chemical Views of Digestion. — Two views con-
cerning gastric digestion contended for first place. One, which may
be designated the mechanical, was espoused by Borelli, who was the
founder of the so-called latromathematical school, which professed to
be able to reduce all the motions and activities of nature to mathe-
matical formulae. Borelli's studies were made on the stomachs or
gizzards of birds. He pointed out the great grinding or pressing force
effected by the muscular coats of the stomach. He compares the ac-
•jon of the fleshy stomach to that of the teeth, and continues: "We
nave already shown that the absolute force of the muscles which close
the human jaw represents a power greater than that of a weight of
350 pounds; therefore, the force of the turkey's stomach is not \e?-^

18 BOERHAAVE

than the power of 1,350 pounds." This estimate of the power of the
' human muscles of mastication, is rather high. Canon in his recent
work places the pressure which the molars are capable of exerting at
270 pounds. Borelli admits, however, that certain animals ''consume
V flesh and bone by means of a certain very potent ferment, much in
^ the same way as corrosive liquids dissolve metals." The iatro-phys-
^ ical school eventually went farther than Borelli and denied that chem-
!; ical action has anything whatsoever to do with digestion, and con-
tended that digestion was mere trituration of the food in the stomach
to a creamy substance known as chyle. Bellini, a pupil of Borelli,
went farther in the beginning of the eighteenth century and endeav-
ored to explain many functions of the human body from mathemati-
cal data. Keill, a member of this cult, calculated from data purely
imaginary the power of each organ. According to him the stomach
had a force of compression so great that to overcome its own resist-
ance must have meant its own destruction. One iatro-physicist esti-
mated the force of the heart as equal to 180,000 pounds; another
placed it at eight ounces. Their calculations were clothed in the im-
posing nomenclature of the exact sciences. This doctrine is said to
have extended to all the universities and medical institutions of
Europe.

The iatro-chemical school, or "chemikers" as they were dubbed
by Guy Patin, a French physician and wit of the time, sought a solu-
tion of all the phenomena of the human body in their flasks and re-
torts. They maintained that the change in the stomach was chiefly
if not wholly a chemical, resulting from the process of fermentation.
It was recognized even at this time that the membrane of the stomach
was glandular in structure, and yet little importance was attached to
the secretion of such membraneous surface.

In 1614 was born Francois de le Boe or Dubois, better known by
his Latin name, Sylvius. He is not to be confused with Jacobus Syl-
vius, the Parisian anatomist, teacher of Vesalius, who lived in the
sixteenth century. The latter Sylvius studied at Sedan and at Basel,
where in 1637 he took his degree. He became professor of medicine
at Leyden, where he exerted a powerful influence until his death in
% 1672. Sylvius, though distinguished as a physician and physiologist,
^ was essentially a chemist. Through his efforts the curators of the
Universtiy of Leyden built for him a ''Laboratorium" which, so far
• as we know, was the first university chemical laboratory. He devoted
a large part of his time to a study of salts, which he learned to rec-
ognize as resulting from the union of acids with bases. Sylvius looked
upon the phenomena of life from a chemical point of view. He was
well versed in that part of physiology derived by deductions from an-
atomy and by experiments on animals. His opinions on the circula-
tion and respiration were orthodox from our modem viewpoint. Har-
vey's teachings entered largely into his thoughts and it was chiefly
through his advocacy that the doctrine of the great discoverer of the
circulation of the blood became established in Holland. The contribu-
tions which Sylvius made to science were essentially chemical.

Boerhaave — Herman Boerhaave, aready mentioned as a contrib-
utor to the chemical knowledge of alimentation, was born in 1668,
near Leyden, where he was educated. His early years were largely

devoted to the classical and oriental studies. He became Ph. D. in 1690,
m.

PATHFINDERS OF PHYSIOLOGY IJ*

having obtained the degree on a thesis, the subject of which was "The
Distinction Between Body and Mind." An illness in the shape of an
obstinate ulcer of the leg turned his attention to medicine, which he
studied along with the ancillary studies, chemistry and botany. He
was graduated M. D. in 1693, and eventually gave up the idea of the-
ology for medicine. In 1701 he was appointed to the chair of medicine
in the University of Leyden. His great ability as teacher caused stu-
dents to flock to his lectures. His worth was quickly recognized by
the authorities of the university who increased his emolument and
endeavored to make his position attractive to prevent him from going
elsewhere. Sir Michael Foster says of him: "Much sought after as
a physician, acute at the bedside, brilliant as an expositor in the pro-
fessorial chair, he was also a great teacher in the sense that in his
daily intercourse with his pupils he was alwaj/s ready to lay his mind
open before them and to let them share his experience and his
thoughts. Russell pays the following tribute to Boerhaave's genius:
"Boerhaave was easily the most remarkable physician of his
age, a man who, when we contemplate his genius, his condition, the
singular variety of his talent, his unfeigned piety, his spotless char-
acter and the impress he left not only on contemporary practice, but
on that of succeeding generations, stands forth as one of the brightest
names on the pages of medical history, and may be granted as an ex-
ample not only to physicians but to mankind." Boerhaave was a
scholar and scientific thinker, too broad to be the slave of one idea.
He was eclectic in the true sense of the term, though he never allied
himself with the medical sect which goes by that name. He had a
mind open to truth wherever it might be sought. He made use
of anatomy, physics and chemistry, but never allowed one to exclude
the other. He made each subservient to the elucidation of physiology.
Boerhaave was not an extreme advocate of either mechanical or
the chemical fermentative school; he recognized that digestion is in
part a solution of some of the constituents of food by means of
various juices, which he, however, regarded not of the nature of 1
fermentation. He denied, however, the acidity of the gastric juice. |
Colored vegetable juices were at the time coming to be used as we \
now use litmus paper, in reaction tests. Boerhaave regarded the )
solution by means of juices only as part of the digestive process; the
remaining process he held consisted of trituration in the stomach, by
which process the nutritive parts of food were expressed. His views
were dominant the early part of the eighteenth century.

An Epochal Year, 1757 — ^The years 1757 was the dividing line be-
tween modern physiology and all that had gone before. It was the
date of the publication of the first volume of Haller's Elementa Phys-
iologia, the eighth volume of which appeared in 1765. Albrecht von
Haller was born at Berne, Switzerland, in 1708. The story is told of
his early precosity, when at the age of four he is said to have ex-
pounded the Bible to his father's servants. Before he was ten, he
wrote in Latin verse a satire on his tutor. Haller's attention had been
directed to medicine after his father's death in 1721, while residing in
the house of a physician in Biel, and in his sixteenth year he entered
the University of Tubingen. Dissatisfied with his progress there, he
went to Leyden ,where Boerhaave was at the height of his fame. He
graduated in 1727, and turned his attention to botany, publishing a

/

\

20 HALLER

great work on the flora of Switzerland. He returned to Berne and be-
gan the practice of medicine in 1729. In 1736 he was appointed pro-
fessor of medicine, anatomy and botany in the newly founded univer-
sity of Gottingen, a position v/hich he had held for 17 years. During
t}iis time he carried on original inve^tigat«on in botany and physiology.
His researches on the formation of bone, the mechanics of respiration,
and the development of the embryo are of the highest importance. Re-
garding Haller as an expositor in physiology, Foster writes : "When
we turn from the preceding writers on physiology and open the pages
of Haller's Elementa, w^e feel that we pass into modern times. Save
for the strangeness of most of the nomenclature, and for no small
differences in all that relates to the chemical changes of the body, we
-, seem to be reading a modem text-book of the most exhaustive kind."
\His chief service, however, was the careful arranging and digesting
? of the theories and facts of physiology up to this time. From his time
physiology became an independent branch of science, to be pursued
for itself rather than as an adjunct to medicine. Regarding Haller's
method of exposition, the same writer goes on to say that "In dealing
with each subdivision of physiology, Haller carefully describes the
anatomical basis, including the data of minute structure, physical
properties and chemical composition, so far as these were then known.
He then states the observations that have been made, and in
respect to each question, as it arises, explains the several views
which have been put forward, giving minute and full references to
all the authors quoted, and he finally delivers a reasoned critical judg-
ment expounding the conclusions which may be arrived at, but not
omitting to state plainly when necessary the limitations which the
lack of adequate evidence places on forming a decided judgment. He
carefully recounts and as carefully criticizes all the knowledge that
can be gleaned about any question. If he feels unable to come to a
decided conclusion he candidly says so."

But we are most concerned at present with what Haller has to
say on digestion. He considered saliva neutral in reaction and pos-
sessing no digestive properties further than the softening of food as
an aid to deglutition. He recognized the importance of the glandular
coat of the stomach, which glands he concluded furnished mucous
only, the true gastric juice being derived from the arteries. He also
concluded that pure gastric juice was neither acid nor alkaline and
refused to regard it as some of his predecessors had done, as a fer-
ment. The acidity he considered a token of the degeneration of the
digested food. Trituration he regards as a useful aid to digestion, es-
pecially where hard grains form part of the food as in birds ; but it
was only an aid.

Bile, he claimed, was not a mere excrement; it was secreted by
the liver and stored for a time in the gall bladder, where it underwent
slight change. Bile is a viscid fluid, bitter but neither acid nor alka-
line. It has the power of dissolving fats, and so acts on a mixture
of oil and water as to form an emulsion. Haller considered the im-
portance of the pancreas due to the fact that its ducts opened into the
intestine in common with the bile duct ; that its fluid softened and
diluted the bile, thus enabling it to mix more satisfactory with the
food. He concluded by prophesying that there may be other func-
tions of pancreatic juice not well known to the physiologists of his
day.

PATHFINDERS OF PHYSIOLOGY 21

Reaumur and His Methods. Rene Antoine Ferchault de Reaumur,
a Frenchman bom in 1683, and described as one of the most notable
men of science of the eighteenth century, is, in chronological se-
quence next most important contributor to the physiology of the
alimentary tract. His name is already familiar to most of us as the
inventor of the Reaumur thermometer. His studies on the gastric
juice at this time are all-important, inasmuch as his methods are
unique. Reaumur had in his possession a kite and took advantage of
the habit of the bird of ejecting from its stomach things swallowed
which it could not digest. The kite was fed pieces of meat secured
in metal tubes. It was found that meat when ejected had no odor
of putrification. Experiments were made with small pieces of bone,
which were completely dissolved when ejected and swallowed by the
kite several times. On vegetable grains and flour, the fluid of the
kite's stomach had apparently little effect. The tubes were filled with
small pieces of sponge, which, when ejected, were squeezed out, thus
enabling the investigator to procure pure gastric juice and to study
it in vitro. He proved that digestion was not putrifaction but some^i
thing really opposed to that process. While Reaumur's experiments W
left much to be ascertained about gastric digestion, he at least favored \
the solvent power of the succus gatricus, by the employment of a J
wholly new method.

Experiments with Gastric Juice. We must look to Italy for the
next contributor to our knowledge of digestion. Parenthetically, it
is of interest to note that the idea of specializing, if it had taken root
at all at this early time, was not markedly apparent. The worker in
the physiology of digestion was equally prominent in almost every
other department of physiological research. Lazzaro Spallanzani
(1729-1799) was one of the most eminent men of his time. Educated
for the church, he was usually known as Abbe Spallanzani. His life
was devoted to experiments, researches and teaching. He was pro-
fessor at Bologna, and afterwards at Pavia. We find him first ex-
perimenting with germ life, with results that disprove the doctrine
of spontaneous generation. His researches in other fields showed that
he had conceived the truly scientific method.

Spallanzani took up Reaumur's methods and most of his results
were achieved by them. Aided by improvements in chemistry, he
was able to make marked advance over his predecessors. His ex-
periments were made on all kinds of animals, fishes, frogs, serpents,
birds, sheep oxen, horses, cats and dogs, and lastly upon himself. Be-
sides hollow tubes, he used hollow spheres, freely perforated, into
which were placed meat and bread, bone or grains of wheat, and the
results of digestion were studied when these were ejected or procured
by opening the animal's stomach. He also attached pieces of meat to
threads, which he would draw from the animal's stomach at fixed in- jc
tervals. He experimented upon himself by swallowing linen bags con- <
taining bread, meat and similar articles, examining the contents after '"1
they had been voided per anum. He procured gastric juice from
himself by producing vomiting on an empty stomach. He repeatedly
tested the action of gastric juice in vitro, keeping the tubes a uniform
temperature by retaining them in his arm pit, using the same food
covered by water as a control. He found that gastric juice acted 1^
more readily upon finely divided parts of food such as crushed grain )

22 SPALLANZANI

or bone which proved trituration only a preparation for solution, and
that it was no further a part of the digestive process.

He found that the gastric juice dissolved the food of animals into
a pultaceous mass or chyme. He observed that heat favored solution
and that in warm-blooded animals certain high temperature was
necessary for the chymification of foods. In Spallanzani's time putri-
faction was considered a form of fermentation. "There are three
kinds of fermentation, the vinous, the acetous, and the putrid." The
action of gastric juice was not putrid; in fact, it tended to arrest put-
refaction. Spallanzani was inclined to believe that the action of the
gastric juice was neither vinous nor acetous. Regarding the reaction
of gastric juice his conclusion was that it was neutral. He believed
that the acidity was due to an abnormality of the stomach contents,
inasmuch as the regurgitation of sour material from the stomach oc-
curred only when something had gone wrong. Spallanzani's failure to
t recognize the acidity of the gastric juice limited his further investiga-
tions. He could only conclude that the action of the gastric juice was
not fermentation, as fermentation was understood at the time.

It is interesting to note that the results of Reaumur and Spallan-
zani were confirmed by Stevens of Edinburgh, who likewise employed
Reaumur's methods of investigation. Stevens experimented on a
"man of weak understanding who gained a miserable livelihood by
swallowing stones for the amusement of the common people." The
man was made to swallow perforated silver spheres containing animal
and vegetable food, raw and cooked, which were examined when void-
ed some 48 hours later. Similar experiments were made on dogs, the
contents of the hollow spheres examined after opening the animal's
C stomach. Stevens concluded that digestion is not the effect of heat,
' trituration, putrification or fermentation alone, but of a powerful sol-
vent secreted by the glandular coat of the stomach.

Summary: Summing up the progress made in the physiology of
digestion during the seventeenth and eighteenth centuries, probably
'--^ no one is more entitled to an audience than Sir Michael Foster ; "Dur-
' , ing the two centuries the seventeenth and the eighteenth, physiologi-
\ cal inquiries, swayed now in one direction by views of chemical fer-
} mentation or effervescence, now in another direction by views of
( mechanical trituration, had come in the end to the conclusion that
digestion was in the main a process of solution of a peculiar charac-
ter, begun and chiefly carried out in the stomach, though assisted by
minor subsequent changes taking place along the intestines. They
who were under the influence of vitalistic doctrines, and these were
< perhaps the more numerous, held the change ^o be the commencement
of, to be the first step in the conversion of food into living flesh and
blood, and spoke of it as a change differing from ordinary chemical
change, without being able to define the exact characters. It was left
^ to the nineteenth century to throw new light on the nature of the
gastric changes and at the same time to show that what took place
in the stomach was not the whole digestion, but only the first of a
series of profound changes taking place along nearly the whole length
of the alimentary canal."

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CHAPTER III.

PHYSIOLOGY OF DIGESTION— WILLIAM BEAUMONT

We have traced the development of the physiology of alimenta-
tion from its crude beginnings, when debate waged as to whether
digestion consisted of mechanical trituration or whether it consisted
wholly of a fermentative process, to the time when some real light
began to be shed upon the subject by experiment with the gastric
secretion itself. No contribution to the subject of gastric digestion
has been of such moment as the work of William Beaumont on the
gastric secretion of the French Canadian, Alexis St. Martin. The story
of Beaumont's life and the circumstances surrounding his work con-
stitute one of the most fascinating chapters in the history of Ameri-
can Medicine.

