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3 1924

THE CELL DOCTRINE.

BY THE SAME AUTHOR.

A GUIDE TO THE PRACTICAL EXAMINATION OF URINE.
For the use of Physicians and Students. Illustrated. Second
edition, revised and improved. Just ready. Price, $1.25.

AN INTRODUCTION TO THE STUDY OF PRACTICAL HIS-
TOLOGY. For Beginners in Microscopy. Price, $1.00; inter-
leaved, $1.50.

IN PREPARATION.

A TREATISE ON DISEASES OF THE KIDNEYS, with especial
reference to Pathology and Therapeutics.

Fig. 4. Fig. 2.

Youngest layer.
Production of formed material from germinal mater in Epithelial cells, trom section through layer of Epithelium
covering papillw of the tongue. ><700.

FORMATION oa PUS.

Yo illustrate the change in germinal matter of an i el, ri 5 from
manner in which the germinal macfer or a normal cell, if pagal Teale with ete may ae e rise to pus,

Fig. 13."

a, Sarcolemma, 6, Contshetile matter.
MUSCLE,

Young. Fully forr-2. Young. Fully formed.
TENDON. CARTILAGE. Ng. 105

+ Fig. th

ELASTIO TISSUE,
The arrow shows the direction in which germinal
matter is supposed to be moving.

wid, 46.

NERVE.

Development of young, dark-bordered herve fibres, at an early
period, show{ng germinal matter and formed material of

elementary parts, *<1800,
Fig. 17,
Oldest part of formed
‘ial.

New centre or nucleolus. mere

Youngest part of
Germinal matter(nuclens). formed material,

.
AMG@EBA,
C .
Pure germinal matter, 5000. ourse of pabulum.

PLATE ILLUSTRATING DR. BEALE’S VIEWS,

H. Sebald, Bo.

THE CELL DOCTRINE:

ITS HISTORY AND PRESENT STATE.

FOR THE USE OF

STUDENTS IN MEDICINE AND DENTISTRY.

ALSO,

A COPIOUS BIBLIOGRAPHY OF THE, SUBJECT.

By JAMES TYSON, M.D.,

PROFESSOR OF GENERAL PATHOLOGY AND MORBID ANATOMY IN THE UNIVER-
SITY OF PENNSYLVANIA; ONE OF THE VICE-PRESIDENTS OF THE
PATHOLOGICAL SOCIETY OF PHILADELPHIA ;

ONE OF THE VISITING PHYSICIANS TO THE PHILADELPHIA HOSPITAL; FELLOW
OF THE COLLEGE OF PHYSICIANS, PHILADELPHIA ;

ETC., ETC., ETC.

SECOND EDITION,
REVISED, CORRECTED, AND ENLARGED.

ILLUSTRATED.

PHILADELPHIA:

LINDSAY & BLAKISTON.
1878.

/ CORNELL
‘UNIVERSITY}|
\\ LIBRARY,

Entered according to Act of Congress, in the year 1878,
By LINDSAY & BLAKISTON,

In the Office of the Librarian of Congress, at Washington, D. C.

CAXTON PRESS OF

SHERMAN & CO,, PHILADELPHIA.

ST

TO

THE MEDICAL CLASS

OF
e

THE UNIVERSITY OF PENNSYLVANIA,

This Little Volume

IS RESPECTFULLY INSCRIBED,
BY

THE AUTHOR.

PREFACE

TO THE SECOND EDITION.

Tue highly favorable and quite unexpected reception
accorded the first edition of this book has stimulated my
interest in the subject, and in preparing a second, I have
sought to improve it as much as possible. In doing so,
many of the original sources of my information have
been re-examined, and from them some additions made
and inaccuracies corrected. The section on the Present
State of the Cell Doctrine, incorporating my own views,
has been entirely rewritten, as was necessitated by the
very important and numerous contributions to the subject
since the first edition appeared.

The bibliography has been increased by the addition
of over three hundred and fifty new references, mostly
to papers contributed directly on the subject since the
first edition was issued. To make room for these, many
of the references included in the bibliography of the
first edition have been omitted where there did not seem
to be a sufficiently close bearing on the subject.

vill PREFACE TO THE SECOND EDITION.

The suggestion of one of the reviewers of the first edi-
tion, that the bibliography should be chronologically in-
stead of alphabetically arranged was carefully considered,
‘and at one time I had concluded to adopt it, but when
I attempted to do so, I found that the inconvenience
resulting from the wide separation of several papers by
a single author, more than offset the advantages of a
chronological arrangement. I therefore adhered to the
original plan.

For valuable assistance in collecting references and
examination of papers I am greatly indebted to my as-
sistant, Dr. H. F. Formad.

1506 Spruce STREET,
October 1st, 1878.

PREFACE

TO THE FIRST EDITION.

THE author has become convinced, by several years’
intimate intercourse with students of medicine, that their
acquaintance with the subjects he has endeavored to in-
clude in this little volume would be facilitated, if the
views, which are now taught and scattered throughout
the often expensive works of their authors, were collected
in a convenient form for study and reference. Taking
it for granted that a knowledge of this subject is of fun-
.damental importance in its bearing upon the study of
physiology and pathology, and stimulated by the frequent
inquiries of students for an appropriate source of infor-
mation, he has prepared what he now submits to them.

He has sought to obtain a continuous history of the
evolution of the “cell doctrine” up to its present state,
without embarrassing his pages with a large number of
isolated facts. He has attempted, however, to secure a
completeness, and to make the work useful to physicians

x PREFACE TO THE FIRST EDITION.

-

and others engaged in research, by careful references, and
the addition of a bibliography, which he has sought to
make accurate and extended. Some authors may have
been overlooked; such the writer cordially invites to
send him references to their own papers, or to those of
others they believe to have a bearing upon the subject.

ILLUSTRATIONS.

Prate.—IuuustRatinc Dr. Braun’s VIEWS.

Figs. 1 to 7. Production of formed material from germinal matter in
epithelial cells, from section through layer of epithelium
covering papille of tongue.

Figs. 7 to 11. Formation of Pus.

Fig. 11. * “ Tendon.

Fig. 12. f “Cartilage.
Fig. 13. " “« Muscle.

Fig. 14, “ “Elastic Tissue.
Fig. 15. o “Nerve. .

Fig. 16. Ameba,
Fig. 17. Illustrating Nutrition of Cell.

INTERCALATED.

Fig. 1. Illustrating Globular Theory. After Virchow.

Fig. 2. Cellular Tissue from the Embryo Sac of Chameedorea Schie-
deana in the act of formation. After Schleiden.