As physicians, we have become familiar with the names of Beau-
mont and St. Martin early in our student career. They have become
inseparably associated with the study of gastric juice and its func-
tions. Standard works on physiology introduce the chapter on diges-
tion with such sentences as: "Gastric fistulae have been made in
human beings, either by accidental injury or by surgical operation.
The most celebrated case is that of Alexis St. Martin, a young Can-
adian who received a musket wound in the abdomen in 1822. Obser-
vations made upon him by Dr. Beaumont formed the starting point of
our correct knowledge of the physiology of the stomach and its secre-
tions."* ''The first fistula of a digestive gland to be the subject of a
thoroughly scientific investigation was one resulting from a gun shot
wound in the stomach of a Canadian hunter. As the consequence of
his accident, the hunter had all the rest of his life a stomach fistula
opening at the upper part of the abdomen, through which the interior
of the stomach could be observed and gastric juice could be obtained.
Beaumont collected a large number of important facts (1825-1833)
concerning the digestive process of the stomach and concerning the
movements of that organ." ''Beaumont's study of St. Martin's stom-
ach showed that in acute catarrh the mucous membrane is reddened
and swollen, less gastric juice is secreted, and mucous covers the sur-
face." Instances might be quoted almost ad infinitum of references
in medical literature to Beaumont's classic study of gastric digestion.

Beaumont; His Early Life: William Beaumont, the third child of
Samuel Beaumont, who had seen active service during Revolution days
prior to the Declaration of Independence, was born November 21st,
1785. There was nothing unusual in his childhood and youth. As he
grew to manhood his sympathies and political leanings were in accord
with those of his father, who was a staunch Democrat and patriot.
While no church record assures us that he was of the faith of his
parents, Congregationalist, his biographer asserts that when the roll
of the drum announced the approaching hour of worship he was
among those who slowly wended their way over the hills on foot or on

*Haliburton's Handbook of Physiology; Tigerstedt's Physiology; Osier's
Practice.

24 BEAUMONT

horseback to the old meeting house. Beaumont was blessed with such
rigorous parental discipline in youth that he explained his lapses in
church attendance in after life by the statement that during his
youth he had made up for a lifetime of church attendance. Further
than that he was a courageous and fearless boy, little is known of his
early life. It is said that he developed deafness, which became more
marked as he grew older, from standing near a cannon which was
being fired, simply to outwit playmates of his own age.

The beginning of last century found Beaumont a boy of fifteen
years. It was twenty-four years since the first birthday of the Ameri-
can Nation. Beaumont's youth was contemporaneous with one of the
most stirring epochs in world history. The United States was begin-
ning to assume an important place among the nations of the world.
Beaumont left home during the winter of 1806-7 with, we are told, an
outfit consisting of a horse and cutter, a barrel of cider, and a hundred
dollars of hard earned money. He traveled Northward, reaching in
the spring of 1807 the little village of Champlain, New York. He was
very favorably impressed with his surroundings and with the people,
who were mostly farmers, and whom he characterized as ''peaceful
and industrious in general." Here he established his "Lares and
Penates" and followed the career of schoolmaster. Coming from one
of the best New England schools his services were much in demand.
While teaching school and during the vacation he found time to de-
vote to medical studies. He had supplied himself with books borrowed
from Dr. Pomeroy of Burlington, Vt., which town was on his itinerary
to Champlain. Beaumont, as many since his day have done, made
teaching a stepping stone to the profession of m.edicine, and an excel-
lent experience it is for the aspiring savant. In 1810 he was apprentic-
ed to Dr. Chandler of St. Albans, Vt. He seems to have exhibited a
wise choice in the matter of preceptor. "Living under the same
roof," writes Dr. Myer, describing the medical education of the
times, "as was customary in the days of medical apprenticeship, the
preceptor could look after both the mind and morals of his pupils.
The fledgeling in return for the instruction received at the hands of
his master, not only compensated him for his trouble, but performed
many of the menial offices of a servant about the house and office.
It was he who prepared the powders, mixed the concoctions, made
pills, swept the office, kept the bottles clean, assisted in operations
and often through main force suppHed the place of the anaesthetic of
, today, in the amputation of limbs and other surgical procedures. He
( rode about with the doctor from house to house, profiting by his per-
\ sonal experience and jotting down on the pages of his note book and on
^r J the tablets of his memory the words of wisdom that fell from his mas-
i ter's lips. * * * He was taught the symptoms of disease, the
( crude methods of diagnosis, the art of prescription writing and the
'process of cupping and bleeding, considered so effective in its day."*
Medical books were rare and expensive, and fortunate was the
student who had access to them. Dissections were rarely performed,
owing largely to the fact of inadequate means of preserving cadavers.
Such were young Beaumont's opportunities.

Beaumont spent the two years of his apprenticeship with dili-
gence, studying the masters. He dissected whenever an opportunity
*Life and letters of Dr. William Beaumont, by Jesse S. Myer.

BEAUMONT ^

afforded, and never lost an opportunity to perform post-mortems. A
perusal of his case histories shows what a careful observer he was —
a qualification of the first importance in a physician. His diploma or
license to practice was granted the second Tuesday of June, 1812, by
the third Medical Society of the state of Vermont. It reads :

By the Third Medical Society of the State of Vermont as by law estab-
lished, William Beaumont having presented himself for examination on the anat-
omy of the human body, and the theory and practice of physic and surgery, and
being approved by our censors, the society willingly recommends him to the
world as a judicious and safe practitioner in the different avocations of the medi-
cal profession. In testimony whereof we have hereunto prefixed the signature
of our president and the seal of the society at the Medical Hall in Burlington on
the second Tuesday of June, A. D. 1812.

CASSIUS P. POMEROY, Secretary. JOHN POMERY, President

Assistant Army Surgeon: In September the same year Beau-
mont joined the army at Plattsburgh, as assistant surgeon under Gen-
eral Dearborn. His old preceptor Dr. Chandler had unsuccessfully
tried to dissuade him from the army service, advising him to settle
down to private practice. Apparently there is a destiny which shapes
our ends. Had he followed the advice of his old master, he would in
all probability have been among the thousands of good men who have
lived their lives through, leaving the world a little better than they
found it, and passed into the silent land, pass and leave no sign to in-
dicate that they have been. But Beaumont followed his own bent and
it was while acting as army surgeon that he made the momentous
discoveries which have placed him among the epoch-makers of medi-
cal history. It is significant to note that more than one army sur-
geon has performed service of an extraordinary nature to medical
science. Fom the times when Machaon and Podilirius rendered aid to
the Greek hosts at ancient Troy to the days of Ambrose Pare, the
army surgeon has been identified with medical progress. A name
honored within recent years in the French service is that of Laveran,
who during his tour of duty in Algeria did a work in connection with
malaria which made possible the work of Sir Ronald Ross, of the In-
dian medical service, and his associates of more recent times. The
spectacle of the Panama Canal and its construction were made pos-
sible by the United States Army medical service. In the British Army
medical service are such names as Sir David Bruce, whose investiga-
tions led to the extermination of Malta fever.

Beaumont's Diary: Beaumont left a diary which is an interest-^^^*
ing description by one on the firing line, of the stormy times of 1812. ^
This graphic account of events of the war by an eye-witness is
reproduced in Dr. Meyer's book. Beaumont was present August, 1814,
at the battle of Plattsburgh, where General Macomb defeated the Brit-
ish under General Provost. The Treaty of Ghent ratified in February,
1815, closed the war. Soon after the close of the war of 1812 Beau-
mont tendered his resignation and in partnership with a Dr. Senter
opened a store in the town of Plattsburgh, which store contained "a
general assortment of drugs, medicines, groceries, dye woods, etc., of
the first quality and choicest selection which they calculate to sell on
liberal terms for cash or approved credit." So runs the advertisement

26 PATHFINDERS OF PHYSIOLOGY

in the local newspaper. In the footnote of the advertisement it is
stated that "Medicines will be put up with accuracy and care." In
December of 1816 Beaumont sold out and afterwards confined himself
entirely to the practice of his profession. He was commissioned by
President Monroe in 1820 and re-entered the military service, when
he was ordered to Fort Mackinac on the Northwestern frontier. He
describes his journey in detail in his diary. His course lay along the
southern shore of Lake Erie to the Detroit river, where he passed
Fort Maiden, near the Canadian town of Amherstburg, opposite Bois
Blanc island. He describes the fort at Detroit as a "regular work of
an oblong figure covering about an acre of graceful slopes." The
parapets are about 20 ft. in height, built of earth and sodded, with
four bastions. The whole surrounded with palisades, a deep ditch
and glacis. It stands immediately back of the town and has strength
to withstand a siege. The Detroit postoffice, corner of Fort and Shelby
streets, stands upon the ground at one time occupied by the above
mentioned fortification. A bronze tablet at the south entrance of the
postoffice gives in brief the vicissitudes of the old fort.

He speaks of crossing over to Sandwich, then a small French vil-
lage. There is no mention of the route again until he reaches Fort
Michilimackinac, which is described as handsomely situated on the
southeast side of the island of this name, on a bluff rising from 100
to 200 feet from the water, almost perpendicular in many places, ex-
tending about half way around the island. The word "Michilimacki-
nac" means "turtle" from the resemblance of Mackinac island on be-
ing approached.

The following entries in his diary throw considerable light on the
character of the man himself.

Sept. 9, 1820. Commenced a diary of conduct on Dr. Franklin's plan, for ob-
taining moral perfection." (Benjamin Franklin appears to have been a favorite
with Beaumont, for he elsewhere quotes him at length.) ^'Reading Shakespeare
today I judged the following extracts worthy of copying; 'Love all, trust few,
do wrong to none; be able for thine enemy rather in power than use; and keep
thy friend under thy life's key; be checked for silence, but more taxed for speech.

"10th. Rose at six o'clock. Visited my patients in village and discharged
garrison duty before 9 a. m. Settled my hospital account, perused scriptures
and Pope's Essay on Man till evening."

Beaumont's diary is an interesting narrative of the times, written
by a keen and practical observer.

The Psychological Moment: Late in the spring of 1822 occurred
the event which made the name of William Beaumont famous in the
annals of medicine. Indians and voyageurs had returned to Mackinac
with the results of the winter's hunting. A strange medley of hu-
manity had gathered at the Amercan Fur Company's trading post.
On the 6th of June a gun was accidentally discharged, its contents
entering the upper abdomen of a young voyageur, leaving a cavity
which would have admitted a man's fist. According to an eye-wit-
ness Alexis St. Martin, for that was his name, fell, as every one sup-
posed, dead. Dr. Beaumont, surgeon of the fort, was called, and ar-
rived shortly after the accident. Shot and pieces of clothing were
extracted and the wound dressed. The surgeon then left with the re-

BEAUMONT 27

mark that the man couldn't live 36 hours. The doctor called again in
the course of two or three hours and found the patient better than he
had anticipated. The patient was removed to the fort hospital where
he eventually recovered, leaving, however, a permanent gastric fistula.
Beaumont's own account of the accident is told in the introduction
to his work on "Experiments and Observations of Gastric Juice."

"Alexis St. Martin, who is the subject of these experiments, was a Canadian
of French descent at the above mentioned time (1822) about 18 years of age, of
good constitution, robust and healthy. He had been engaged in the service of
the American Fur Company as a voyager and was accidentally wounded by the
discharge of his musket on the 6th of June; the charge, consisting of powder and
duck-shot, was received in the left side of the youth, he being at a distance of not
more than one yard from the muzzle of the gun. The contents entered posteriorly
and in an oblique direction, forward and inward, literally blowing off integuments
and muscles of the size of a man's hand, fracturing and carrying away the anter-
ior half of the sixth rib, lacerating the lower portion of the left lung, the dia-
phragm and perforating the stomach. The whole mass of materials forced from
the musket, together with fragments of clothing and pieces of fractured ribs,
were driven into the muscles and cavity of the chest. I saw him in 25 or 30 min-
utes after the accident occurred, and on examination found a portion of the lung
as large as a turkey's egg protruding through the external wound, lacerated and
burned; and immediately below this another protrusion which, on further ex-
amination, proved to be a portion of the stomach lacerated through all its coats
and pouring out the food he had taken for his breakfast through an orifice large
enough to admit the forefinger."

Beaumont's hospital and bedside notes give a complete history
of the case.

Being destitute and without friends or relatives, Alexis St. Mar-
tin became a pauper on the town of Mackinac. It was at last decided
to ship him to his native town, Montreal, nearly one thousand miles
away. Beaumont, however, rescued him from misery and inevitable
death by taking him into his own family. "During this time, says
his benefactor, I nursed him, fed him, clothed him, lodged him and
furnished him with every comfort and dressed his wounds daily and
for the most part twice a day." It should be realized that Beau-
mont endeavored to close the wound; that when all other means
failed he suggested incising the edges of the wound and, "bringing
them together by sutures, an operation to which the patient would
not submit."

Not until three years after the accident did the idea of perform-
ing a number of experiments appear to occur to the mind of Beau-
mont. In 1825 he began to realize the importance of this case which
had fallen to his care, when it occurred to him what a great service
to humanity might result from this accident. About this time Beau-
mont describes the situation as follows:

'He (St. Martin) will drink a quart of water or eat a dish of soup and then
by removing the dressings I frequently find the stomach inverted to the size
and about the shape of a half-blown rose, yet he complains of no pain, and It
will return itself or is easily reduced by gentle pressure. When he lies on the
opposite side I can look directly into the cavity of the stomach and almost see
the processes of digestion. I have frequently suspended flesh, raw and wasted,
and other substances into the perforation to ascertain the length of time re-

28 PATHFINDERS OF PHYSIOLOGY

quired to digest each, and at one time used a tent of raw beef instead of lint
to stop the orifice, and found that in less than five hours it was completely
digested off as smooth ai)d as even as if it had been cut with a knife."

Then his resolve to make use of the case as a means of study-
ing gastric digestion takes shape as follows:
P "This case affords an excellent opportunity for experimenting on the gastric

\ fluid and process of digestion. It would give no pain nor cause the least un-
p \ easiness to extract a gill of fluid every two or three days for it frequently flows
I out spontaneously in considerable quantities. Various kinds of digestible sub-
/ stances might be introduced into the stomach and then easily examined during
/ the whole process of digestion. I may, therefore, be able hereafter to give some
I interesting experiments on these subjects."

Recognition of Michigan Medical Society: The Medical Society
of the territory of Michigan was the first body to recognize the work
of William Beaumont. The following letter dated from Detroit an-
nounced his election as an honorary member of the Michigan Territor-
ial Medical Society.

"Dr. William Beaumont, United States Army, Michilimackinac.

Detroit, March 3, 1825.

"Sir: — It is with much pleasure that I transmit to you as an extract from
the minutes of the medical society of this territory at a meeting held at the home
©£ Capt. Woodworth in the City of Detroit on Monday, 7th ultimo; Dr. William
Beaumont, of the United States Army, duly proposed by Dr. Pitcher and unani-
mously elected by ballot an honorary member of this society.'

"Whereupon it was ordered that the secretary be directed to inform Dr.
Beaumont of his election as aforesaid.
"I remain, sir, with much respect,

*'Your most obedient servant,

"JOHN S. WHITING,
"Secretary of the Medical Society of the Territory of Michigan."

The first experiments were carried on at Mackinac and were
continued at Fort Niagara, to which place Beaumont was removed.
While on a visit to Burlington, Vt., as one of his master's household,
A Alexis, whose interest in science had long ago reached the vanishing
J point, ran away and was lost to his benefactor for some time. This
ungrateful act on the part of the French-Canadian proved a sore dis-
appointment to our ''Backwoods physiologist." His experiments up
to this time were to estimate the length of time required for the
digestion of certain kinds of food, which were suspended in the stom-
ach by means of silk threads and withdrawn from time to time to note
5 the changes in the substances. He found that food would digest more
I quickly in the stomach than when mixed with gastric juice in vitro.
Four years after St. Martin's unceremonious departure, Beau-
mont got in communication with him. In the meantime Alexis had
married and became the father of two children. The doctor took
him, his wife and two children into his own home, where Alexis did
duty as a common servant when not employed for purposes of experi-
C mentation. Beaumont's laboratory equipment consisted of a thermo-
(^ meter, a few open mouthed vials and a sand bag. His observations

BEAUMONT 29

were made with a true spirit of inquiry and with no particular hypo- ,
thesis to support. Fifty-six experiments were made between Dec.
6th, 1829, and April 9th, 1831. Alexis, with his wife and family, were
permitted to return home to Quebec on the promise to appear when
again wanted. Beaumont had felt that he had accomplished about all
he was able in his researches on gastric digestion, and he longed to
go to Europe a year and take St. Martin with him, that the work
might be pursued farther by more competent physiologic chemists.
The brevity of his furlough precluded the idea of going abroad and
instead he remained in Washington with Alexis where he found his
surroundings very congenial. Access to the works of European
physiologists in the library and recognition from many of the promi-
nent men at the capital made his sojourn pleasant. '>>. ^

Between Dec. 1st., 1832 and March 1st, 1833, we find recorded (f ^
116 experiments, some in confirmation of what had been done before. [ '
He tested the temperature of the stomach when full, when fasting, \
when exercising, when resting, also the length of time required to \
digest various food substances. He also experimented to disprove /
the old theory of maceration or mechanical trituration.