Fig. 3. From the Point of a Branchial Cartilage of Rana Esculenta.
After Schwann. °

Figs. 4 to 12. Formation of Nuclei and Cells from Molecules, accord-
ing to Bennett.

Fig.

Fig.

Fig.
Fig.

Fig.

. 14,

15.

f6.

17.
18,

19.

ILLUSTRATIONS.

. Diagram of the Investment Theory. From Virchow.
. 13.

Formation of Pus from subcutaneous connective tissue.
From Virchow.

Formation of Pus from interstitial connective tissue of mus-

cle. From Virchow.

Development of Cancer from connective tissue. From Vir-
chow.

Connective Tissue Corpuscles anastomosing one with the
other. From Virchow.

Formation of Elastic Tissue, according to Virchow.

Formation of Connective Tissue, according to Schwann and
Henle. From ‘Virchow.

Formation of Connective Tissue, according to Virchow.

From Virchow.

THE CELL DOCTRINE.

Tue idea that animals and plants, however com-
plex their organization, are really composed of a
limited variety of elementary parts, constantly re-
curring, was appreciated by Aristotle, who was born -
384 years before Christ, while it appears to have
been little more clearly conceived by the acknowl-
edged father of medical science, Galen, who lived
400 years later. Aristotle distinguished as “ partes ~
similares,” those structures, such as bone, cartilage,
fat, flesh, blood, lymph, nerve, ligament, tendon,
membrane, vessels, nails, hairs, and skin, which
were not confined to one part of .the body, but
distributed throughout it generally. He applied
the term “partes dissimilares” to the regions of
the head, neck, trunk, and extremities. Fallopius
of Modena, 1523-1562, to whom we are indebted for
our knowledge of the conceptions of Galen in regard
to these “ partes similares” or “simplices,” has fur-
ther developed the subject of general anatomy in his
“‘ Lectiones de Partibus Similaribus Humani Corpo-
ris.” These, however, plainly do not correspond
with the “elementary parts” or “ cells” of the pres-
ent day. As Prof. Huxley says in his valuable essay
‘ 2

14 THE CELL DOCTRINE.

on “ The Cell Theory,” they were ultimate to Fallo-
pius, because he could go no further, “though it is,
of course, a very different matter whether we are
stopped by the imperfection of our instruments of
analysis, as these older observers were, or by having
really arrived at parts no longer analyzable.”* These
“ partes similares ” really correspond to the “ tissues”
of the present day, which are collections of elementary
parts. The conceptions of these older writers with
regard to the “ vital endowment” or “independent
vitality ” of their similar parts or tissues, were sin-
gularly correct, and correspond almost identically
with those held by the majority of physiologists of
the present day.

Further than this, however, the anatomists of the
period of Fallopius could not go—not because, as we
now well know, they had arrived at parts no longer
analyzable, but because of their imperfect means of
analysis. :

It is probable that the magnifying properties of
lenses were known to the Egyptiaus, as well as the
Greeks and Romans, over 2000 years ago; since
a table of refractive powers is introduced into
his “Optics” by Ptolemy, since Aristophanes, the
Athenian poet (B.C. 500), speaks of “burning
spheres” of glass as sold in the grocers’ shops of
Athens, and since both Pliny and Seneca refer to
lenses and their magnifying properties ; while lenses
themselves have been found in the ruins of Nineveh,

* The Cell Theory—a Review, by T. H. Huxley ; Br. and For-
eign Med. Chir. Rev. for October 1853, No. xxiv.

THE CELL DOCTRINE. 15

Hercnlaneum, and Pompeii. But it is quite certain,
also, that they did not become available as com-
pound inicroscopes until about 1590, when the Jan-
sens, father and son, of Holland, are said to have in-
vented the compound microscope. Fontana, in 1646,
writes that he had invented the microscope in 1618.
Galileo, as early as 1612, is said to have sent a micro-
scope to King Sigismund of Poland, though whether
it was his own invention, or made after the pattern
of another, is not easily determined. In 1685,
Stelluti published a description of the parts of a bee
he had examined with the microscope, and although
George Hufnagle is said to have published in Frank-
fort, in 1592,a work upon insects, illustrated by fifty
copper plates, it is highly probable that these, as well
as very many most important observations made after
the invention of the compound microscope, were
made with the simple instrument.*

It is impossible to estimate the assistance the
microscope has been to us in opening up the minute
structure of animals and vegetables, and in thus af-
fording a reliable basis on which to build a doctrine
of organization. Prof. Iuxley ‘further says, ‘ The
influence of this mighty instrument of research upon
biology, can only be compared to that of the galvanic
battery,in the hands of Davy,upon chemistry. It has
enabled proximate analysis to be ultimate.”+ But it
is more than this. Since, as he correctly states, it

* For an interesting and exhwustive history of the invention of
the compound microscope, see Das Mikroskop, Theorie, Gebrauch,
Geschichte nnd gegenwartiger Zustand desselben. Von P. Harting.
In drei Banden. Braunschweig, 1866. Dritter Band, ss. 1-85.

+ Huxley, loc. citat., p. 290.

%

16 THE CELL DOCTRINE.

has enabled proximate physical analysis to become
ultimate, it corresponds, not to the galvanic battery
alone, but to all the appliances made use of in ulti-
mate chemical analysis.

The time prior to the invention of the compound
microscope may be considered as the first period in
histology ; that between this date and that of the ob-
servations of Schleiden and Schwann (1838), inclu-
sive, the second period ; while the time subsequent to
these observations becomes appropriately the third
period. Notwithstanding the imperfect state of in-
struments during-quite two hundred years after the
invention of the compound microscope, a flood of
facts was added to our knowledge of the minute
structure of living things.

Borellus, of Pisa, seems first to have used the mi-
croscope in the examination of the higher animal
structures, about the year 1656, but his observations
were grossly misinterpreted in his attempt to adapt
them to the prevailing idea of the day, that diseases
were caused by animalculee in the blood and tissues.
Asa result, he describes pus-corpuscles as animalcules,
and even says he has seen them delivering their eggs.

According to Boerhaave, Swammerdam had recog-
nized the blood-corpuscle in the frog in 1658.