Seeks Assistance of Two Leading Scientists: In 1833 Beaumont
sought the assistance of two of the leading scientific men of the
United States, Robley Dunglinson, professor of physiology. Univer-
sity of Virginia, and Benjamin Silliman, professor of chemistry at
Yale. Thanks to Beaumont's painstaking and methodical nature, the
correspondence between the two and himself had been carefully pre-
served, and it constitutes an excellent account of the physiology of
the period. A sample of gastric juice from St. Martin's stomach was
sent Dunglinson for analysis with the request to convey to the giver
the results and to refrain from publishing anything that would antici-
pate the labors of Beaumont himself. He is assured that the profes-
sor has but one desire in the prosecution of his profession, by teaching
and practice to benefit his fellow men, which could always be done
with due credit without forestalling his coadjutors in the field of
science, or arrogating to himself merit to which he might be but sec-
ondarily entitled. Dunglinson found the sample of gastric juice to
contain "free muriatic and acetic acid and phosphates and murates .
with bases of potassa, soda, magnesia and lime and animal matter
soluble in cold but not in hot water."

Professor Silliman, to whom a bottle of gastric juice was also
submitted, suggested that a sample be sent to Professor Berzelius, of
Stockholm, Sweden, "as the man of all others best qualified to investi-
gate a subject of such deep interest to mankind." Accordingly a
bottle of the digestive fluid was packed for shipment. Beaumont's
disappointment may be imagined when it was known that the parcel
was delayed over two and a half months. This he learned about the
time he was patiently awaiting the results of the Swedish professor's
investigations. In the meantime Beaumont had received a letter from
Professor Silliman enclosing an abstract of a portion of a system of
chemistry by Berzelius, important as presenting a clear idea of the ^
knowledge of the physiology of digestion at that time (1833). The \
communication states, among other things, that Prout, Tiedeman and j
Gmelin gave the best notions on the subject of gastric juice and ex- ^

30 PATHFINDERS OF PHYSIOLOGY

plained the contradictory statements of other authors; at one time it
was said to be very fluid clear and neutral in reaction ; then alkaline,
then acid. Prout in 1824 declared the gastric juice to contain free
hydrochloric or muriatic acid, the result of an experiment made on
the contents of the stomach of an animal killed soon after eating.
Gmelin and Tiedeman also established the presence of free hydroch-
loric acid. The fluid of the empty stomach was found to be slightly
acid, sometimes neutral and the acidity was in proportion to the quan-
tity, becoming very acid when food had been swallowed. According
to Gmelin and Tiedeman, the salts of gastric juice were principally
sodium chloride and potassium chloride in small quantities, hydro-
^ chlorate of ammonia and a little sulphate of potassium. The com-
i munication concludes with the assertion that "no organ for the special
[secretion of the gastric juice has yet been discovered."

Berzelius' Reply Disappointing: Through Professor Silliman,
Beaumont eventually heard from Berzelius, whose letter was dated
July, 1834. The communication upon which such great expectations
were placed was wholly disappointing. It was in the main an apology
for the writer's inability to work with the gastric fluid with prospects
of results of any value, owing to the time which had elapsed since
its secretion and its arrival at his laboratory, to the possible alteration
on account of summer heat, and to the inadequate quantity received.

Nothing but the utmost zeal and love for the work could account
for the persistence with which Beaumont pursued his researches. He
felt not only the handicap of inadequate resources and facilities for
experimentation, but St. Martin was a source of canstant annoyance
to him. He would leave his master and benefactor, often absent for
several years, when by overtures in the shape of money he would be
prevailed upon to return and furnish the precious fluid for his mas-
ter's investigation. Beaumont's lot was cast at a time when it was
difficult, almost impossible, to obtain government grants for the pro-
motion of education. His work, therefore, has been accomplished al-
most entirely at his own expense.

Attains Fame Through His Stomach: St. Martin lived the life
of the French Canadian habitant mostly in poverty, though physically
he was, the larger part of his life, in good condition. Nine years after
his notable accident, we are told, he took his family in an open canoe
via the Mississippi, passing St. Louis, ascended the Ohio River, then
crossed the state of Ohio to the lakes and descended the Erie and
Ontario and the River St. Lawrence to Montreal, the trip consuming
the interval from March to June. He was able to engage in manual
labor requiring considerable strength and endurance. Perhaps his ex-
treme poverty is due to lack of thrift and to intemperance, for we are
told that he indulged immoderately in the "glass that cheers."

The longevity of the habitant is evidenced in St. Martin, for
he lived twenty-eight years after the death of Beaumont. St. Mar-
tin's death occurred in his eighty-third year. Sir William Osier, at
the time, (1880) a resident of Montreal, reading of his death, wrote
the local physician and parish priest urging them to secure for him
the privilege of an autopsy, and at the same time offering a goodly
sum for the stomach, which he intended to place in the Army Medi-
cal Museum at Washington, but his entreaties were of no avail, the

BEAUMONT 31

body was interred eight feet below the surface of the ground, after
being detained at home much longer than the usual period, so that
decomposition setting in, might baffle the doctors, and prevent any
attempts at resurrection.

Beaumont Resigned From Army: William Beaumont resigned
his position as army surgeon in 1839. He continued, however, to
attend the families of the officers at St. Louis, where he made his
home. Owing to the distance from St. Louis of his successor, who
was stationed ten miles away, he presented an account to the War
Department for professional services covering a period of a few
months, which services he conceded "irregular and informal," but
"correct and just." On receipt of his account the surgeon-general
threatened either to ignore the bill or to deduct the amount from
the salary of Beaumont's successor. The manner in which Beau-
mont received the threat showed the independent nature of the man.
He declared the surgeon-general's view at "absurd opinion, con-
tracted view, narrow-minded vindictive spirit and petty tyrannical
disposition," of the "weak, waspish and wilful head of a medical de-
partment," and congratulated himself over having the "privilege of
detesting a man, the motives and the mind from which such egregius
folly, parsimony and injustice could emanate and be promulgated."
The Surgeon-General was, however, unyielding, and Beaumont's claims
were unrecognized.

Though severed from the War Department, he still had a very
lucrative practice, and what is above any monetary consideration, de-
voted friends, and was very happy in his domestic relations. The
following paragraph quoted in Dr. Myer's Life and Letters of Beaa-
mont gives a splendid estimate of his character:

"Dr. Beaumont possessed great firmness and determination of purpose.
Difficulties which would have discouraged most men, he never allowed to turn
him from his course. These he did not attempt to evade but to meet and overcome.
He posseTssed more than any man I ever knew, a knowledge almost intuitive of hu-
man character. You might have introduced him to 20 different persons in a
day, all strangers to him, and he would have given you an accurate estimate of
the character of each, his peculiar traits, disposition, etc. He was gifted with
strong natural powers which, working upon an extensive experience in life, re-
sulted in a species of natural sagacity, which I suppose was something peculiar
to him not to be attained by any course of study. His temperament was ardent
but never got the better of his instructed and disciplined judgment, and when-
ever or however employed, he always adopted the most judicious means of ob-
taining ends that were always honored. In the sick room he was a model
of patience and kindness; his intuitive perceptions guiding a pure benevolence
never failed to inspire confidence. Thus, he belonged to that class of physicians
whose very presence affords nature a sensible relief."

He died on April 25th, 1853. His death was considered the re-
sult of injuries he received by slipping on icy steps while making a
professional visit. What a satisfaction such a life must be, and the
resignation with which one might approach the infirmities of old age
and one's final destiny. And indeed a few months before the end he
breathed forth this beautifuly symphony:

32 PATHFINDERS OF PHYSIOLOGY

"Myself and wife, not unlike John Anderson my Jo, have climbed the hill o'
life togither, and mony a canty day we've had wi' ane anither. But now we
maun totter down life's ebbing wane in peaceful quiet ease and compitence, with
just so much selfishness and social sympathy as to be satisfied with ourselves, our
children and friends, caring little for the formalities, follies and fashions of the
present age. * * * Come when it may, we only ask God's blessing on our
frosted brows and hand in hand we will go to sleep together."

DR. BEAUMONT'S BOOK.

I am fortunate in having before me an original copy of Dr. Beau-
mont's work. The title page bears the following description: "Ex-
periments and Observations on the Gastric Juice and the Physiology
of Digestion, by William Beaumont, M. D., Surgeon in the United
States Army. Plattsburg. Printed by F. P. Allen, 1833." The vol-
ume is dedicated to Joseph Lovell, M. D., Surgeon General of the
United States Army. The work comprises 2B0 pages, 122 of which
deal with "Preliminary Remarks on the Physiology of Digestion."
The remainder deals with Experiments and Observations on the Stom-
ach of Alexis St. Martin. The first part is divided into seven sections,
as follows: 1st, Of Ailment; Section two of Hunger and Thirst;
Section three of Satisfaction and Satiety; Section four of
Mastication, Insalivation and Deglutition; Section five of Digestion
by Gastric Juice; Section six of the Appearance of the Villous Coat
and of Motions of the Stomach; Section seven of Chylification and
Uses of the Bile and Pancreatic Juice. There are three illustrations,
consisting of crude wood cuts of the gastric fistulse. The typograph-
ical appearance of the work should be onsidered creditable consider-
ing the printing art at the time. The conclusion of the second part of
the work contains 51 inferences made from the foregoing experiments
and observations. Of these I shall quote a few :

That digestion is facilitated by minuteness of division and tenderness of fibre
and retarded by the opposite qualities.

That the quantity of food generally taken is more than the wants of the
system require, and that excess, if persevered in, generally produces not only
functional aberration but disease of the coats of the stomach.

That bulk as well as nutriment is necessary to the articles of diet.

That oily food is difficult of digestion, though it contains a large proportion
of the nutrient principles

That stimulating condiments are Injurious to the healthy stomach.

That the use of ardent spirits always produces disease of the stomach if
persevered in.

That the agent of chymification is the gastric juice, which acts as a solvent
of food and alters its properties.

That the action of gastric juice is facilitated by the warmth and motions of
the stomach.

That it coagulates albumin and afterwards dissolves the coagulum.
ciples.

That the gastric juice is secreted from vessels distinct from the mucous
follicles.

That bile is not ordinarily found in the stomach and is not commonly neces-
sary for the digestion of food, but assists in the digestion of oily foods.

:yj^

BEAUMONT 33

That the inner coat of the stomach is of pale pink color, varying in its hues
according to its full or empty state.

That the motions of the stomach produce a constant churning of its contents
and admixture of the food and gastric juice.

That these motions are in two directions, transverse and longitudinal.

Beaumont failed, however, to ascribe any digestive function to
the saliva. He maintained that food finely divided placed directly
into the stomach was as completely digested as that which entered
by the oesophageal route.

When he began his work the status of the physiology of
digestion had been very well described by William Hunter; "some
physiologists will have it that the stomach is a mill ; others that it is
a fermenting vat ; other again that it is a stew pan ; but in my view
of the matter it is neither a mill, a fermenting vat nor a stew pan, but
a stomach, gentlemen, a stomach." When William Beaumont com-
pleted his labors there was a marked advance in knowledge of the
digestive process. Among the most important results of his work
was his complete and accurate description of the gastric juice, whi
has been quoted in so many text books since his day.

"Pure gastric juice when taken directly out of the stomach of a healthy adult,
unmixed with any other fluid, save a portion of the mucus of the stomach with
which it is most commonly, perhaps always combined, is a clear, transparent fluid;
Inodorous; a little saltish, and perceptibly acid. Its taste, when applied to the
tongue, is similar to mucilaginous water, slightly acidulated with muriatic
acid. It is readily diffusible in water, wine or spirits; slightly effervescent with
alkalies, and is an effectual solvent of the materia alimentaria; it possesses the
property of coagulating albumin in an imminent degree; it is a powerful anti-
septic, checking the putrefaction in meat; and effectually restorative of healthy
action when applied to old foetid sores and foul ulcerating surfaces."

His work confirmed the observation of Prout, that the acid con'
tents of the gastric secretion was hydrochloric. He recognized the
fact that the elements of the gastric juice and the mucus of the stom-
ach were a separate secretion. He established by direct observation
the marked influence of mental states on the secretion of gastric juice
and on digestion. His was the first comprehensive and thorough
study of the motions of the stomach ; and to quote Osier : "His study
of the digestibility of different articles of diet in the stomach remains
today one of the most important contributions ever made to practical
dietetics."

A German edition of the work was issued in 1834. In 1838 Sir
Andrew Combe, an eminent English physician, published an English
edition of the work, so as to give it greater publicity in the British
Isles. Probably no fairer or more impartial estimate of the value
of Beaumont's contribution to science has been made than that of
Sir Andrew in his preface to the British edition. Answering the
objection that Beaumont had made no original discovery in the phys-
ology of digestion, this advocate claims that by "separating the truth
clearly and unequivocally from the numerous errors of fact and opin-
ion with which it was mixed up, and thus converting into certainties
points of doctrine in regard to which positive proof were previously
inaccessible, he has given to what was doubtful or imperfectly known

V

S4 PATHFINDERS OF PHYSIOLOGY

a fixed and positive value which it never had before, and which, being
once obtained, goes far to furnish us with a clear connected and con-
sistent view of the general process and laws of digestion."

CLAUDE BERNARD, PHYSIOLOGIST.
1813—1878

CHAPTER IV.

GLYCOGENIC FUNCTION OF THE LIVER— VASOMOTOR
NERVES— CLAUDE BERNARD.

"For a man to be an investigator of the first order two gifts are perequisite
it is not merely necessary to possess a well-ordered and what we may term a
philosophic imagination, to possess a mind that is capable of balancing phenomena,
seeing their relationship and deducing problems that have to be solved and the way
in which to solve them; there must be something more, namely, a mechanical
ability, a love for technique, and a capacity to construct and manipulate the ap-
propriate Instruments. This is particularly necessary in connection with physio-
logical resarch." — ^Adaml

The real life of every notable character lies in the story of his
achievement, rather than in how he passed his days. Human interest,
however, loves to dwell on the details of how he moved among his
fellowmen and the vicissitudes that befel him on his path through
life. Often in the lives of our greatest men these details which con-
stitute the human touches have not been recorded. Not everji John-
son has his Boswell, and we must content ourselves with the frag-
mentary data that have been preserved. Such has been the fate of
Claude Bernard, the first centenary of whose birth is now the sub-
ject of commemoration.

Early Life and Education — Let me give a brief summary of his
life. He was bom on July 12th, 1813, of humble parentage ; his father
owned a small farm at St. Julien, near Lyons, France. The vintage of
the little estate which was situated in the wine district of France, pro-
vided the family revenue. The property eventually came into the
hands of the son, who spent his summers there within view, on clear
days, of the white summits of the Alps. Bernard received his early
education at his native village and afterwards at Lyons. His educa-
tion was, however, cut short by necessity, which turned him to prac-
tical pharmacy as a means of earning a living. The young man pos-
sessed that "fine frenzy'* which makes "the lunatic, the lover and the
poet" of "imagination all compact,*' and was on the point of giving up
the calling which had engaged his attention for two years, for litera-
ture. His literary aspirations drew him towards the dramatic art, and
it is hard to predict what the future physiologist might have given te
the world had not the divine flame been smothered by a more prosaic
career of investigator. He was the author of a comedy, "The Rose of
the Rhone," which had met with a certain amount of success. But des-
tiny had reserved Bernard for another and very different calling.
He submitted his work to the great French critic, St. Marc Girardin,
who, while recognizing its merit, advised the young aspirant to lit-
erary fame to pursue a more lucrative calling, to engage in some
pursuit in which he could earn his bread and to court the Muses only
in his leisure moments. "You have studied pharmacy," said the
critic, "study medicine; you will thereby much more surely gain a
livelihood." Bernard followed this advice with heart and soul, de-

1

Se PATHFINDERS OF PHYSIOLOGY

fraying his expenses by tutorage. The literary longings began to
fade as the young savant waxed warm with his medical studies.
Anatomy and physiology claimed the greater portion of his attention
and energy. His remarkable manual dexterity, in which he was par-
ticularly fortunate, rendered his dissections of singular completeness
and value. The chaotic condition of physiology of the time (1840)
served to awaken in his mind a desire to solve problems by direct
^ experimental appeal to nature. He was one of the first to ^employ
animal experimentation, or vivisection. In 1841 he attracted the
attention of the great IMajendie, then the leading physiologist of
France, also Professor of Medicine in the College of France. Ma-
jendie is described as being in manner abrupt and even rough and
rude. At first he took little notice of Bernard, his new interne, but
was soon impressed with the young man*s dexterity and skill. One
day while Bernard was busy at his dissecting, Majendie blurted out:
"I say, you, there. I take you as my preparateur at the College of
France." And it was not long before the master had occasion to say
in his gruff way as he left the class-room: "You are a better man
than I am." Bernard's career as physiologist may be said to date
from this appointment in 1841.