Malpighi,* between 1661 and 1665, had seen the
blood-corpuscle‘in the hedge-hog, had witnessed the
circulation of the blood, and had published observa-
tions upon the minute structure of the lungs, which
he had even compared to a racemose gland,t of the

* Malpighi, Opera Omnia. London, 1686.
+ Fort, Anatomie et Physiologie du Poumon, considere comme

THE CELL DOCTRINE. 17

kidneys, spleen, liver, and membranes of the brain,
and with some of these structures his name has
become inseparably associated. In 1667, Robert ‘
Hooke* pointed out the cellular structure of plants,
and Malpighit further elaborated the same subject
with considerable accuracy in his “ Anatome Planta-
rum,” in 1670, He showed that the walls of the
“ cells” or “ vesicles,” were separable, that they could
be isolated, and gave to each the name “ utriculus,”
believing also the “cell,” or “utriculus,” to be an
independent entity. The latter observert also recog-
nized the blood-corpuscle. Leeuwenhoek, in 1673,§
described these corpuscles with considerable accuracy,
not only in man, but also in the lower animals. He
also demonstrated the capillaries, examined most of
the tissues, and made the discovery of the sperma-
tozoids, which he conceived to be spermatozoa or
sperm animals, and of different sexes.

Theory of Haller, 1757.—No attempt, however,
seems to have been intelligently made at building up
the tissues by an ultimate physical element, to cor-
respond with the ‘atom ” of the inorganic chemist,
prior to that of Haller. He resolved the solid parts
of animals and vegetables into the “ fibre” (fibra),

un organe de Secretion. Puris, 1867, Preface; or a notice of Dr.
Fort’s book, by the writer, in American Journal of Medical
Sciences, October, 1869. 2

* Hooke, Rob., Micrographia. London, 1667.

+ Malpighi, Anatome Plantarum. London, 1670.

t Malpighi, Opera Posthuma. London, 1697.

@ Leeuwenhoek, Opera Omnia seu Arcana Nature detecta.
Tom. ii, p 421. Leyden, 1687. Vel Opera Omnia, etc., Lugd.
Batav., 1722.

Qx

18 THE CELL DOCTRINE.

and an “organized concrete.” To the former he as-
signs the most important position, asserting that it
is to the physiologist what the line is to the geome-
trician; that a “fibre,” in general, may be con-
sidered as resembling a line made up of points,
having a moderate breadth, or rather as a slender
eylinder.*

The second elementary substance of the human
body according to Haller, the “ organized concrete,”
must not be lost sight of, as appears to have been
the case with many eminent authorities who have
attempted to give his views. This, he says,is a
mere glue, evasated and concreted, not within the
fibres, but in the spaces betwixt them, in illustration
of which it is stated, that cartilages seem to be
scarcely anything else besides this glue concreted.
But these views of Haller were clearly not based upon
microscopic observation, though the microscope had
been for some time in use. For Haller himself tells
us that the fibre is invisible, and to be distinguished
only by the “mind’s eye,”—invisibilis est ea fibra,
sola mentis acie distinguimus.t No allusion.to the
cell beyond the imperfect description of the blood-

* Haller, Elementa Physiologia, vol. i, lib. i, sec. i, Lausan.,
Helvet., 1757.

+ Asingular discrepancy exists between these words of Haller
and those found in both the Latin and English editions of the
‘elegant compend”’ of Haller’s works printed in Edinburgh, the
former in 1766, and the latter (an edition in the possession of the
writer), in 1779, under the inspection of William Cullen, M.D.
In the latter, we have the following: ‘ The solid parts of animals
and vegetables have this fabric in common, that their elements, or

THE CELL DOCTRINE. 19

corpuscles and spermatozoids appears to have been
made by Haller. .
Theory of Wolff, 1759-74.—Better founded, in
being based upon observation, was the theory of
Wolff, and it contained many of the elements of
truth. For an available exposition of these views,
physiologists are much indebted to Prof. Huxley,
who in the able review already cited, has pre-
sented them as agreeing partially, also, with his
own. The doctrine of Wolff, as given by Prof.
Huxley, is as follows: “Every organ is composed, at _-
first, of a mass of clear viscous, nutritive fluid, which
_ possesses no organization of any kind, but is at most
composed of globules. In this semifluid mass, cavi-
ties (Blaschen, Zellen) are now developed ; these, if
they remain rounded or polygonal, become the sub-
sequent 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 in the
young budding organs of an animal. Both cells and
vessels may subsequently be thickened by deposits
from the ‘ solidescible ’ nutritive fluid. In the plant,
the cells at first communicate, but subsequently be-
come separated from one another; in the animal,
they always remain in communication. In each case..
they are mere cavities and not independent entities ; or-
ganization is not effected by them, but they are the visible
results of the action of the organizing power inherent in

the smallest parts we can see by the finest microscope, are either
fibres or an organized concrete.”’!

1 First Lines of Physiology. By the celebrated Baron Albertus Haller, M.D,
Translated from the correct Latin edition, and printed under the.inspection of
William Cullen, M.D, Edinburgh, 1779,

20 THE CELL DOCTRINE.

the living mass, or what Wolff calls the vis essentialis,
For him, however, this vis essentialis is no Archeus,
but simply a convenient name for two facts which
he takes a great deal of trouble to demonstrate ; the
first, the existence in living tissues (before any pas-
sages are developed in them), of currents of the nu-
tritious fluid determined to particular parts by some
power which is independent of all external influence ;
and the second, the peculiar changes of form and
composition, which take place in the same manner.””*
-Two points are here particularly to be observed as
cardinal,——first, the non-independence of cells, either
anatomically or physiologically; that they are effects,
passive ‘results, and not causes of a vitalizing or or-
ganizing force ; second, that organization takes place
from the “differentiation” of the homogeneous
living mass in these parts, through the agency of the
vis .essentialis or inherent vital force. The radical
difference between these principles of development
and those generally held at the present day, will be
better appreciated when these latter have been
worked out. An acknowledged error may, however,
be pointed out,—the probable result of the inferiority
of the instruments of that day—-that of supposing
the cells of plants and animals in all instances to
communicate when in their youngest state, and in
the latter to continue thus in communication through-
out life. It will be observed, also, that this theory
involved the spontaneous origin of the cell, that is,
independent of any previously existing cell.

* Huxley, loc. citat., p. 298-4. Wolff, O. F., Theoria Genera-
tionis, 1759. Ed. Nova, Halae, 1774.

THE CELL DOCTRINE. 21

The theory of Wolff, however, full as it was of origi-
nal conception, and based on actual observation,seemed
to claim little attention, and would have been still
less Known but for the labors of Prof. Huxley. The
“ fibre” theory of Haller was still further expanded,
and that fibres were the groundwork of nearly all
the tissues, continued the prevailing view, until the ,
latter part of the eighteenth century, and there are
few of the older Physiologies even of a later date,
which do not contain an account of it. Naturally,
it maintained itself longest in the case of the fibrous
tissues, since the appearances of these tissues, when
examined by the highest powers, are those of struc-
tures apparently composed of fibres.

Oken, 1808.—The first clear expression with regard
to the cellular or vesicular composition of animal or-
ganisms as well as vegetable, comes from the physical
school in the language of Oken, who, as early as
1805, in his work on “ Generation,” refers to elemen-~
tary parts as “ vesicles;” and who says in his “ Pro-
gramm tber das Universum” in 1808, “ The first
transition of the inorganic to the organic is the con-
version into a vesicle (Blaschen), which I, in my
theory of generation, have called infusorium. Ani-
mals and plants are throughout nothing else than mani-
foldly divided or repeating vesicles, as I shall prove
anatomically at the proper time.” This most ex-
plicit statement seems also to have been overlooked,

The Globular Theory, 1779-1842.—The reaction
which took place at the date referred to against the

: “fibre” theory, culminated in the “ globular ” theory,
due less to speculation than erroneous methods of ob-

22 THE CELL DOCTRINE.

servation and imperfect instruments. Leeuwenhoek*
(1687) early announced the “ globular” structure of
the primitive tissues of the body, but the “ globule”
apparently attracted little notice until this period of
reaction against the “fibre,” when it claimed the
attention of Prochaskat (1779), Fontanat (1778), the
brothers Wenzel§ (1812), Treviranus| (1816), Bauer
(1818 and 1823), Heusinger** (1822), MM. Prevost
aud Dumas,jt Milne-Edwardstt (1823), Hodg-
kin§§ (1829), Baumgartner|| (1830-42), Frederick

* Leeuwenhoek, op. citat.

+ Prochaska, De Structura Nervorum, Vind., 1779. Opera
min., Pars i. :

{ Fontana, Sur les Poisons, 1787, ii, 18; Abbandlung tiber das
Viperngift, das Amerikanische Gift, u. s. w. Aus dem Italien.
Berlin, 1787.

@ Weuzel, Joseph and Charles. De structura cerebri. Tubing.,
1812,

|| Treviranus, Vermischte Schriften, Anatom. und Physiolog.
Inhalts Bd. i. Gottingen, 1816.

{ Bauer, Philosoph. Transac. for 1818, and Sir E. Home’s Lec-
tures on Comparative Anatomy, vol. iii, Lect. iii. London, 1823.

** Heusinger, System der Histologie. Thl.i Eisenach, 1822-4

+t MM. Prevost and Dumas, Bibliothéque Universelle des Sci-
ences et Arts, T. xvii.

tt Milne-Hdwards, Mémoire sur la Structure Elémentaire des
Principaux Tissues Organiques des Animaux. Paris, 1828. Also,
Recherches Microscopiques sur la Structure Intime des Tissues Or-
ganiques des Animaux, in Ann. des Sci, Nat. December, 1826.

#2 Hodgkin, in Grainger’s Elements of General Anatomy. Lon-
don, 1829. Also Hodgkin and Fisher’s translation of M. Edwards
“Sur les Agens Physiques.”” London, 1882. Hodgkin’s Lectures
on the Morbid Anatomy of the Serous and Mucous Membranes.
London, 1836, p. 26. Am. Ed., Philadelphia, 1838, vol. i, pp.
17-18.

|||| Baumgartner, K. H., Beobachtungen iiber die Nerven und
das Blut in ihrem gesunden und Krankhatten Zustande, February,

THE CELL DOCTRINE. 23

Arnold* (1836), Dutrochett (1837), Raspailt (1839) ;
all except Hodgkin admitting in greater or less de-
gree the importance of the globuleas an ultimate phys-
ical element; while it is evident, also, that there
was much confusion in the use of terms, the words
globule, granule, and molecule,§ being often indiscrimi- ”
nately used, and the word globule sometimes used
to indicate what is now clearly recognized as the
“ cell.”

1830. His views are further elaborated in his Beitrige zur Phy-
siologie und Anatomie. Aus der Lehre von der Gegensazen in
den Kraften in lebenden thierschen Kérper, ein Grundriss zur
Physiologie und zur allgemeinen Pathologie und Therapie, 2te
Aufluge, besonders abgedruckt. Stuttgart, 1842.

* Arnold, Friedreich, Lehrbuch der Physiolugie des Menschen.
Erst. Theil, Zurich, 1836.

t Dutroulet, Mémoires pour servir a l’Histoire Asatemigna et
Physiologique des Végétaux et des Animaux, t. ii, Atlas. Paris,
1837.

} Raspail, Recherch. sur la struct. et ledevelopm. de la feuille et
du trone, et sur la struct., et devel. des tissus Animale, Paris, 1887.

3 The German aiiglione of this period, and even more recent
times (Henle, 1841, Virchow, 1858), at least in speaking of the
development of histology, seem to use indiscriminately the terms
granule or molecule and glubule, whereas they are morphologically
something distinct. A globule is usually held to be a body which,
under the microscope, is more or less spherical in form, possessing
a bright centre, and dark outline,—the width of this outline being
directly as the difference between:the refracting power of the
globule itself and that of the menstruum in which it floats. Thus,
the dark outline of a globule of oil floating in water is wider than
that of the same glubule floating in glycerin. A granule or
molecule, on the other hand, is indeterminate in size and shape, and
appears as a mere dot under the highest powers of the microscope.
It is true that what appears as a granule under a low power, may
appear as a glubule under a higher.

24 THE CELL DOCTRINE.

Prochaska,* in 1779, described the brain as made
up of globules eight times smaller than blood-glob-
ules. In the year 1801, the philosophic mind of
Bichat elaborated his excellent classification, but he
seems to have made no original investigations in
minute structure, or to have adopted any special
theory of an ultimate physical element. The bro-
thers, Joseph and Charles Wenzel,t in 1812, de-
scribed the brain as composed of. globules of small
size. Among the earliest histologists worthy of
mention, is Treviranus,{ whose elements, according
to Henle, were first, a homogeneous, formless matter ;
second, fibres; third, globules (kiigelchen). Mr.
_ Bauer,§ quoted as a most experienced microscopic
observer by Sir Everard Home, in 1818, and again
in 1823, described the ultimate globules of the brain
and of muscular fibre as of the size of a globule of
blood when deprived of its coloring matter, or about
aoa Of an inch in diameter. The fibre was excluded
as an ultimate element of organization by Heusinger|
in 1822-4, who started al] tissues from the globule,
still, however, retaining the formless material of
_ Haller and Treviranus. Heusinger formed the fibre
by the linear apposition of his globular elementary
parts, and even explained how canals and vessels
were formed by a similar arrangement of vesicles
which had originated from the globules. The ac-
count given by Henle of the method in which Heu-

* Proschaska, Opera Minora, Part I, p. 342

+ Wenzel, op. citat., p. 24, { Treviranus, op. citat.