Claude Bernard was of a retiring, silent nature, difficult to under-
stand and often misunderstood. Michael Foster described him as
"tall in stature, with a fine presence and a noble head, the eyes full
at once of thought and kindness ; he drew the look of observers upon
him wherever he appeared. As he walked the streets passers-by
might be heard to say, *I wonder who that is ; he must be some dis-
tinguished man.* "

The Productive Period — Bernard had shown the precious metal of
his genius before he was far on in his twenties. Nearly all of his great
achievements were accomplished during the period of his life which
ended with 1860. The essential results of his two greatest discoveries,
the glycogenic function of the liver and the vaso motor nerves were
gained prior to 1850, before he was 37 years old. He is illustrative of
Osier's declaration that the world's best and most important work
was mainly done by young men, for further example: Mor-
gagni's germinal idea, which made him the father of modem path-
ology, came to him when he was scarcely twenty; Auenbrugger
began his work upon percussion when he was under twenty-five ; Laen-
nec undertook the problem of constructing a system of auscultation in
his early twenties and published his book when he was not yet thirty-
five.

All significant work in medicine has had its basis in observation,
not theory. Men have been prone to theorize too much and to observe
too little. For two thousand years the learned men of Europe de-
bated as to whether this or that place was the site of ancient Troy, or
whether there ever was such a place at all. It remained, however, for
a retired man of business, Schliemann, to decide the question. He
said, "Let us go and see," and, at the expense of a few thousand
pounds, he went and found Troy and Mycenae and revealed or dis-
covered the whole matter — "The most tremendous and picturesque tri-
umph of the scientific method over mere talk and pretended historic
learning," says Ray Lancaster, "which has ever been since human

GLYCOGENIC FUNCTION OF LIVER 37

record has existed." Emerson has said: "I am impressed with the
fact that the greatest thing a human soul ever does in this world is
to see something and tell what it saw in a plain way. Hundreds of
people can talk for one who can think, but thousands can think for
one who can see. To see clearly is poetry, philosophy and religion all
in one." And we might add, that rare quality of mind which enables
its possessor to_see clearly is the sine quo non of the true scientist.

Gastric Digestion — Among Bemard*s earliest investigations was
that of gastric digestion. It was important chiefly as a prelude to the
momentous discoveries he afterwards made. He was the first to in-)
quire into the differences to be found between the digestive appara-i
tuses and functions of plant-eating and meat-eating animals — ^between \
the herbivora and camivora. The former thoroughly masticate their
food, while the latter bolt theirs. This instinct is explained by the fact
that the food of the plant-eating animal contains a relatively large
amount of starch, requiring thorough admixture of saliva as an aid to
its digestion. Those animals subsisting on meat-protein do not require
the aid of the saliva, which accounts for the rapidity with which they
devour their food. From this Bernard turned to study the function
of the pancreaticjuice. Up to this time the pancreas had been passed.^
over in silSh'ce by the physiologists of the day. He demonstrated its (
three-fold action: **He showed that it, on the one hand, emulsified, \
and, on the other hand split up into fatty acids and glycerine, the }
neutral fats that are discharged from the stomach into the duodenum.
He proved it had a powerful action on starch, converting it into
sugar." The study of the_a£iJiQjajpf ..the^ancreatic juice upon proteins
begun by Bernard wasfcontinued by ffiEiiij, his pupil, who investi-
gated the action of extracts of the" giandr Pancreatic juice as se-
creted does not possess proteolytic powers. This change under normal
conditions is brought about by the activating substance, enteroki-
nase, contained in the succus entericus producing as soon as the
pancreatic juice enters the gut, the change from the inert typsinogen
to trypsin, thus acquiring an activity over proteins superior to that
of any other digestive juice. (Starling). Up to Bernard's time the
principal role of digestion had been confined to the gastric juice. With! j^
his discoveries it became clear that the action of the gastric juice on y
the food in the stomach was simply preliminary to intestinal diges- i
tion and that the chief work in the preparation of the food for ab- J
sorption was accomplished by the pancreatic juice.

Discovers Glycogenic Function of Liver — Important as were his
numerous contributions to our knowledge of physiology, Claude Ber-
nard is probably best known as discoverer of the glycogenic function
of the liver. The story of his discovery is interesting and well worth
relating. The dominant opinion among physiologists when Bernard be-
gan his work was to the effect that animals and plants presented a
chemical contrast to each other. The plant built up such organic com-
pounds as fats, carbohydrates and proteins out of inorganic elements ;
the animal feeding on the plant received these organic compounds into
its body resolving them into inorganic substances, at the same time
using that resolution for the needs of life. While the animal modified
vegetable proteins, carbohydrates and fats so as to give them an ani-
mal character, it never made anything new. It was maintained that

38 PATHFINDER OF PHYSIOLOGY

the animal body never manufactured any of these three compounds,
that all or any of them present in the animal body had been taken into
it with its food.

Such was the current belief among physiologists of France at

^ the beginning of the fourth decade of last century. The first heresy

was uttered by Liebig who proved that the fat accumulated in the

bodies of fattened geese exceeded greatly the quantity of fat in the

intake of food, and furthermore that when a cow was fattened, the

excreta during the fattening period contained as much fat as the food

taken. At this time Bernard undertook his researches on the physi-

r* ology of sugar. His first discovery was that cane sugar acted upQn

\ by the gastric juice was changed into dextrose (glucose). It was his

2 intention to study the tliree great claisses of foods, but he found it nec-

j essary to confine his attention to the carbohydrates owing to the fas-

/ cinating problems suggested by diabetes. He set about to discover the

Vcause of the excess of sugar in diabetes with the hope of finding a

remedy for the disease.

Having previously satisfied himself that no dextrose was present

; in the alimentary canal, or in the portal blood, Bernard fed a dog on

meat only; killing the animal at the height of digestion he found to

his great astonishment the blood loaded with dextrose.

rv "Why!" said he, "if I have made no mistakes I have in this ex-

S periment come upon the production of sugar; the liver produces sugar.

^If the result I have got is confirmed on repetition of the experiment,

the liver is the sugar-producing tissue. It manufactures sugar out

of something that is not sugar, and within it lies the secret of dia-

i betes. This is a big thing of which I have got hold. I must make

// sure that I have made no mistake in the experiment, and then push

^/ forward as far as possible the lead thus given me."

C Bernard's results were confirmed by numerous experiments. He

\ determined that the sugar in question was dextrose, responding to all

' j the tests for dextrose. He also discovered that while this hepatic

/ sugar did not come direct from the food, it was influenced in regard

J to its quantity by the nature of the food. Starling, however, main-

/ tains that in some animals, the carnivora, tne liver can continue to

/ supply sugar to the blood on a diet which includes only proteins and

I fats. Yan Noorden explains the fact that proteins yield sugar, by

V the presence of a carbohydrate group in the protein molecule, which is

split off during pepsin-hydrochloric acid digestion.

Bernard eventually came to the conclusion that sugar was not

formed immediately from the elements whatever they might be which

the blood brought to the liver, but from some substance existing in

the liver tissue which was capable of being converted into sugar...J[]j

1857 he announced^iQ, the -scientific world the discDverv of glycogen.

* Though he made known each step in his discoveries which extended

•p over a number of years, he had the satisfaction of telling the whole

j^ story in his own writings, never having experienced the humiliation

which is sometimes the lot of pioneers, in seeing their leading con-

,^ ceptions worked out by other minds. To ^uote liia^,IiiiQgxaph^^^lr

C Michael Poster, "Bernard in the matter of glycogen not onlylaidThe

J first stone but left a house so nearly finished that other men have been

^ able to add but little."

VASO-MOTOR NERVES 39

"No less pregnant of future discoveries," says this biographer,,
"was the idea suggested by this newly found-out action of the hepatic)
tissue, the idea happily formulated by Bernard aa /internal secretion.''
No part of physiology is at the present day being more fruitfully
studied than that which deals with the changes the blood undergoes
as it sweeps through the several tissues." The study of these in-
ternal secretions constitutes a path of inquiry which has within re-
cent years been pursued with conspicuous success.

To Bernard we owe the discovery of the remarkable fact that
temporary diabetes may be caused by puncture of the fourth ven-
tricle. This glycosuria was formerly attributed to direct stimula-
tion of the liver through its nervous connections. It has been found.
however, that if the left adrenal is cut off from the left sympathetic
nerve, no sugar appears in the urine after the medulla has been punc-
tured, and it is now believed that the stimulus is transmitted by the
left sympathetic nerve to the left adrenal, whence it is passed to the
right adrenal by the connecting nerves. As a consequence of the
medullary puncture the adrenals secrete more actively and the in-
creased flow of the adrenal secretion in its turn brings about an
excessive output of sugar by the liver.* A number of toxic influences
possibly act in the same way, the glycosuria to which they give rise
being partly the result of the action they exert on the diabetic center
in the medulla, and partly an effect of their stimulating action on the
sympathetic nerves, or on the adrenals directly, thus, in any case,
causing hyperfunction of the chromaffin system, with consequent
overproduction of sugar by the liver.

Discovery of Vaso-Motor Nerves — Next in importance to the dis-
covery of glycogen was Bernard's discovery of the vaso-motor
nerves. "To Claude Bernard," says Sir Michael Foster, "we owe the
foundations of the vaso-motor system. He made known to us the ex-
istence of vaso-motor nerves and he also made known to us that vaso-
motor nerves are of two kinds, vaso-constrictor and vaso-dilator — ^the
two fundamental facts of vaso-motor physiology." The importance
of this discovery can hardly be over-estimated when we consider that
there is scarcely a physiological problem of any magnitude which does
not sooner or later involve vaso-motor questions. The vaso-motor
nerves presiding as they do over the contraction and dilation of the
walls of the blood vessels, assume an important role in such functions
as gastric digestion, blood pressure, heat processes, blushing and
various other congestions, or on the other hand, the significant blanch-
ing of an organ as in sudden fright.

■^ ' Among Bernard's minor investigations which might be mentioned
is that, into the physiological action of curare, a black resenoid ex-
tract prepared by the South American Indians from the bark of strych-
nos toxifera and used to poison arrows. Owing to its poor diffusi-
bility through animal membranes curare i^ harmless taken into the
alimentary canal, though the minutest quantity introduced into a
wound is fatal., Since Bernard's time curare has become an instru-
ment in the hands of the physiologist to enable him to abolish tempo-
rarily the movements of the skeletal muscles, enabling him to carry
out experiments which could not be made without such aid.

The precise action of carbonmonoxide gas in asphyxia no one
understood until Bernard investigated the matter. His experiments

♦Futcher Journal A, M. A. Decemher 21, 1912.

40 PATHFINDERS OF PHYSIOLOGY

led him to conclude that C 0 was rapidly poisonous to animals owing
to the fact that it instantly displaced the oxygen of the red corpuscle
and could not itself be subsequently displaced by oxygen. The animal
died because the red corpuscles were, so to speak, paralyzed and cir-
culated as inert bodies devoid of the power of sustaining life.

A Friend of Pasteur — It is interesting to note that at a time
when physiological opinion favored spontaneous generation, vitalism
and such theories, the independent mind of Claude Bernard foresaw
what subsequent decades of physiological research have found to ap-
proximate the truth on such subjects. He was a firm friend of Pas-
teur, whom he ably seconded in his efforts to disprove spontaneous
generation.

A man is great in proportion to the obstacles he is able to sur-
mount. The subject of this paper illustrates the truth that one who
possesses in a high degree the qualities of genius will succeed in spite
of his surroundings. His early education, neither adequate nor con-
ducive of the best, together with the keen struggle for a livelihood,
and in his early career, the apathy of an unappreciative age and labor-
atories with meagre equipment, were obstacles which bring into relief
the rare qualities that he possessed. Contrast such a condition with
the magnificent equipment and endowment of modern scientific re-
search and the facilities for training as they exist today !

Bernard's life was far from being strewn with roses. He was mar-
ried to a wife who was non-appreciative of his genius. She saw noth-
ing in what to her was empty honor, the homage of the scientific
world, when the means which make for affluence were not forthcom-
ing. His two daughters became estranged from him and it is said
that one of them who was still living within the last ten years, joined
that silly sentimental class of antivivisectionists and endowed hos-
pitals for dogs and cats to atone for the crimes of vivisection which
her father had committed. Not only lacked he the sympathy which
"in true marriage lies," but he began his work at a time when the
physiologist had need of a "real passion for his science and in order to
ward off fatal discouragement had to possess his soul of high courage
and great patience. So soon as the experimental physiologist was dis-
covered he was denounced ; he was given over to the reproaches of his
neighborhood and subjected to the annoyance of the police;" Bernard
suffered all this.

But conscientious work well performed is not without its rewards
and perhaps the greatest is the satisfaction of "something attempted,
something done." He was a greater man than Majendie, whose re-
searches were made more or less at random and who had described
himself as a "rag picker by the dust heap of science." Bernard always
made his experiments with a definiteness of purpose. His contribu-
tions to physiology have been greater in number and importance than
those of any other investigator. Later in life he enlisted the friena-
ship of Emperor Napoleon UI., which resulted in two well equipped
laboratories which greatly facilitated his work. His academic oppor-
tunities included professorships in the College of France as well as a
chair at the Sorbonne. In 1868, he was admitted to the Academy of
France and made one of the "Immortals."

SKILL AS EXPERIMENTER 41

The Quest for Truth — As already mentioned, Bernard possessed
a faculty that contributed in no small degree to his success as phy-
siologist. Huxley has described an educated man as one whose hand
is the ready servant of his will. Often, too, great stress is laid upon
the purely intellectual qualities and too little upon that manual dex-
terity which is so essential to successful work in the laboratory.
In fact medicine itself is an art as well as an ensemble of sciences,
and the art is as important as the science. As much depends upon
the skillful use of the senses, and in surgery, skill in manipulation,
as upon the well trained mind. The extreme nicety with which Ber-
nard performed his dissections excited the astonishment as well as
the admiration of his associates. It was this faculty which first won
him the favor of Majendie. A clumsy experiment is apt to be a poor
experiment barren of results, and a patient's chances of life may be
jeopardized by an operation poorly performed.

Bernard was active until the end. On what proved to be his death-
bed he worked at the revision of proofs of a volume of lectures on
operative physiology. He died oiL-the tenth of February, 1878, and
was laid in the grave with all the pomp and ceremony of a state fu-
neral. Gambetta eulogized him as one who had never allowed himself
toloe led away either by party spirit or by the dogmas of a school, or
by private feelings. Bernard's work is a model of patient persevering
investigation, experiment and research, an unprejudiced and disinter-
ested quest for truth. He lived up to and fulfilled the ideals with
which he began his career, ideals aptly expressed : "Truth like beauty
is when unadorned, adorned the most." Such ideals have inspired men
of light and leading of all time; they inspire medicine today, ideals
old yet always new, and we may say with Kipling :

"The men bulk big on the old trail, our own trail, the out trail.
They're God's own guides on the Long Trail, the trail that's always new. '

;J

CHAPTER V.

RESPIRATION

he" ancients speculated upon the physiology of respiration;
Aristotle (384 B. C.) contended that the function of breathing was to
aol the blood. It was noticed that animals over-heated from exertion
breathed more rapidly, hence the inference. Galen (131-203 A. D.)
also maintained that the air inspired served to regulate and to cool
down the innate heat of the heart; that the peculiar action of the
chest wall seen in respiration introduced into the blood the air re-
quired for the regeneration of vital spirits in the left side of the
heart, whence by the arterial route they were distributed through-
out the body. Galen also recognized the necessity of ridding the body
of "fulginous vapors'* produced by the innate fire in the heart which
act was accomplished by expiration. In the latter part of the fifteenth
century, Leonardo da Vinci, painter, mathematician and naturalist,
disproved the fallacy that air simply cooled the blood in respiration.
He found that air was consumed by fire and that animals could not
live in a medium incapable of supporting combustion. This is the
first record in the history of science which pointed to the fact that
the function of air in respiration depended upon its chemical com-,
position and not upon its physical properties.

r^ It is evident that no real advance could be made in the physiology^
\of breathing until the circulation of the blood had been demonstrated.
Furthermore, this department of the science of physiology lagged
until the chemist appeared on the scene. Harvey had pointed out
that as the blood went to the lungs from the right side of the heart
thence to the left auricle a marked change took place, the blood as-
suming a bright arterial hue. The cause which resulted in this
peculiar change, Harvey was unable to discern, nor did it become
known until a much later day, when scientists became familiar with
the characteristics and constituents of atmospheric air.

Mechanics of Respiration. The first real knowledge on
the mechanics of respiration we owe to Borelli. Applying
the knowledge of muscular contraction on the one hand,
and atmospheric pressure on the other, he taught that inspira-
tion consisted of the entrance of air into the chest by virtue of at-
mospheric pressure, the thorax being enlarged by the muscular con-
traction of its walls; expiration consisted mainly in a cessation of
muscular contraction. Borelli broke with the ancient view that the
function of breathing was the cooling of the excessive heat of the
heart or the ventilation of the vital flame. "So great a machinery
and vessels and organs of the lungs," he continues, "must have been
instituted for some grand purpose ; and that we will try to expound,
if possible, though we shall stammer as we go along." Again he in-
, sists, "Air taken in by breathing is the chief cause of the life of ani-
j mals." It is more important than the heart and the circulation of the
blood.

^,^_-BX)

BERT BOYLE 43

The Work of Boyle, Now we turn to the English
school. Robert Boyle "(1627-1691), perhaps the most re-
nowned physicist of his time, by means of the air pump
jmade many researches on the "spring" of air. He showed,
among other things that a flame was extinguished in a partial
vacuum and that in a more complete vacuum not only the flame but
the lives of small animals such as the mouse ceased very quickly.
Here we see that the phenomena connected with the burning candle
closely resembled the phenomena of life ; furthermore that air what-
ever it might be, and not the mechanical movements of the chest
wall was necessary for the continuance of life. Boyle lived, at Ox-
ford for many years and while there made important improvements
in the air pump and in a long series of experiments with it made vari-?
ous discoveries in the properties of air and the propagation of sound
He was at the same time an ardent student of theology. He was ad-
vised to enter the church, but declined, feeling that his writings on
religious topics would have greater weight coming from a layman
than from a paid clergyman. As a man of science he was the ^rst
to carry out the principles of Bacon's Novum Organum.

The next step was taken by Robert Hooke, who was for some
time assistant to Boyle. xHooke was bom on the Isle of Wight, in 1635.
He was destined for the church, but ill-health diverted his career into
other channels, which gave scope for his precocious mechanical
genius. His personal appearance is described as very unattractive ;
his hair being in dishevelled locks over his haggared countenance. He
possessed an irritable temper and was much given to spending his
time in solitude.

To him Boyle was endebted for valuable work in connection with
the perfecting of his air pump. He was one of the earliest and most
zealous users of the microscope ; a volume entitled Micrographia, con-
tains an account of his many ''Observations Made on Minute Bodies
of varied kinds by magnifying glasses." Hooke's microscopic studies
on cork lead to the adoption of the term "cell" as the histologic unit.
He was curator of the Royal society, at a meeting of which he demon-
strated before the Fellows an experiment on artificial respiration,
which had been made before and many times since. The uniqueness
of the experiment consisted in the important conclusions which Hooke
made. The experiment consisted in opening the thorax of a dog and
substituting the movements of the chest wall by respiratory move-
ments accomplished by means of hand bellows, the nozzel inserted in
the trachea. This proved that the mechanical movements of the
chest wall were only of a secondary importance and that the whole
business of respiration was carried on in the lungs. This fact was fur-
ther proven by inflating the lungs to their utmost capacity and keep-
ing them distended by a powerful blast allowing the air to escape con-
tinually through minute holes pricked in the lungs. This showed that
life could be maintained even in the absence of the artificial move-
ments so long as the parenchyma of the lungs were so subjected to a
fresh supply of air. Therefore the secret of the change from venous
to arterial blood depended upon the exposure of the blood to fresh
air which was in the course of life accomplished by the bellows-like
action of the chest wall and diaphragm.

44 PATHFINDERS OF PHYSIOLOGY

Change in Color of Venous to Arterial Blood. — Richard Lower,
1631, concluded that the change in color, venous to arterial, blood was
due to the exposure of the blood to the air in the lungs ; he drew the
further conclusion that the change in color was due not to the ex-
posure alone, but to the fact that the blood took up some of the air;
that is, according to Lower, arterial blood differed from venous in
that it contains air. The blood gave up its "fresh air" in the course
of the circulation, hence the necessity of a constant supply of fresh
air for the maintenance of life. **Were it not for this, we should
breathe as well in the most filthy prison as among the most delightful
pastures." * * * ^j^^ fact," he continues, "where a fire bums
readily there we can easily breathe." Note that there was no men-
tion that only a part of the air was taken up by the blood. The com-
mon knowledge of the time was that air was a simple substance, not
a mixture of several elements as we know it today.

Mayow and His Researches. — ^The next contribution to the sub-
ject of respiration was that of John Mayow, bom in London in 1643.
Mayow was a lawyer by profession) and science was his avocation.
Many valued contributions to medical science were made by men
whose lives were spent in other callings. Priestly who discovered
oxygen was a Unitarian minister; Schleiden, whose name is con-
nected with the cell theory, was a lawyer; Schwann was a botanist;
Metchnikoff is a biologist. Thus many of the important discoveries
germain to medicine were made by men whose work was inspired by
the fascination of the subject in hand — ^the avocation of their leisure
moments. Of Mayow it was said he took his degree in law and
"became noted for his practice therein." Mayow's published works
consisted of four tracts — de sal nitro et spiritu nitro aero ; de respi-
ratione ; de respiratione f eotus in utero et ovo ; de motu musculari et
spiritibus animalibus. He showed that it was not the whole air
which was necessary for respiration, but only a portion, and that par-
ticular constituent of the air which has since become known as oxy-
gen. In the language of the chemists of his time, for he was essen-
tially a chemist, Mayow endeavors to prove "that this air which sur-
rounds us, and which, since by its tenuity it escapes the sharpness of
our eyes, seems to those who think about it to be an empty space,
is impregnated with a certain universal salt, of a nitro-saline nature,
that is to say, with a vital, fiery, and in the highest degree fermen-
tative spirit." The word "salt" was used by the seventeenth century
chemist to designate any substance not distinctly metallic or Jiquid.

Mayow sums up the conditions necessary for combustion; "con-
cerning fire.it must be noted that for the ignition of this it is neces-
sary that igneo-aereal (evidently oxygen) particles should either pre-
exist in the thing to be. burnt or should be supplied from the air.
Gunpowder is very easily burnt by itself by reason of the igneo-aereal
particles existing in it. Vegetables are burnt partly by means of the
igneo-aereal particles existing in them, partly by help of those
brought to them from the air." This early chemist recognized the
fact that in combustion we have a chemical combination with the
substance burnt, and as a result an actual increase in weight. He
experiments with antimony, which he bums by focusing the sun's
rays by means of a lens; by weighing the substance he finds an in-
crease in weight which he attributes the "insertion into it of igneo-

MAYOW 45

aereal particles during the calcination. As we shall see more than a
century later Lavoisier arrives at the same conclusion.. But Mayow
did not stop here. He proceeded to point out the identity of burning
and breathing:

"If a small animal and a lighted candle be shut up in the same vessel, the
entrance into which, of air from without is prevented, you will see in a short
time, the candle go out; nor will the animal long survive its funeral torch.
Indeed, I have found by observation that an animal shut up in a flask together
with a candle will continue to breathe for not much more than half the time
than it otherwise would, that is without the candle. * * * The reason why the
animal can live some time after the candle has gone out seems to be as follows:
The flame of the candle needs for its maintenance a continuous and at the same
time a sufiiciently full and rapid stream of nitro-aereal particles. Whence it
comes about that if the succession of nitro-aereal particles be interrupted, even
for a moment, or if these are not supplied in adequate quantity, the flame pres-
ently sinks and goes out. Hence, so soon as the igneo-aereal particles begin to
reach the flame scantily and slowly, it is soon extinguished. For animals, on
the other hand, a lesser store of the aereal food is sufficient, and one supplied at
intervals, so that the animal can be sustained by aereal particles remaining after
the candle has gone out. Hence it may be remarked that the movements of the
collapsed lungs not a little help towards the sucking of the aereal particles
which may remain in the said flask, and towards transferring them into the
blood of the breathing animal. Whence it comes about that the animal does not
perish until just before the aereal particles are wholly exhausted. * * * We may
infer that animals and fire deprive the air of particles of the same kind."

Mayow's account of the mechanics of respiration would need lit-
tle or no revision for a modem text book on physiology. He showed
that the air entered the lungs during respiration solely by atmospheric
pressure. He makes use of the experiment whereby a collapsed
bladder is placed into a bell- jar, the bladder expanding as the air in
the jar is exhausted by means of an air pump. He taught that in in-
spiration the chest is enlarged by the descent and contraction of the
diaphragm and by the raising of the ribs. Mayow further tackles
the raison d'etre of breathing in which he shows that something
necessary to sustain life passes from the air into the blood. "We
have no right," said he, "to deny the entrance of air into the blood
because on account of the bluntness of our senses we cannot actually
see the vessels by which it makes its entrance."

These extracts go to show how mature the views of the seven-
teenth century school of English physiologists, Boyle, Hooke, Lower /^
and Mayow in particular, were. Mayow by his nitro-aereal or igneo- ^
aereal substance evidently meant oxygen. Their work was, however,
allowed to slumber, until the scientific path was retraveled by their
successors nearly a century later.

Summary ^rior to the Beginning of the Eighteenth Century. —
Vto H5hnont^(1648) had discovered some of the properties of car-
bohdioxide. lie showed that a gas was formed from fermentation
or the combustion of carbon and from the action of vinegar on cer-
tain carbonates, and that this gas was incapable of supporting com-
bustion. Boyle (1670), as we have seen, proved that air was neces-
sary to ^e life of all animals, even those which lived under water.
Bernoulli, j at a later date, showed that the existence of aquatic ani-
^ale depended upon air held in solution in water. Hooke exposed

46 PATHFINDERS OF PHYSIOLOGY

the lungs of a living animal and maintained the vital pi^ocesses by
means of artificial respiration, showing that the vital processes de-
pended upon a continual change of air in the lungs. Fracassati drew
attention to the fact that the red color of the upper surface of a clot
was due to its exposure to air. Mayow (1674) advanced the view
that air contained a principle capable of supporting combustion, and
which, absorbed in respiration, changed venous into arteral blood
and was the cause of heat developed in animal bodies.

Eighteenth Century School. — Among the early eighteenth cen-
tury contributors to our knowledge of respiration was Stephen
Hales, bom 1677, who, by the way, was not connected with the med-
ical profession. He received his M. A. degree at Cambridge in 1703,
and Bachelor of Divinity in 1711. He was a clergyman by profession,
a calling which he followed until his death in 1761. He is chiefly
known as the inventor of a "ventilator," by means of which fresh air
was introduced into jails, mines, hospitals, and ships' holds. Four
years after the introduction of Hales' invention into the Savoy prison
only four prisoners died, whereas the mortality before its introduc-
tion had been as high as one hundred a year. Devoted as was Hales
to the church, he was even more devoted to science. He was the first
to determine blood pressure by actual experiment on the living animal.

Next in chronological sequence is Joseph Black, an eminent chem-
ist born at Bordeaux in 1728, where his father was engaged in the
wine trade. Both parents were of Scotch descent. In 1746 Black en-
tered the University of Glasgow, where he studied chemistry under
Dr. Cullen. He, however, graduated from the University of Edin-
burg in 1754. In a graduation thesis he proved that the causticity of
lime and the alkalis is due to the absence of carbonic acid present in
limestone. He did not use the term carbondioxide but instituted the
term "fixed air." The former name was first used by Lavoisier in
1748. Black's work was a distinct contribution to chemistry. In
1756, he became professor of anatomy and chemistry at Glasgow, but
shortly become professor of the Institutes of Medicine. In the mean-
time he practised his profession and found opportunity for original
investigation. In 1766 he was transferred to a similar position in
Edinburgh. His lectures were noted for their clearness and what is
perhaps the best testimonial to any lecturer, his classes became the
largest and best attended in the university. Though of delicate con-
stitution, by constant care he lived to the fairly ripe age of seventy-
one.

Black had been anticipated in his discovery of "fixed air" by Van
Helmont, whose researches had been made a century earlier r-Jn other
words, he had re=discovered the gas later to be known as C(P. By
using clear lime water, he was able to show that "fixed air" was given
off in fermentation, in expiration and that it was a product of burn-
ing charcoal. The chemical formula for clear lime water is Ca (OH) 2,
which in the presence of "fixed air," CO^, becomes Calcium Carbon-
at, CaCQS, which is precipitated as chalk, and water (H20). The
result of the chemical reaction is, of course, a reduction in the caus-
ticity of the original substance. "

I quote the following extracts from his treatise on chemistry:

"I had discovered that this particular kind of air, attracted by alkaline sub-
' stances, is deadly to all animals that breathed it by the mouth and nostrils to-

PRIESTLY 47

gether, but if the nostrils were kept shut I was led to believe that it might be
breathed in safety. I found for example that when sparrows died in it in ten or
eleven secondsi, they would live in it three or four minutes when the nostrils
were shut by melted suit. And I convinced myself that the change produced on
wholesome air "by breathing it consisted chiefly, if not wholly, in the conversion
of part of it into fixed air. For I found, that by blowing through a pipe into
lime water, the lime was precipitated, and the alkali was rendered mild. ♦ * ♦
In the same year I found that fixed air is the chief part of the elastic matter,
which is formed in liquids in the vinous fermentation. Van Helmont has indeed
said this. But it was at random that he said it was the same with the Grotto del
Cane in Italy (but he supposed the identity because both are deadly), for he had
examined neither of them chemically, nor did he know that it was the air dis-
engaged in the effervescence of alkaline substances with acids. I convinced
myself of the fact by going to a brew house with two phials, one filled with dis-
tilled water and the other with lime water. I emptied the first into a vat of
wort fermenting briskly, holding the mouth of the phial close to the surface of
the wort. I then poured some of the lime water into it, shut it with my finger,
and shook it. The lime water became turbid immediately."

Black goes on to criticise Van Helmont's pronouncements as
mere chance statements. He, himself, verified all his conclusions by
repeated experiment.

As Black re-discovered under the term "fixed air" that which
Van Helmont had recognized a century before, so Mayow's igneo-
aereal salt or spirit was re-discovered by Priestly and Lavoisier.

Priestly and His Dephlogisticated Air: With the name of Joseph
Priestly, perhaps more than any other, is associated in the modern
mind the discovery of oxygen, though he did not make use of the
term. Of him Frederick Harrison has said:

"If we choose one man as a type of the intellectual energy of the eighteenth
century we could hardly find a better than Joseph Priestly, though his was not
the greatest mind of the century. His versatility, eagerness, activity and hu-
manity; the immense range of his curiosity in all things physical and social; his
place in science, in theology, in' philosophy and in politics; his peculiar relation
to the Revolution, and the pathetic story of hisi unmerited sufferings, may make
him the hero of the eighteenth century."

He was bom near Leeds, England, in 1733, and died in the United
States in 1804. His boyhood was uneventful. His family was de-
scribed as "simple, sober, honest. God-fearing folk, staunch Calvinists
and deeply religious." The son inherited these qualities and entered
the ministry as a Unitarian preacher, an act which was particularly
offensive to the orthodoxy of the time. Benjamin Franklin, to whom
Priestly is endebted for the incentive for scientific study, refers to
him in a letter as an "honest heretic." And continuing in Franklin's
charactersitic style, he says:

"I do not call him honest by way of distinction, for I think all the heretics
I have known have been virtuous men. They have the virtue of fortitude, or
they would not venture to own their heresy; and they cannot afford to be dif-
fident in any of the other virtues, as that would give advantage to their many
enemies; and they have not like orthodox sinners, such a number of friends to
excuse or justify them. Do not, however, mistake me. It is not to my good
friend's heresy that I impute his honesty. On the contrary 'tis his honesty that
has brought upon him the character of heretic."