% Bauer, op. citat. || Heusinger, op. citat., p. 112.
{| Henle, Allgemeine Anatomie. Leipzig, 1841, p. 128.

THE CELL DOCTRINE. 25

singer built up his fibres and vessels is interesting
and important, since there is in these views an ap-
proximation to the truth. “As the result of an
equal contest between contraction and expansion,
there arises the globule, of which all organisms, all
organic parts, are originally composed. By a stron-
ger exercise (Spannung, tension) of power, there
originates from the often more homogeneous globule,
the vesicle. Where in an organism globules and a
formless mass are present, the globules arrange them-
selves according to chemical(?) laws and form fibres.
Where vesicles arrange themselves, there arise canals
and vessels.” In the latter sentence one cannot fail
‘to note a close approximation to the truth, though
the facts upon which the theory was based are partly °
false and partly misinterpreted.

But the observations and writings of ‘Milne Ed--

wards* may be looked upon as having given, more
than those of any other author, position and popular-
ity to the “globular theory.” He examined all the
principal tissues, and announced that the fibres of the
then so-called cellular (fibrous) tissues, membranes
composed of these fibres, muscle and nerve, were
composed of globules of about the same size, from
avav tO zslyy Of an inch in diameter; whence he
concluded that these spherical corpuscles, by their
aggregation, constituted all organic textures, vege-
table or animal, and whatsoever their properties or
functions. There’ is little doubt but that many of
these so-called globules described by Edwards were

* Edwards, loc. citat.
8

26 THE CELL DOCTRINE.

really cells, seen with indifferent instrumeuts, and
further distorted by the glare of direct sunlight.

Similar, as regards the element of organization,
were the views of Baumgirtner* and Arnold,t who
were joint observers.

They consideredt the fundamental elements of or-
ganization to be the formative globule (Bildungskugel',
and the molecular granule (Molecular-kugelchen).
The first is primarily formed by a simple aggrega-
tion of smaller granules first represented by the gran-
ules of the yolk united by a formless material. The
“molecular granule” arises from a breaking up of
the “formative globule.” A modification of the
“ formative globule” out of an aggregation of which’
the entire embryo is first formed, is the hematoid
body (Hamatoidkorper). This is a nucleated dis-
coid body with a distinct ring-like border (geringtes
Kérperkern mit einem Ringe).

Further, “Out of these two kinds of globules and
out of formless material,” says Baumgartner, “all
tissues are formed, namely, the tissue-fibres (threads)
out of the molecular granules, and the hematoid
bodies out of the formative globules and newly-
formed tissue-fibres. The molecular form is not
everywhere equally expressed, which is owing in the
first place to the fact that often the molecular gran-

* Baumgirtner, loc. citat.; also, Virchow, Cellular Pathology,
Am. Ed. of Chance’s Translation. Philadelphia, 1863, p. 53.
+ Arnold, loc. citat.; also, Virchow, Cellular Pathology, Am.
Ed. of Chance’s Translation. Philadelphia, 1863, p. 53.
| f Baumgiirtner, Beitrage zur Physiologie und Anatomie, 1842,
p. 36. :

THE CELL DOCTRINE. 27

ules more or less fuse together, and second, that
formless material surrounds the molecular granules
and makes their ontline indistinct. The theory here
brought forward of the fundamental form of organi-
zation in animals may well be called the globular
theory.” P. 88.

It is evident that the “formative globule,” or at
least the modification of it called the Hamatoidkér-
per, is nothing more nor less than the nucleated cell,
which, however, Baumgirtner did not admit, contend-
ing as lateas 1842 against the cell doctrine ; asserting
also (p. 40, op. citat.) that in the development of tissues
the formative globule never divides to form two, in
other words, that there is no such thing as cell di-
vision.*

Arnold also says thatt all fluid and solid parts of
the human body are resolvabie first into a fluid or
half fluid material of no determinate form, and sec-
ond, into granules which are more or less completely
spherical, and in all solid structures appear for the
_ most part as minute globules. The granules, which
are the second more important element, occur not
only in all fluids and solid parts of the completely
formed human organism, but they are also the origi-
nal and essential constituents of the human embryo.
Out of these by their aggregation are formed the most
complex tissues of the organism.

* To the student desiring to pursue further this very interesting
subject, with the argument against the cell theory by Baumgirt-
ner, I would recommend the perusal of the very interesting ‘ Bei-
trige zur Anatomie und Physiologie” alluded to.

+ Lehrbuch der Physiologie des Menschen, 1836, p. 82.

28 THE CELL DOCTRINE.

The error of these and other observers seems to
have been clearly pointed out by Dr. Hodgkin,”
though much importance was still attached to the

Fie, 1.

“0

Illustrating the Globular Theory.

A, Fibre, composed of elementary granules (molecular granules), drawn up in
a line. B, Cell, with spherically arranged granules. (After Virchow, slightly
modified.)

globule as an element of organization (but perhaps
from this time forward, more in the stricter sense of
the term granule), which has continued, in this latter
sense, to the present day.

It should be mentioned that in 1828 Déllingert

— gnnounced that the tissues of the body are built up

of blood-corpuscles, which move in wall-less (wandlos)
channels in these tissues.

From the foregoing facts it is evident that for
some time prior to the year 1838, the cell had come
to be quite universally recognized as a constantly re-
curring element in vegetable and animal tissues,
though as yet little importance had been attached to
it as an element of organization, nor had its charac-
ters been clearly determined. As stages in its grow~
ing importance may be mentioned, the demonstration
of the cellular structure of plants by Robert Hooke,

* Hodgkin, loc. citat,
{ Dollinger, Ignaz, Vom Kreislaufe des Blutes, 1828.

THE CELL DOCTRINE. 29

in 1667, the further elaboration of this subject by
Malpighi, and his statement that each “ utriculus ”
was an independent entity, the very clear statement of
Oken in 1808 with regard to the cellular composition
of animals and vegetables, the description of Heusin-
ger, in 1822, of the mode of formation of vessels by the
apposition of vesicles, already referred to, and the an-
nouncement, though erroneous, of Déllinger, in 1828,
that the body is built up of blood-corpuscles which
move in wall-less (wandlos) channels in the tissues.

Raspail, 1837.—Singularly near the truth did Ras-
pail* approach, in 1837, when he tells us that in the
condition of development there are vesicles or cells,
endowed with life and the property, almost unlimited,
of producing out of themselves other cells of the same
structure and similar endowments, of spherical form,
and capable of taking up oxygen when exposed to
the atmosphere ; that the cell membrane in its fresh
state is structureless. Yet he considers the organic
cell as made up of granules or atoms, spirally ar-
ranged about an ideal axis, comparing the cell with
the crystal, and speaks of organization as crystalliza-
tion in vesicles (crystallization vesiculaire).

Dutrochet, 1887.—Similar was the view of Dutro-
chet,t who divided the component parts of the body
into solids and fluid. The solids were formed by the
aggregation of cells of a certain degree of firmness ;
the liquids, as the blood, are also made up of cells
which, however, float freely among each other, an
there are also tissues in which the cells are so feebly|:

£

i

* Raspail, op. citat. f Dutrochet, op. citat.
3*

30 THE CELL DOCTRINE.

united, that one can scarcely tell in what class to
place them. The contents of the cell may be more
or less solid, but the highest degree of vitality is
only compatible with liquid cell contents. Muscular
fibres, and the remaining animal fibres, are cells much
elongated. And he considers the same general plan to
prevail in the animal and vegetable. The approach of
“both of these observers to the truth is striking.
Both, however, either failed to detect the nucleus or
to attach any importance to it. They failed also to
lay down a law of organic development, and their
views were soon forgotten.
v Discovery of the Nucleus, 1833.—-A most important
contribution to the anatomy of the cell was made be-
fore this, however, in the discovery of the “ nucleus,”
by Dr. Robert Brown, of Edinburgh, whose paper,
“ Organs and Mode of Fecundation in Orchidee and
Asclepiadex,” appeared in the Transactions of the
Linnean Society of London, in 1888. He failed, how-
ever, to appreciate its importance, though its dis-
, covery was another fact added to those necessary to
complete the data on which has been founded the
so-called ‘cell theory.”

Meyen, 1836.—Meyen* sought to establish the
opinion that the cell is formed of spiral fibres which
lie closely upon one another, founding his view upon
his own observation.

Since the discovery of the nucleus, by Dr. Robert
Brown, in the vegetable cell, it had been recognized
by many observers in various pathological, as well as

* Meyen, Pflanzenphysiologie, Bd. i, 1886.

ry

THE CELL DOCTRINE, 31

healthy animal cells, and in the germ cell or ovule of
birds, as early as in 1825, by Purkinje ;* while Pur-
kinje,f Valentin, and Turpin,§ had actually called
attention to the relations of the animal and vegetable
cell to each other.

The pre-existence of the nucleus, and the gradual
development of the cell about it, Valentin had at-¥
tempted to demonstrate in the case of pigment cells,
C. H. Schultz] in the blood-corpuscle, Rudolph Wag-
ner in the egg, and Henle in epithelium, all before
the work of Schleiden had appeared. Miller had
also insisted on the analogy between the cells of the
chorda dorsalis and vegetable cells. Valentin, too,
had said, when describing the nucleus.of epidermic
cells, which he was the first to point out, that they
reminded him of the nucleus of the cells of vege-
table tissues.** Not only this, but Armand de Qua-
trefagestt and Dumortiertt had actually observed the
origin of young cells from’ the full grown, in the

* Purkinje, J., Ev. Symbole ad ovi avium historiam ante ificu-
bationem, cum duobus lithographs. Vratis., 1825.

+ Purkinje and Raschkow, Meletemata circa Mammalium Den-
tium Evolutionem. Diss. Inaug. Vratis., 1835, p. 12.

t Valentin, Ueber den Verlauf und die Enden der Nerven, aus
den Nov. Act. Nat. Curios., vol. xvii; besondersabgedruckt. Bonn,
1836.

@ Turpin, Ann. d. Sci. Nat., 2 ser. vii, 207.

|| Schultz, C. H., Miiller’s Archiv fir Anatomie, Physiologie
und Wissenschaft. Med., p. evii, 1837.

{ Stricker, Manual of Human and Comparative Histology,
New. Syd. Soc. Translat., 1870, p. 1.

**® Valentin, Nov. Act., N. C. xvii, pt. I, p. 96.
+t Quatrefages, ‘Amiates des Sci. Nat., 2 ser. ii, p. 114.
tt Dumortier, Annales des Sci. Nat., 2 ser. vii, p. 129.

82 THE CELL DOCTRINE.

embryo of the freshwater snail, while Valentin had
furnished examples of the development of fibres
out of cells in the muscular fibres, and in the sub-
stance of the crystalline lens. In fact, as stated by
Dr. Waldo J. Burnett, in his admirable paper,* Val-
entin “ perceived the true physiological rélations of
eells as far as he well could without apprehending
the grand fact that the nucleated cell is the funda-
mental expression of organic forms.”

Virchow had also compared the whole organism
to a free state containing individuals endowed with
equal privileges if not with equal powers.{

SCHLEIDEN AND SCHWANN, 1838.

It was reserved for Schwann to accomplish this mas-
terstroke in observation and generalization, through
the intermediate results of Schleiden, without whose
observations on vegetable structures, the true position
of the cell would probably have remained undetected
for some time longer. Schleiden, in 1838, clearly
pointed out the formation of cells in vegetable struc-
tures, according to a single and uniform method, and
elaborated the theory of development of which the
cell was the unit, and which Schwann immediately
extended to animal tissues.

* Burnett, W. J., The Cell; its Physiology, Pathology, and
Philosophy, as deduced from original investigations. To which is
added its History and Criticism. A prize essay, read before the
American Medical Association, and published in vol. vi of its
Transactions. Philadelphia, 1853.

+ Stricker, op. citat., p. 2.

THE CELL DOCTRINE. 33

A formidable obstacle for some time in the way of
a law of development, applicable to animal and vege-
table tissues, was the opinion, long entertained, that
the growth of animals, whose tissues are furnished
with vessels, is essentially different from that of plants ;
an independent vitality being ascribed to the elemen- :
tary particles of vegetables growing without vessels.
So firmly was this believed, that the ovum, which
exhibited undoubted evidences of an actual vitality
at one period of its growth, was said by all physiolo-
gists to have hada plant-like growth. This obstacle
was removed in 1837, by Henle,* who showed that
an actual growth of the elementary parts of epithe-
lium took place without vessels.