48 PATHFINDERS OF PHYSIOLOGY

Priestly was thirty years old when Franklin was sixty. Priestly
like Franklin was well informed on a variety of subjects. He wrote
learnedly on politics, religion and on science, particularly on pneu-
matic chemistry. Boswell dubbed him a "literary Jack-of-all-trades,"
and he was busy with proof sheets until the day of his death. His
pamphlets on politics and religion were so much opposed by the orth-
odox theologians of his day that they answered his arguments by
burning his house and dispoiling his belongings, a peculiar way that
the so-called orthodox theology has had in the past of dealing wih
those bold intrepid spirits who have dared to stand for what they
believed to be the truth. His home surroundings in Birmingham
became so unpleasant that in self-defense he set sail for America,
here to breathe the atmosphere of civil and religious freedom. He
was offered the professorship of chemistry in the University of Phila-
delphia, but the following year moved to Northumberland, a town on
the Susquehanna, a hundred and thirty miles northwest of Philadel-
phia. He lived and worked until his death, which occurred in Feb-
ruary, 1804.

Priestly endeavored to change back to its original condition, air
that had been breathed, or which had failed to support the flame of a
candle. He eventually succeeded by means of vegetation. First he
experimented by placing a sprig of mint into a glass jar standing in-
verted over a vessel of water. Parenthetically, Priestly invented the
pneumatic trough, which has been found so convenient in experiment-
ing with gases. When the sprig of mint had been growing some
months, the air within the vessel would not extinguish a flame nor
act deleteriously to small animals, such as the mouse, placed therein.
The growing plant really contributed to the flame or the animal that
was placed in the vessel. Further experiment showed that a growing
plant placed in a vessel in which a flame had been extinguished would
in time render the atmosphere in the jar capable of supporting either
flame or animal life. This lead him to conclude: "That plants, in-
stead of affecting the air in the same manner with animal respiration,
reverse the effects of breathing and tend to keep the atmosphere
sweet and wholesome when it is become noxious in consequence of
animals either living and breathing or dying and putrifying in it."

Priestly's researches might have been more fruitful in results
had he not been dominated by the phlogiston theory, a term devised
by^ Stahl. Phlogiston, from phlogistos, burnt, was a hypothetical
principle of fire regarded as a material substance. Every combustible
substance was a compound of phlogiston and the phenomenon of com-
bustion was due to a separation of the compound into its component
elements.

Priestly was able to obtain the same gas by heating mercuric
oxide, and from red precipitate. But he could not get away from
the phlogiston theory. Air supported combustion because it took up
phlogiston given out by the burning body. Common air supported
combustion in proportion as it was free from phlogiston. He pre-
pared oxygen in 1774, that is he discovered that the gas he prepared
was part of the common air, which supported life and combustion.
Venous blood was blood laden with phlogiston. Blood exposed to de-
phlogisted air gave up its phlogiston and became bright arterial blood.

LAVOISIER 49

Some idea of the scope of Priestley's researches may be inferred
from the mere catalogue of his discoveries. He is^ -credited with dis-
covering^ dephlogisticated air toxygen) hydrochloric acid, sulphur
dioxide, nitrosulphuric acid, sulphuretted hydrogen, and the isolation
of amonia gas.

/^Lavoisier and His Work. — Antoine Laurent Lavoisier was born
in( i^ari^^in 1742^ jben years later than the date on which Priestly first
saw^thcr^M of day. As scientist his career was practically contem-
poraneous with that of Priestly, who made the same momentous
discovery, working independently. In 1775, a year after Priestly had
prepared his dephlogisticated air (oxygen), Lavoisier published his
paper "On the nature and principle which combines with metals dur-
ing their calcinat ionT" Irfthis paper he showed that metals on being
"burnt did not give up phlogiston to the air but took something from
the air; they on becoming metallic oxides, increased^ iiTweighT. La-
voisier deall-the death- blow jto the phjogiston theory and was in a
sense the real discover of oxygen. ~Iltl)r6ved'Ehat the principle which
combined with metals when calcined was the principle of acidity. He
says: "I shall therefore designate dephlogisticated air, air eminently
respirable, when in a state of combination or fixedness by the name of
'acidifying principle,' or, if one prefers the same meaning in a Greek
dress, by that of 'oxygine' principle." Lavoisier discovered oxygen
and gave it the name by which it will henceforth be known. He made
further experiments in connection with respiration which he con-
cluded to be "a combustion, slow it is true, but otherwise perfectly
similar to the combustion of charcoal." He eventually saw, however,
that some of the oxygen inspiredhad other use than the production
of carbon dioxide.

It was not, however, until the early decades of the nineteenth

century that the view that oxidation took place in the lungs gave

way to the accurate theory of tissue respiration. In 1837, Gustave

Magnus proved that both venous and arterial blood contained oxygen

^and carbon dioxid.

Hydrogen was discovered by Cavendish in 1781, when he also
discovered the composition of water. Nitrogen was discovered in
1772 by Rutherford. Oxygen was prepared by Priestly in 1774 and
recognized by Lavoisier the following year. Carbonic acid gas, or car-
bon dioxide was first discovered by Van Helmont in 1640 and redis-
covered and defined by^lack in 1757.

7^

CHAPTER VI.

THE NERVOUS SYSTEM.

The progress of knowledge of the nervous system has been very-
slow. Most of the other viscera were known to the ancients before
the brain was recognized. The word "brain" is not to be found in the
Bible. The ancient Hebrews evidently looked upon the heart as the
seat of the soul. The kidneys were the habitation of the mind, while
the tender emotions were referred to the bowels. Plato was perhaps
the first to assign the supreme seat of the mind to the brain, but his
views were purely speculative, inasmuch as he confounded the sub-
stance of the brain and of the spinal cord with the marrow of bones.
Aristotle, about 335 B. C, examined the brain for himself and con-
cluded that its function had nothing whatever to do with the mind,
but that it was a refrigerating organ which cooled the blood for the
heart. He reasoned according to the knowledges of his time. The
brain was apparently an insensible and inexcitable organ as contrast-
ed with the heart, which is the opposite. Hippocrates recognized how
soon animals became unconscious from the loss of blood, or how
changed by blood poison or by the heated blood of fever; hence the
inference by Aristotle that the conscious mind resided in the blood
and that the great central organ, the heart, was the seat of the soul.
The arteries (from the etymology, air tubes or wind pipes) found
empty after death, were supposed to carry air or ^^ethereal" spirits to
the rest of the body. It was this great blunder that delayed for cen-
turies, virtually until Harvey's time, all progress of knowledge of the
true function of the heart. Hippocrates maintained that the brain
was a gland. With this supposition subsequent writers ventured the
suggestion that the brain secretion was a subtile fluid which they
designated "animal spirits.** The authority of such names as Hip-
pocrates and Aristotle forbade first hand investigation for fully five
centuries. It must not be overlooked, however, that amid all this
guessing, Alcmaeon (about 500 B. C), an anatomist and physiologist,
taught that the brain was the seat of the mind and that all sensation
traveled to the brain by means of the nerves. He spoke of the nerves
as "tendons" which misconception held sway until Descartes, the
philosopher, showed the difference between tendons and nerves.

About 300 B. C. sprung up the Alexandrian school of anatomists
and physiologists of whom Herophilus and Erastistratus were chief
who dissected the brain and traced to it the nerves as Alcmaeon had
done. They even went so far as to distinguish nerves of sensation
and nerves of motion, but were still hampered by Alcmaeon's "ten-
dons." "When Greece fell under the subjection of Alexander, mind went
into exile, and its first asylum was the city of the conqueror." Under
royal patronage the study of anatomy and physiology and surgery
made great progress. Galen spoke of Herophilus and Erastistratus as
possessing more accurate knowledge of the human body than any one
before their time. Herophilus was the first anatomist of importance
in the annals of medicine. He is said to have discovered the lacteal

THOMAS WILLIS 61

vessels, and the construction of the eye, including the retina. Galen
speaks of Herophilus as having a very intimate knowledge of the
anatomy of the nervous system. The term, "torcular Herophili"
signifies the "press" or dilation at the junction of the superior longi-
tudinal, lateral and occipital sinuses first described by Herophilus.
Herophilus and his associates performed vivisection upon condemned
criminals. Not only did medicine progress during this early period
(about 300 B. C), but literature, philosophy, mathematics, natural
history and astronomy flourished as well under the patronage of
Ptolemy. A great part of the record of this fruitful period was lost
during the seventh century of the Christian Era, with the destruction
of the great Alexandrian library.

"The Brain the seat of Thought and Sensation:" Galen, A. D.
160, overthrew Aristotle's theory in regard to the brain and showed
it to be the seat of thought and sensation. Aretaeus (170 A. D.)
taught that the brain controlled the muscular movements of the body
by means of nerves originating in the brain. He recognized the
crossing of the nerves so that injury to one hemisphere pro-
duced paralysis on the opposite side. If injury occurred in the
cord below the medulla the paralysis was on the same side as the in-
jury. The seat of the soul was, however, in the heart.

Andreas Vesalius (1514-1564) declared that the "brain in ap-
propriate structures, and in organs properly subserving its work
manufactures the animal spirit which is by far the brightest and
most delicate, and indeed is a quality rather than a natural thing.
* * * Nerves serve the same purpose to the brain that the great
artery does to the heart." The nerves he regarded as the "busy at-
tendants and messengers of the brain." Vesalius, however, is free in
the use of such terms as "vital soul," "vital spirits," "animal spirits,"
which meant so much to the physiologist of his day and so little to
us of the twentieth century. While these meaningless terms make a
great deal of his work unintelligible, yet there abound throughout
gleams of truth as we understand it today. He showed that by sever-
ing a nerve or by ligation it was possible to abplish_the action of
the nerve upon the muscle. Regarding the brain he says : ~**But how
the brain performs its functions in imagination, in reasoning, in
thinking and in memory, I can form no opinion whatever."

Nearly a century later we come to the conclusions of von Hel-
mont and of Descartes which were much less to the point than the ex-
pressed opinions of Vesalius. One placed the seat of the soul in the
pylorus ; the other in the pineal gland.

Malpighi devoted much attention to the histology of the nervous
system but said practically nothing about the functions of the nerves.

Thomas Willis: Perhaps the most important investigator of
the seventeenth century into the anatomy and physiology of the
nervous system was Thomas Willis. He was born in Wiltshire, Eng-
land, in 1621, educated at Oxford where he graduated with the degree
of M. A., 1642. He eventually entered upon the study of medicine
and on graduation was appointed to a professorial chair in Oxford.
Here he taught, practised medicine and pursued his scientific re-
searches. In 1666 he located in London where in the language of a

52 PATHFINDERS OF PHYSIOLOGY

contemporary "he became so noted and so infinitely resorted to, that
never any physician before went beyond him or got more money year-
ly than he." Willis possessed a practical knowledge of the structure
and functions of the brain, both in health and disease. His name to-
day is familiar to all students of anatomy in the "circle of Willi^,"
which designates the combined arterial structure at the base of the
brain. Sir Michael Foster is inclined to depreciate the work of Willis.
The value of his book is much above the worth of the author. It ap-
pears that WilHsHhirst for fame was much greater than his love for
truth. Richard Lowfer, a contemporary was the real man of science
of his day. Willis is said to have appropriated the work of Lower
and other earnest men and to have published it as hts own. Through-
out the work of this period we still have to deal with ^he "corporeal
soul/' "animal spirits," "sensitive sottl*'^ ^d similar phrases.

Muscle Irritability: Frances Glisson, an Englishman, born, 1597,
came upon the truth of the relation of nervous influence to muscular
contraction. Educated at Cambridge, he became a Fellow and lec-
turer in Greek in his Alma Mater. On the publication of Harvey's
work, in 1628, Glisson determined to turn his attention to medicine,
and six years later he received his M. D. degree. He did not go
abroad as Harvey did but pursued his medical studies in London. He
was soon appointed Regius Professor of physic at Cambridge, but it
seems did not spend much time there, as the social atmosphere was
not congenial to him. Cambridge was strongly Royalist, while Glis-
son was a very pronounced Presbyterian. He served in a professional
capacity in London during the great plague of 1665. He died at the
ag^ of eighty years. He is probably best known for his work on the
Jiver. His name is familiar to us in connection with the capsule cov-
( ering that viscus. Glisson's studies on the liver lead him, to his dis-
covery regarding the peculiar properties of muscle tissue.

Explaining how the bile is discharged into the intestine only
when it is needed, he shows that the secretion is greater when the
gall bladder and passages are irritated, hence they must possess the
power of being irritated. For this peculiar property he suggests the
term irritability. The idea was not seized by contemporary physi-
ologists, hence Glisson's work remained dormant until the following
century, when Haller made use of the term, and since his day it has
become established in physiology and has played an important part in
the development of both physiology and pathology.

Goll and Phrenology: A name which has received but slight at-
tention at the hands of biographers is that of Franz Joseph Gall or
Goll, best known as the founder of the pseudo science of phrenology,
or "bumpology" as it has been contemptiously called. Goll was born
in 17^8. He took his degree in medicine at the University of Vienna,
in 1785, where his studies on brain and mind began. He was an acute
observer of phenomena and from a collation of observed facts was
the first to demonstrate that the brain was the organ of the whole
mind. The modem phrenologist with whom we are more or less
familiar, is a disciple of Goll; his name will be remembered as as-
sociated with the discovery of certain areas in the spinal cord. Goll
died in Paris in 1828.

BELL AND MAGENDIE 53

Bell and Magendie: One of the greatest names in connection
with the anatomy and physiology of the nervous system is that of
Sir Charles Bell. In fact, his discovery has been placed in importance
in the satne efess as that of William Harvey. Charles Bell was bom
at Edinburgh, Scotland, in 1774. After graduating from the Uni-
versity of Edinburgh he began the study of medicine under his elder
brother John, who had already achieved distinction as anatomist.
After graduating he devoted himself to anatomy and surgery. He
eventually moved to London, where he worked into a very lucrative
surgical practice. His first published work (1798) bore the cumber-
some title ,and it was the custom of writers of the time to preface
their work with a sentence descriptive of its contents, of "A System
of Dissections Explaining the Anatomy of the Human Body, the
Manner of Displaying its parts and their varieties in Disease.*' Four
years later Bell published a series of engravings of original drawings
showing the brain and nervous ^stem. His drawings are worthy of
special mention. His skill as anatomical artist rivaled that of anato-
mist. He was also the author of a work entitled "The Anatomy of
Expression," the object of which was to describe the arrangement
by which the influence of the mind is propagated to the musculature
of the face and to give a rational explanation of the muscular move-
ments which accompany the various emotions and passions. He
emphasized to the physician and surgeon the importance of a
knowledge of facial expression in diagnosis, to ascertain the nature
and extent of bodily suffering. In these days of the clinical labora-
tory and multifarous other clinical methods, the ability to make a
diagnosis by observation alone which amounted to intuition with the
old-time clinicians, is a lost art. This work, which was illustrated
by himself, had a wide circulation in his day.

Charles Bell's most important work, however, was the discovery
of the double system of nerves Issuing from the spinal cord. He dis-
covered that in the nerve trunks are special sensory filaments to
transmit impressions from the periphery of the body to the sen-
sorium and motor filaments to convey motor-impressions from the
brain or other nerve centres to muscle. He demonstrated that the /
anterior roots of the spinal cord were motor and the posterior roots/
sensory.

While in London, he was Professor of Anatomy, Physiology and
Surgery in the College of Surgeons. He was knighted by William
IV. He returned to Edinburgh 1836 where he became professor of
anatomy and surgery. His name is associated with the disease
which he was the first to accurately describe, paralysis of the sev-
enth nerve— "Bell's Palsy.*' He died in 1842.