Taking up the nucleus as discovered by Robert
Brown, Schleiden,t in reference to its function, ap-
plies the name cytoblast (zyts, a cell, @Aastos, a bud
or sprout), or “cell bud,” and in a careful study of
its anatomy, discovers that “in very large and beau-
tifully developed cytoblasts, there is observed a
small, sharply defined body, which, judging from the
shadow which it casts, appears to represent a thick
ring, or thick-walled hollow globule.” { One, two, three,
and even four of these may be present. Without fur-
ther present comment than that these characters, as

* Henle, Symbole ad Anatomiam vill.intest. Berol., 1887.

t Schleiden, Beitrage zur Phytogenesis, Miiller’s Archiv, 1838,
p. ii; Contributions to Phytogenesis, Sydenham Soc. Transl., p.
233. ;

{ The term nucleolus or nucleus-corpuscle (Kernkérperchen),
seems to have been first applied by Schwann. (See Introduction
to Schwann’s Researches, Syd. Society’s Translation.) .

34 THE CELL DOCTRINE.

given by Schleiden, are by no means constant, it is
plain that what is commonly known as the nucleolus
is here intended, to the discovery of which we are
therefore indebted to him, though Valentin also
claims its discovery at an earlier period.* He fur-
ther states that the observations he has made upon
all plants, lead him to the conclusion that these small
bodies are found earlier than the cytoblasts.
According to Schleiden, when starch, which is
superfluous nutritive material deposited for future
use, is to be employed in new formations, it becomes
dissolved into sugar or gum, which are convertible
into one another. The sugar appears as a perfectly
transparent fluid, not rendered turbid by alcohol,
and receiving from tineture of iodine only so much
color as corresponds to the strength of the solution.
The gum is somewhat yellowish, more consistent,
less transparent, and coagulated into granules by
tincture of iodine, assuming a pale yellow color,
which is permanent. In further progress of organi-
zation, in which the gum is always the last immedi-
ately preceding fluid, a quantity of exceedingly mi-
nute granules appears in it, most of which, from
their exceeding minuteness, appearing as black points.
It is in this mass that organization takes place,
though the youngest structures are composed of
another distinct, homogeneous, perfectly transparent
substance—so transparent as to be invisible when

* Valentin, ‘ Outline of the Development of Animal Tissues,”
in Wagner’s Elements of Physiology, translated by Drs Willis.
London, 1844, p. 214; Leipzig, 1889; where he refers to Valen-
tin’s Repertorium, vol. i, p. 143.

THE CELL DOCTRINE, 85

not surrounded by opaque or colored bodies,—and
continuing thus after pressure. This substance, which
frequently occurs in plants, Schleiden calls vegetable
gelatin, and considers as slight modifications, pectin,
the basis of gum tragacanth, and many of the sub-
stances usually enumerated under the term vegetable
mucus. It is this gelatin which is ultimately,
through the agency of the nucleus, converted into
the actual cell-wall, or structures which consist of it
in a thickened state, and into the matter of vege-
table fibre.

There are two situations in the plant in which
new organization may be observed most easily and
clearly, in consequence of there being cavities closed
by a simple membrane, Ist, in the large cell, which
subsequently contains the albumen of the seed, the
embryonal sac, and 2d, in the extremity of the pollen
tube, from which the embryo itself is developed. The
embryonal sac never contains starch originally, but
probably in most instances the saccharine solution or
gum. The pollen always contains starch, or repre-
senting it, a semi-granulous substance identical with
the small granules in the gum above alluded to, which
Schleiden calls mucus.

In both of these situations the above-mentioned
minute mucus-granules are very soon developed in
the gum, upon which the solution, previously homo-
geneous, becomes clouded and more or less opaque.

Single, larger, more sharply defined granules next
become apparent, A, Fig. 2, constituting the nucleoli,
and soon after the cytoblasts or nuclei, B, appear, look-
ing like granulous coagulations about the granules.

36 THE CELL DOCTRINE.

Thecytoblasts then grow considerably in the free state,
C, but so soon as they have attained their full size,
a delicate transparent vesicle rises upon their surface,
assuming the relation of the watch-crystal to a
watch, D, E. This is the young cell, which at first

Fie. 2,

Cellular Tissue, from the embryo sac of Chamzdorea
«
Schiedeana, in the act of formation.
A, Formative substance, gum, mucus-granules, nu-

clei of cytoblasts (nucleoli).

B, Cytoblasts.

C, Single and free cytoblast, more highly magni-
fied.

D, Cytoblast with cell forming in it.

E, Same, more highly magnified.

TF, Cytoblast isolated after destruction of cell.

From Schleiden’s “ Beitrige zur Phytogenesis.”

represents a very flat segment of a sphere, the plane
side of which is formed by the cytoblast, and the
convex side by the young cell, which is placed upon
it somewhat like a watch-glass of a watch. In 4
natural medium it is distinguished almost by this
circumstance alone, that the space between its con-
vexity and the cytoblast is perfectly clear and trans-
parent, and probably filled with a watery fluid, and
is bounded by the surrounding mucus-granules, which
have been aggregated at its first formation, and are
pressed back by its expansion, as shown in D, E. But
if these young cells be isolated, the mucus-granules
may be almost entirely removed by shaking the
stage. They cannot, however, be absent for any
length of time, for in a few minutes they become

THE CELL DOCTRINE. 37

completely dissolved in distilled water, leaving only
the cytoblasts behind. The vesicle gradually ex-
pands and becomes more consistent, and with the
exception of the cytoblast, which always forms a
portion of it, the wall now consists of gelatin. The
entire cell then increases beyond the margin of the
eytoblast, and quickly becomes so large that the
latter at least merely appears as a small body inclosed
in one of the side-walls in such a manner that the wall
of the cell splits into two lamine, one of which passes
exterior and the other interior to the cytoblast. That
upon the inner sideis generally the more delicate, and
in most instances only gelatinous, and is also absorbed
simultaneously with the cytoblast. Within these cells,
again, new cytoblasts arise, grow, and form young
cells, which grow and fill up the mother cells, and
finally cause the latter to disappear. This is endogen-
ous cell formation, while the formation of cells external
to other cells constitutes exogenous cell formation.
But according to Schleiden * the entire growth of the
plant consists only of a formation of cells within
cells.”* No other method of formation of new cells
seems to have been conceived by him. For although
the multiplication of cells, by fissiparous division of
previously existing cells, had been demonstrated by
Mirbel,f and confirmed by Von Mohl,} and the seg-

F

* Loe. citat., p. 257.

+ Mirbel, Recherches sur la Marchantia, 1833, Schleiden, how-
ever, says distinctly (op. cit. p. 232), “‘ Mirbel does not make any
allusion to the process of cell formation.”

f¢ Von Mohl, Entwicklung und Bau der Sporen der Kryptogam.
Gew., Flor., 1833.