A name of only less importance than that of Sir Charles Bell is
that of Magendie. Magendie has been considered the greatest phy-
sician France had produced down to his day. His work on physiology
written while in his early thirties was almost immediately translated
into English and German. It was a valuable work, inasmuch as it \
was based upon experimentation. He was the first continental in-
vestigator to discover therfimction of the spinal nerves, and accord-
ing to Gorton, contributed more to the knowledge of the nervous
system than any of his distinguished predecessors. Magendie was

54 PATHFINDERS OF PHYSIOLOGY

bom at Bordeaux, France, in 1783, studied medicine in Paris, where
he became demonstrator, and eventually professor of anatomy in the
College of France. He died in 1855. ^

Magendie is described as being abrupt in manner, even to rude-
ness. His brusque manner has been referred to in his relations with
his understudy, Claude Bernard. He seems, however, to have been
a brilliant if not very methodical worker. He refers to himself as
a ragpicker by the dust heap of science. His work on the nervous
system was parallel with that of Sir Charles Bell, and the scope of
the work of both is epitomized in the well-known Bell and Magendie
Law to the effect that the spinal roots may be divided into afferent
and efferent, the anterior roots carrying impulses only from the
spinal cord to the periphery, while the posterior roots carry impulses
from the periphery to the central nervous system; a nerve fibre can-
not be both motor and sensory; we may have both nerve fibres in a
single nerve trunk but the fibres in each case are isolated and con-
duct impulses only in one or other direction.

To Claude Bernard, associated with Magendie in the College
of France, we owe the discovery of the vaso-motor nerves.

Broca and the "Speech Center." — In 1861 Paul Broca, an immi-
nent French surgeon, proved that there is a definite locality in the
brain which is the seat of articulate speech. This is known today as
"Broca's Convolution." Nine years later, thanks to the labors of such
men as Hitzig, Ferrier and Charcot, it was shown that each of the
special senses has its anatomical seat in the brain. It was also found
that each volutary muscle or group of muscles could be made to con-
tract by the excitation of certain "centers" or localities in the surface
of the brain. Regarding later progress in brain physiology, Gorton
says: "It is worth while to note the stride anatomy has made dur-
ing the closing years of the nineteenth century, especially in knowl-
edge of the central nervous system of man and animals. Early in the
last decade of the century the subject was taken up by German and
Italian anatoniists, Waldeyer, Nissl, Marchi, Golgi, His, Apathy and
others. To Waldeyer we are indebted for the doctrine of neuron as
applied to nerve cells, from the Greek word "neuron,' signifying unit.
According to this doctrine every cell is a unit having an independent
existence, distinct and apart from other cells, though related to them,
and may degenerate and die without affecting the existence of the
others. Meynart estimates that "the cortex of the cerebral hemis-
pheres ^lone <;ontains_twelve hundred millions of ganglionic cells;"
and Donaldson states that three thousand miltion: cells "is a modest
estimate of the total number of these neurons in the central nervous
system." The doctrine of neurons has been assailed as applied to com-
parative histology by the distinguished Apathy, and defended among
others by Barker, of Johns Hopkins University." The invention of
staining processes afforded a powerful impetus to the study of nerve
tissues.

By means of animal experimentation Flourens, Luciani and
Horsley determined the function of the cerebrum. Removal of the
cerebrum from a frog or pigeon caused all its voluntary movements
to cease, but did not interfere with the reflexes or the negative func-

PATHOLOGIC STATES OF THE BRAIN 55

tions. The same investigators found that if the cerebrum was re-
moved and the cerebellum left, the animal has sense of appreciation,
but fails in muscular coordination. Stephen Hales showed that the
spinal cord is necessary for reflex movements and Marshall Hall work-
ed out the whole problem of reflexes. Galvani (1791) studied reflexes
by applying electric stimuli to frogs' legs.

Pathologic States of Brain and Nervous System — Apropos of the
development of knowledge of the physiology of the nervous system is
the evolution of our knowledge of its pathologic states. The insane
have suffered much owing to Jgnorance and misconception on the
partx)f^the~sane^ Ancient nationsTookedlipoh tHe msane aslpossess-
ed of evil spirits or as "possessed of devils." Later the Greek, Alex-
andrian, and the Roman, looked upon the insane man as a sick man
and he was accordingly treated by means of drugs, baths, exercise and
other hygienic measures. A great retrogression took place during the
second or third centuries of the Christian era. Theories of demoniac
posses&ion again held sway, with the result that the insane were sub-
jected to the utmost cruelty. This attitude continued throughout the
Middle Ages. In fact, no marked advance was made until the eigh-
teenth century. Various places of custody were maintained for the
insane where they were confined in dungeons, badly clothed and bad-
ly fed. The first real advances in their care were made by Philip
Pinel, in France. Tuke, in England, and Benjamin Rush, of America,
near the end of the eighteenth century. Pinel in 1793 substituted a
system of non-restraint and humane treatment for blows and punish-
ments. William Tuke, member of the Society of Friends, was mak-
ing similar reforms in England. Stahl, early in the eighteenth cen-
tury, insisted on the essentially sinful character of insanity and this
attitude found echo in Heinroth in the early nineteenth century.
Religious theories have little by little given place to physiological and
psychological explanations until today the insane man is regarded as
a sick man. Insanity implies disease organic or functional, just as
do other abnormal manifestations.

Note: I am indebted to F. X. Dercum's work on Mental Diseases, 1913,
for the data of the last paragraph.

CHAPTER VII

THE CELL THEORY

"The cell theory furnishes the starting point for all modem
studies in biology and enables all students to speak the same
language," says a twentieth century writer. The recognition of the
fact that animals and plants are constructed on a similar plan must
be placed among the most important discoveries of the nineteenth
century prolific as that century has been in scientific achievement.
"No other biological generalization," says Professor Wilson,
referring to the cell theory, "save only the theory of or-
ganic evolution has brought so many diverse phenomena
under a common point of view, or has accomplished more for the
unification of knowledge." By the term "cell-theory" is understood
the teaching that all animal and plant tissues are composed of units
known as "cells," which term as we shall see is inappropriate so far as
the actual things designated by it are concerned. The cell-theory is
a generalization which places animals and plants on a basis of similar-
ity of structure.

Anticipated in the Seventeenth Century: The cell doctrine was
anticipated as far back as the seventeenth century, for it is to a
worker of the mid-seventeenth century that we are endebted for the
term "cell." Robert Hooke, an English microscopist, experimented
with cork, which he declared to be made up of "little boxes or *cells*
distinguished from one another." He made thin sections by means
of a pen knife and found them to be all "cellular or porous in the man-
ner of a honeycomb." Malpighi and Leeuwenhoek, in the seventeenth
century, made drawings which have been preserved showing the cell
structure of plants; we may therefore conclude that the cell theory
announced in 1838, was foreshadowed by seventeenth century work-
ers. Wolff, an acute scientific observer in 1759 worked out the
identity of plants and animals, as shown by their development. Hux-
ley summarizes Wolff's view of the development of elementary parts
as follows: "Every organ, according to him, is composed at first of
a little mass of clear viscous nutritive fluid which possesses no or-
ganization of any kind, but is at most composed of globules. In this
semi-fluid mass cavities are now developed; these if they remain
round or polygonal, become the subsequent cells; if they elongate,
the vessels; and the process is identically the same whether it is ex-
amined in the vegetating point of a plant or the young budding organs
of an animal."

Bichat's Contribution: Though his connection with th^e cell
theory is open to question, the name of Bichat is deserving of mention
in discussing it. Marie Francois Xavier Bichat, bom in France in
1771, is noted as the founder of histology. He studied in Paris under
the great surgeon Desault. He was himself made professor of anatomy
at the age of twenty-six years, a position which he held until death
relieved him of his labors at the early age of thirty-one. It is related

M. SCHLEIDEN,

THEODOR SCHWANN,
Co-founders of the Cell-Doctrine. From Locy Biology and Its Makf&fs.

THE CELL THEORY, 1838 57

that he won the attention and admiration of his chief by making a
complete extemporaneous report of one of Desault's lectures. Bichat
was a most admirable character; he has been des(Jribed as of "mid-
dling stature, with an agreeable face, lighted by piercing and expres-
sive eyes," and as being "in all relations of life most aimable, a
stranger to envy or other hateful passions, modest in demeanour, and
lively in his manners which were open and free." Two of his works,
his treatise on the membranes and his general anatomy are important
as the foundation of histology, or the minute anatomy of the tissues.
After the ennunciation of the cell theory Bichat's work took on a new
phase, namely that of microscopic study of the tissues. Schwann's
cell theory was in reality an extension of his work. Bichat's claim
for credit in connection with the cell theory has been called into ques-
tion inasmuch as his investigations were done without the aid of the
microscope.

The Cell Theory, 1838: During the first three decades of the
Nineteenth century there accumulated a great mass of unconnected
observations on the microscopic structure of both animals and plants.
"We must clearly recognize," said Tyson, "the fact that for some
time prior to 1888 the cell had come to be quite universally recognized
as a constantly recurring element in vegetable and animal tissues,
though little importance was attached to it as an element of organiza-
tion, nor had its character been clearly determined."

Eighteen hundred and thirty-eight was an epochal year in biolog-
ical science, chronicling as it does the ennunciation of the cell-theory
by Schleiden and Schwann the result of the combined efforts of
botanist and animal biologist. The work of Schwann, however, was
more comprehensive and important than that of Schleiden, and to
him, therefore, belongs the greater honor.

M. Schleiden was educated for the legal profession and had engaged
in the practice of law. He soon abandoned it for medicine, but after
graduation devoted himself to the study of botany. Locy describes
his work in 1837, stating that he arrived at a new view in regard to
the origin of plant cells. This new view though founded upon er-
roneous observations and conclusions served to provoke discussion.
His work acted like a ferment, we are told, in bringing about new ac-
tivity. Schleiden was noted for his alertness in entering upon con-
troversies, a trait which better befits the lawyer than the man of
science whose sole concern should be the quest of truth. His replies
to his adversaries were at times vitriolic and he often indulged in bit-
ter personalities. Perhaps his legal training was responsible for this.

His methods of investigation were sound, based as they were on
experiment and observation. He conceived the necessity of studying
the development of plants in order to understand their anatomy and
physiology. The nucleus of the plant cell was discovered in 1831, by
Robert Brown. Schleiden seized upon the nucleus as the starting
point of new cells but wrongly supposed that the new cells started
from a small clear bubble on one side of the nucleus. And yet it was
through these inaccurate observations of Schleiden that his co-found-
er, Schwann, arrived at his general conclusions. An incident is re-
lated of the two dining together one October evening when Schleiden

58 PATHFINDERS OF PHYSIOLOGY

took occasion to relate to his friend his observations and inferences.
Schwann was impressed at once with the similarity to his own ob-
servations on animal tissues. They at once proceeded to Schwann's
laboratory where sections of the spinal cord were examined. Schleiden
recognized the nuclei as similar to those he had found in plant cells.

Theodor Schwann: Schleiden and Schwann seem to have been the
most diverse personalities. The former was pugnacious and always
ready to take up the gauntlet in controversy; the latter was one of
the mildest of men. We are endebted to Henle, a name familiar in
microscopic anatomy, for what we know of the life of Schwann. This
is Henle's description of him: "He was a man of stature below the
medium, with a beardless face, an almost infantile and always smiling
expression, smooth dark brown hair, wearing a fur trimmed dressing
gown, living in a poorly lighted room on the second floor of a restau-
rant which was not even of the second class. He would pass whole
days there without going out, with a few rare books around him, and
numerous glass vessels, retorts, vials and tubes, simple apparatus
which he himself made. Or I go in imagination to the dark and
fusty halls of the anatomical institute where we used to work till
night fall by the side of our excellent chief, Johann Muller. We took
our dinner in the evening, after the English fashion so that we might
enjoy more of the advantages of daylight."

Johann Muller: The mention of Johann Muller is worth a mo-
ment's digression. Muller, the son of a poor shoemaker, was bom at
Coblentz in July, 1801. Perhaps it was the meagerness of his worldly
possessions, for have not all the followers of Saint Crispin been men
of lowly estate, that served to bring out the true metal of his charac-
ter. Surmounting the disadvantages and lack of opportunity of
youth he became eventually one of the great teachers and master
minds of German science. The inspiration derived from a great
teacher or personality is difficult to comprehend much less to explain.
Harvey was influenced by his association with Fabricius; Bernard
was similiary inspired by Magendie. The dominant physiological mind
during the first half of the nineteenth century was that of Muller.
He was the great trainer of anatomists and physiologists. Among
desciples during his professorship at Berlin were Virchow, the patho-
logist; Du Bois Reymond and Brucke, the physiologsits ; Henle, the
anatomist; Helmholtz, and Leiberkuhn. All became distinguished
scholars and professors in German universities. In glowing tribute
to his master, Helmholtz said: "Whoever comes in contact with men
of the first rank has an altered scale of values in life. Such intellec-
tual contact is the most interesting event that life can offer."

Muller's manner and gestures in the classroom reminded his
hearers of a Catholic priest. The way he impressed the scientific men
of his time is best evidenced by the numerous tributes accorded his
memory. Verworn says: "He is one of those monumental figures
that the history of every science brings forth but once. They change
the whole aspect of the field in which they work and all later growth
is influenced by their labors." And of his monumental work the
Handbook of Physiology, which appeared in 1833, the same eulogist
writes : "This work stands today unsurpassed in the genuinely philos-

JOHANN MULLER 59

ophical manner in which the material, swollen to vast proportions by
innumerable special researches was for the first time sifted and elab-
orated into a unitary picture of the mechanism within the living or-
ganism. In this respect the handbook is not only unsurpassed but
unequalled."

To sum up and to sift the accumulated knowledge of a depart-
ment of scientific endeavor is truly a herculean task, one requiring
the impartiality of a judge and energy and zeal for the work that
amounts to genius. Haller performed a similar service for physiology
in his day.

Johann Muller a "vitalist:" Attempts have been made to ac-
count in some more or less satisfactory way for the phenomena of life.
Two theories have engaged the attention of scientists — vitalism and
the chemico — physic or mechanistic theory. The majority of scien-
tists of the present day maintain that living organisms are mere
machines, as opposed to the theory of vitalism which presupposed the
presence of some "life" principle. The chemico-physicist today sees
nothing that may not be explained by the ordinary laws of physics
and chemistry. The tendency in all science is to express the less sim-
ple in terms of the more simple. Every activity of living substance
is accompanied by molecular or chemical changes in its composition,
such as oxidation (combustion) so that chemical activity, which is
the source of energy, and all vital manifestations are physico-chemical
in nature. Haller, in 1700, defined vitalism or vital force as a life
principle which possessed the ability to originate energy, which meant
that an organism was not wholly dependent upon the food which it
consumed for its energy.

The scientists of the period 1810 to 1850 were, for the most part,
adherents to the mechanical explanation of the phenomena of life.
During this time also, the vitalistic theory was not without its advo-
cates who were among the pupils of the great idealist philosopher,
Schelling. Such men as Johann Muller, the physiologist; Von Baer,
the embryologist, and Liebig, the chemist, were said to be close ad-
herents to the vitalistic theory. It was not, however, until 1847, the
date of publication of the researches of Helmholtz on conservation of
energy that vitalism received a stunning blow. Sir Michael Foster A
explains Muller's vitalistic leanings by declaring that, "He was a
vitalist only in the sense that he was theoretically of opinion that
even when the physico-chemical analysis of vital phenomena had
been pushed as far as it could, there would still remain a large residue
which could not be explained by any such analysis, however complete.".
In view of the fact that his great pupils were noted for their effort to
solve physiological problems by physico-chemical means, the explana-
tion is plausible. It might be stated that Schwann, as well as other
pupils of Muller, had recourse to vitalistic explanations only
when their means of analysis proved inefficient.

"The graven image, vitalism," says Starling, "has acted as a con-
tinual check on the growth of man's knowledge and control of his
environment just as the hypothesis of special creation would impede
all research into the relationships of animals and plants, so vitalism
would stay the hand of the physiologist in his endeavors to determine
the changes which occur within the living organism."

SO-, , "- PATHFINDERS OF PHYSIOLOGY

Muller died in 1857. Vircho:^)^, at his obsequies in Berlin, indulged
in the following panegyric over his master :

"My feeble powers have been invoiced to honor this great man,
whom we all, representatives of the great medical family, teachers and
taught, practitioners and investigators, mutually lament and whose
memory is still so vividly with us. Neither cares by day nor labors
by night can efface from our mind the sorrow which we feel for his
loss. If the will made the deed, how gladly would I attempt the hope-
less task of proper appreciation. Few have been privileged, like my-
self, to have this great master beside them in every stage of de-
velopment. It was his hand which guided my first steps as a medical
student. * * * But how can one tongue adequately praise a man
who presided over the whole domain of the science of natural life ; or
how can one tongue depict the master mind, which extended the
limits of his great kingdom until it became too large for his own un-
divided government ? * * * ^^Q hsiYe to inquire what it was that
raised Muller to so high a place in the estimation of his contempor-
aries ; by what magic it was that envy became dumb before him, and
by what mysterious means he contrived to enchain to himself the
hearts of beginners and to keep them captive through many long
years ? Some have said that there was something supernatural about
Muller, that his whole appearance bore the stamp of the uncommon.
That this commanding influence did not wholly depend on his extra-
ordinary original endownments is certain from what we know of the
history of his mental greatness."

Years of Discovery: Such was the mind from which Schwann de-
rived his inspiration. The middle of the nineteenth century was the
golden age — ^the Periclean age — of physiology in Germany. To quote
further from Schwann's biographer (Henle) : Those were great days.
The microscope had been brought to such a state of perfection that it
was available for accurate scientific observation. The mechanics of its
manufacture had besides just been simplified to such a degree that
its cost was not beyond the means of the enthusiastic student even
of limited means. Any day a bit of animal tissue, shaved off with a
scalpel or picked to pieces with a pair of needles might lead to im-
portant ground breaking discoveries."

After the publication of his work on the cell theory, Schwann
was appointed professor in the University of Louvain, where he re-
mained nine years, after which he received a similar appointment in
V 1 the University of Liege. His "Microscopical Researches into the Ac-
Jj cordance in the Structure of Plants and Animals," though of somewhat
cumbersome title, is one of the great classics of biology. He proves
the identity in structure of animals and plants by direct comparison
of their elementary parts. His conclusion is that "the elementary
parts of all tissues are formed of cells in an analogous, though very
diversified manner, so that it may be asserted that there is one univer-
sal principle of development for the elementary parts of organisms,
however, different and that this principle is the formation of cells."

Virchow and "Cellular" Pathology: Any account of the cell
theory must needs be incomplete with the omission of the name and
work of Rudolph Virchow. Virchow was bom in 1821 of humble
parentage, his father eking out a livelihood from the combined oc-

DISCOVERY OF PROTOPLASM 61

cupations of farmer and small shopkeeper. The son who received the
academic training of his day was of an active restless temperament.
Virchow's was a mind open to new ideas, of liberal and independent
views on medicine, politics and religion. Hi_s._open sympathies with
the reform tendencies in 1848 were such that he was bbliged^to leave
, Berlin 'for Wurzburg, where he taught pathology and did much orig-
Cinal work therein. He was recalled to Berlin in 1856, when he was
made professor of pathology in the university. The scope of his ac-
tivities may be seen when it is considered that he was also a member
of the Reichstag, where he became leader of the opposition and a
vigorous antagonist of Bismark. As chairman of the finance com-
mittee, Virchow is credited as the author of the Prussian Budget
system. He took a leading part in the politics of his city; and the
fact that from being one of the most unsanitary cities Berlin has
come to be one of the most healthful spots has been attributed in
great measure to his insistance on sanitary reform. Virchow stands
in much the same relation to pathology as Schwann to histology. He
has been called the "Father of Modern Pathology." He established^
"The true and fertile doctrine that every morbid structure consists of
cells which have been derived from pre-existing cells," or as he him-
self expressed it: "Omnis cellula e cellula." His chief work was his \
cellular pathology published in 1858; in it he applied the cell theory J
to diseased tissues. He died in 1903.

The cell theory incomplete as first announced: When William
Harvey published his discovery of the circulation, so complete was
his self-appointed task that little was left for future workers. The
glycogenic function of the liver is known and understood by us
practically as proclaimed by Claude Bernard. The cell doctrine has a
vastly different history. As announced by it's co-founders, it was
far from being complete. Among other inaccuracies they attached
too much importance to the cell wall. The word "cell" implies a wall-
ed enclosure. The cell of honeycomb or the cell of a penal institution
are examples which suggest themselves. The fundamental declara-
tion that all parts of plants and animals are built of similar units or
structures has been substantiated. This is'perhaps the only portion
of the theory that has not been profoundly changed.

The Discovery of Protoplasm: Perhaps of equal importance to
the cell-theory was the recognition of protoplasm. Huxley called it
"the physical basic of life." FeHx Dujardin recognized this sub-
stance, which is the basis of vital activity, in 1835. He discovered in
lower animal forms a jelly-like substance which he called "sarcode."
Dujardin was born in 1801 at Tours, France. He was trained to fol-
low the trade of his father, namely, that of watchmaker, and the
manual dexterity thus acquired served him in good stead in the later
vocation of his life. He was an adept with the microscope and pos-
sessed no small ability as sketch artist. He showed early a love for
the natural sciences. His contributions to science cover a range of
topics. He was perhaps the greatest authority of his day on proto-
zoology. He died in 1860.

Schleiden saw protoplasm but called it gum. Cohn, in 1850,
taught that "protoplasm" of plants and "sarcode" of lower animal
life were the same thing. Max Schultze, in 1861, confirmed Cohn's

62 PATHFINDERS OF PHYSIOLOGY

position and added that the cell consisted of little units of protoplasm
surrounding" a nucleus. The nucleus was first described by Fontana,
in 1871. It was regarded as a normal element of the cell by Robert
Brown in 1883. It was eventually seen that many cells, especially
animal cells, are without a cell wall, hence the conclusion that the so-
called "wall" is not an essential feature of the "cell." When the cell
wall is absent the protoplasm is the cell. The nucleus was found to
be within the substance of the cell and not within the cell wall.
Schultze -defined the cell as a globule of protoplasm surrounding a
nucleus. From being regarded as an element of structure merely, the
cell has come to be recognized as the physiological unit within which
all physiological activity takes place.

Perhaps the most authoritative as well as the most recent defi-
nition of protoplasm is the following significant paragraph by Star-
ling:

"Though it may be convenient to have a word such as protoplasm signifying
simply 'living material,' it is important to remember there is no such thing as a
single substance — protoplasm. The reactions of every cell as well as its organiza-
tion are the re,sultant of the molecular structure of matter of which it is built up.
The gross methods of the chemist show him that the composition of the proto-
plasm of the muscle cell is entirely different from that of a leucocyte or white
blood corpuscle. The finer methods of the physiologist show him that every sort
of cell in the body has its own manner of life, its own peculiarities of reaction
to uniform changes in its surroundings. No individual will react in exactly the
same manner as another individual even of the same species, and the reactions
of the whole organism are but the sum of the reactions of it's constituent cells.
There is not one protoplasm therefore, but an infinity of protoplasms and the use
of the term can be justified only if we keep this fact in mind and use the word
merely as a convenient abbreviation for any material endowed with life. Even-
in a single cell there is more than one kind of protoplasm. In its chemical
characters, in its mode of life, and in its reactions, the nucleus differs widely
from the cytoplasm. Both are necessary to the life of the cell and both must
be regarded according to our present ideas as 'living.' In the cytoplasm itself
we find structures or substances which we must regard as on their way to proto-
plasm or as products of the break down of protoplasm; but in many cases it is
impossible to say whether a given material is to be regarded as lifeless or as re-
active living matter. Even in a single cell we may have differentiation among
its different parts, one part serving for the process of digestion while other parts
are employed for purpose of locomotion. Here again there must be chemical
differences, and therefore dtflerent protoplasms."

A statement of the cell theory at the present time (1913) must
include four conceptions: (1) The cell as a unit of structure; (2) The
cell as a unit of physiological activity; (3) The cell as embracing all
hereditary qualities within its substance; (4) The cell in the histori-
cal development of the organism."

Students of cytology have sought to find out if any uniformity
of organization of protoplasm exists. Accordingly we have a number
of explanations or theories regarding its structure. Altmann pro-
posed the granular theory. By the employment of certain hardening
reagents he demonstrated dense masses of spherical or rod-shaped
granules in all the cells of the body. In these he located the various
vital functions, the sum total of which constitute the life of the cell.

THE NUCLEUS 63

He further maintained that these granules could come only by division
of pre-existing granules. He parodied Virchow's famous phrase
omnis cellula e cellula into omne granulum e granulo.

The fibrillar theory presupposes net-work or clusters of fibrils
known as "spongio-plasm" (sponge plasm) in contra -destinction to
clear or structureless matter filling in the meshes of the net to which
the name "hyaloplasm (glass plasm) has been given.

In the Alveolar Theory of Butschli the author regards the so-
called granules as products manufactured by the hyaline protoplasm
and stored up as spherules so that the protoplasm between the drop-
lets form an alveolar partition — hence the name of the theory.

Discussing the question as to the fluJdityLQ£_proto^asm Starling
regards it as "essentially :fluidm fihAracter, the form andligidity
which are acquired by most cells being due to chemical and physical
differentiation occurring in its fluids/*

The cell consists of cytoplasm and nucleus. Cytoplasm (cell plasm)
is a term formulated by Kolliker in 1863. Though not so applied
when first used, it has come to mean the living substance of the cell
body other than the nucleus. Cytoplasm contains, for the most part,
substances apparently foreign to the cell proper. In the cytoplasm
of plant cells, for example, are stored up starches and oils. Most
nerve cells contain various shaped bodies which, it is alleged, repre-
sent stored up energy. The passive bodies in the cytoplasm are sup-
posed to represent some form of latent energy upon which the cell
may draw. In the cells of any green leaf are to be found spherical
masses which play a most important role in the lives of not only plants
but of animals as well. By the action of the sun's rays a chemical
change takes place in these bodies known to botanists as chloroplasts
by which carbondioxide and water are broken down, decomposed and
immediately synthetized into a different substance — carbohydrate,
starch, which will respond to the well known iodine test for starch.
Carbohydrate is one of the food principles. Fats are also made and
stored in the form of oils. In spite of the fact that the atmospheric
air surrounding the plant contains an abundance of free nitrogen.
The plant cells are unable to make use of it. Nitrogen must be first
combined as a nitrate, become dissolved in the soil and taken up by
the roots of the plants, or in the case of water plants, by special cells,
before the green matter in the leaf can be transformed into protein.
The plant, therefore, has power to make foods out of the
chemical elements of air and water when these elements are
properly combined. This is the only source of food of both plant
and animal and it is the result of cellular activity.

The Nucleus: The nucleus has been recognized as a most es-
sential part of the cell. It not only takes part in the complex process
of cell division but dominates the rest of the cell. It is not my pur-
pose to enter upon a discussion of the morphology and physiology of
the animal and vegetable cell, further than it is necessary to trace the
various stages of the history of its revelation from its earliest recog-
nition to the present. The reader is referred to the numerous excel-
lent text books on the subject.

64

PATHFINDERS OF PHYSIOLOGY

ILLUSTRATIONS SHOW DIAGRAMATICALLY THE CELL AND
INDIRECT CELL DIVISION.

Fig. 1.

Fig. 2.

Fig. 3.

Fig. 4.

Fig. 5.

Fig. 6.

Fig. 7.

"The first change in the appearance of the nucleus which indicates that a
division is about to take place, consists in a rearrangement of the chromatin net
work, which now takes place on the appearance of a tangled thread (Fig. 2).
The outwardly directed loops of this skein often correspond to the seperate por-
tions into which the thread eventually breaks up. The thread gradually grows
shorter and thicker, and presently becomes divided into a number of pieces known
as chromosomes. In the chromosomes the shortening and thickening process is
continued until these bodies arrive finally at the form of stumpy rods, each of
which, often becomes bent into the form of a horse shoe. Meanwhile the nuclear
membrane, breaks down, so that the hyaline substance of the nucleus becomes
continuous with that of the cell body surrounding it. A fresh phenomenon now
becomes visible. A spindle-shaped arrangement makes it's appearance consisting
of a number of minute fibrils which connect together two points — the poles of
the spindle — situated at opposite ends of the cell. The chromosomes now change
their position so that they come to be in the plane of the equator of the spindle,
and about this line each chromosome splits longitudinally into two great por-
tions (Fig. 4 and 5). This splitting in the case of each chromosome takes place
in the equatorial plane of the spindle, so that one member of each pair of daugh-
ter chromosomes faces towards one pole of the spindle and the second towards
the other pole. The members of each pair of daughter chromosomes now begin
to move away from one towards the two poles of the spindle, and as they do so
the first indication of a dividing wall between the second new cells begins to
make its appearance in the equatorial plane. Arriving at the poles, the daughter
chromosomes begin to elongate and to put out processes which finally meet and
fuse with those of their neighbors to form the chromatin reticulum of the new
nuclei. (Fig. 7.) Surrounding each new nucleus, thus developing at either pole
of the now rapidly disappearing spindle, a new nuclear membrane makes it's ap-
pearance; the dividing wall in the position of the equator of the spindle develops
into a complete partition in tfie case of plants. (The animal cell is without a
cell wall.) The division into two new cells is thus completed. (Fig. 8.) Each
new cell is provided with a nucleus into which has entered precisely its fair
share of the chromatin which was present in the parent nucleus."
— Illustration and description after Locke.

THE CELL IN HEREDITY 65

The discovery of the various dyes and tissue stains afforded a
wonderful stimulous to the microscopic study of tissues as well as to
bacteriological studies. It is hard to conceive of much progress in
bacteriology without this aid. Dyes for staining protoplasm were first
prepared in 1868. The property of taking up a stain gave rise to the
invention of a number of new names for which scientists have as
usual drawn freely from the Greek. To designate that protoplasm
which stained deeply, we have the term "chromatin." The word
"achromatin'* has been applied to protoplasm," which will not absorb
the dye. Certain rod-shaped bodies situated within the nucleus,
which stain more deeply than any other portions are known as
"chromosomes."

The Cell In Heredity — Within recent years the subject of here-
dity has claimed the attention of biologists and its practical applica-
tion has become of intense interest to the laity, advances in our
knowledge of heredity are already producing results. They have
revolutionized agricultural methods as shown in the marked improve-
ment of animals and plants. It is impossible of realization what are
the potentialities in regard to the improvement of the human race.
Eugenics is as yet in its infancy. The past ten years has witnessed
the production of voluminous literature on eugenics and its kindred
subject heredity.

Smallwood in his latest work states that "Whatever may be
the ultimate analysis of the problem of heredity, there can be no
hesitation in stating that the transmitted characters exist potentially
in the protoplasm of the cell. From the egg of a robin only a robin
will develop, from the ovum of an oak only an oak will grow and dur-
ing the growth each follows its own successive developmental stages
even to the minutest details. It has been well said ^nature never yet
made two eggs or two sperms exactly alike.* The cells which give
rise to new organisms are the germ cells, sperms and ova. These
differ greatly in shape and size — some of the sperm cells being but
one one-hundred-thousandths the bulk of the ovum and yet the pa-
ternal characters are easily recognized in the adult. * * * The
cells of the body are divided into body plasm and germ plasm." Germ
plasm might be looked upon as the immortal in man in as much as it
is continuous. After the germ plasm has given rise to a new individ-
ual, some of it is left behind to participate in the formation of a new
offspring, so as Davenport puts it, "There is really no inheritance
from parent to child but^^rent and child resemble each other because
they are derived from the same plasm, they are chips of the same
old block; and the son is half-brother of the father by another
mother."

As the cell has been called "The physiological unit," and proto-
plasm "the physical basis of life," the chromosomes have been proven
the physical basis of heredity. They are very definite and import-
ant organs. The number which make their appearance at each cell
division is the same in all the cells of any given creature and is con-
stant for the cells of the members of any given species.

"The remarkable fact," says Wilson, "has been established that
every species of plant or animal has a fixed and characteristic num-

66 PATHFINDERS OF PHYSIOLOGY

ber of chromosomes, which regularly occurs in the division of all of its
cells, and in all forms arising by sexual reproduction, the number
is even/'

Whatever the offspring is, it is potential in the fertilized ovum.
If this is the contribution of each parent, the role performed by the
mother is that of custodian of her embryonic charge until birth. Her
power- to alter it in any way is as futile as that of the father. The
parent is rather the trustee of the germ plasm than. .the- producer of
the child. Sir Michael Foster once said, "The animal body is in reality
a vehicle for the ova; and after the life of the parent has become
potentially renewed in the offspring, the body remains as a cast-off
envelope whose future is but to die." The germ plasm is "the lighted
torch handed on from one runner to another." Et quasi cursores
vitai lampada tradunt. This equally true of plant life, where the
plant matures and dies leaving the future offspring potentially in the
seed. How characteristics are transmitted from ancestor to offspring
is not known.

NOTE: — It has been estimated that the number of cells entering into the
composition of the body of an adult human being is about twenty-six million
five hundred thousand millions (26,500,000,000,000).

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