4

38 THE CELL DOCTRINE.

mentation of the egg had been observed even earlier
(1824) by Prevost and Dumas, all before the inves-
tigations of Schleiden had been made, the latter
author considered the apparent growing across of the
partition walls an illusion, and that the young cells
escape observation in consequence of their transpar-
ency, until, at a late stage, their line of contact is re-
garded as the partition wall of the parent cell; while
even Schwann states somewhat hesitatingly what is
now so generally admitted.* This is the cell theory of
Schleiden, which he assumes to be the universal law
for the formation of vegetable cellular tissue in the
phanerogamia. At that time the cryptogamia had
not been examined, and Schleiden had not: then ex-
pressed his views in reference to the cambium.

The merit of Schwann consisted in applying this
theory to animal tissues, his conclusions being based
upon the study of the formation of the chorda dor-
salis and cartilage, and a comparison of their cells
with those of vegetable tissues. Thus, in a cyto-
blastema, either structureless or minutely granulous,
“a nucleolus is first formed; around this a stratum

‘ of substance is deposited, usually minutely granulous,
but not yet sharply detined on the outside. .As new
molecules are constantly being deposited in this stra-
tum between those already present, and as this takes
place within a precise distance of the nucleolus only,
the stratum becomes defined externally, and a cell
-nucleus, having a moreor lesssharp contour, is formed.
The nucleus grows by a continuous deposition of new

* Schwann, op. citat, Introduction, p. 4.

THE CELL DOCTRINE. 39

molecules between those already existing, that is by
intussusception. (See Fig. 3,¢.) If this go on equally
throughout the entire thickness of the stratum, the

Fie. 8.

From the point of a Branchial Cartilage of Rana esculenta.
(From Schwann.)
nucleus may remain solid ; but if it go on more vigor-
ously in the external part, the latter will become
more dense, and may become hardened into a mem-
brane, and such are the hollow nuclei.”*

When the nucleus has reached a certain stage of.
development, the cell is formed around it. The fol-
lowing is the process by which this takes place:
“ A stratum of substance, which differs from the cyto-
blastema, is deposited upon the exterior of the nu-
cleus. (See Fig. 3, d.) In the first instance, this
stratum is not sharply defined externally, but be-
comes so in consequence of the progressive deposition
of new molecules. The stratum is more or less thick,
sometimes honfogeneous, sometimes granulous: the
latter is most frequently the case in the thick strata
which occur in the formation of the majority of ani-

* Schwann, op. citat., p. 175.

40 THE CELL DOCTRINE.

mal cells. We cannot, at this period, distinguish a
cell cavity and cell wall. The deposition of new
molecules, between those already existing, proceeds,
however, and is so effected that when the stratum is
thin, the entire layer, and when it is thick, only the
external portion, becomes gradually consolidated into
amembrane. The external portion of the layer may
become consolidated soon after it is defined on the
outside; but, generally, the membrane does not be-
come perceptible until a later period, when it is
thicker and more defined internally; many cells,
however, do not exhibit any appearance of the for-
mation of a cell membrane, but they seem to be solid,
and all that can be remarked is that the external por-
tion of the layer is somewhat more compact.*
“Immediately that the cell membrane has become
consolidated, its expansion proceeds as the result of
the progressive reception of new molecules between
the existing ones; that is to say, by virtue of a growth
by intussusception, while at the same time it becomes
separated from the cell nucleus..... '. The inter-
space between the cell membrane and the cell nu-
cleus is at the same time filled with fluid, and this
constitutes the cell contents. During this expansion
the nucleus remains attached toa spot on the internal
surface of the cell membrane.” Though, according to
Schwann, in animal cells the nucleus is never covered
by a lamella passing over its inner surface, as is the
case with the vegetable cell according to Schleiden.

* Schwann, op. citat., p. 176. Stricker also informs us (Syden-
ham Soe. translation, p. 5) that the corpuscles of mucus and pus,
even in the eyes of Schwann, had no cell-wall.

THE CELL DOCTRINE. 41

Thus is formed the animal cell according to
Schwann, and although its method is very similar to
that of Schleiden, both as to endogenous and exoge-
nous cell formation (for Schwann did not restrict
cell genesis to endogenous cell formation), we have’
quoted his own paper because he is plainly fuller and
more precise in his descriptions. The object of each
observer was, however, the same with regard to the

\tissues studied; the additional object of Schwann
being to show that all organisms,'whether animal or
vegetable, ave formed on a common principle, and that
this principle is origin from cells,—that the various
tissues of the plant and animal, however simple or
complicated, are all combinations of these cells, modi-
fied in adaptation to the special peculiarities of tis-
sues.

The conception of Schleiden was truly original,
though its application was less difficult in conse-
quence of the simplicity of vegetable tissues. The
conception of Schwann was easier, in being the re-
flection of that of Schleiden, while its application
was more difficult, in consequence of the great diver-
sity of animal tissues; so difficult that he acknowl-
edged that “there are some exceptions, or at least
ditterences, which are as’yet unexplained.” This need
not surprise us when we recollect that one of theablest
modern exponents of the cell theory admits the diffi-
culty of its application to some of the so-called higher
tissues.* Indeed, the careful reader of Schwann’s

* Virchow, Cellular Pathology, Chance’s Translation. Am.
Edit., Philadelphia, 1863, p. 78.

4%

42 THE CELL DOCTRINE.

researches cannot but be surprised at the accuracy of
the observations of this histologist, nor can he fail to
realize how comparatively few have been the changes
necessitated in his descriptions, or the method of ap-
plication of his theory to the formation of the differ-
ent tissues; while the portion of the theory of
Schleiden and Schwann which does not accord with
the latest expression of the cell doctrine, is not so
much that which pertains to the formation of tissues
from existing cells as that which relates to the
method in which they supposed the cells to origi-
nate; which, it will be recollected, was by a species
of spontancous generation of the essential parts of the
cell, in a homogeneous cytoblastema.

A difference in the anatomy of the cell as given by
Schwann and physiologists of the present day, is
seen in the location of the nucleus by the former,
who places it not merely eccentrically, but actually
“separated from the surface only by the thickness of
the assumed cell-wall.”* At the present day, the
situation of the nucleus, though usually central, is
known to be not unvarying. Again, the primary
}and absolutely essential presence of the nucleolus, as
well as the universal presence of the cell-wall, may be
considered characteristics of Schleiden and Schwann’s
idea of the cell, which are now no longer insisted
upon.

As already stated (p. 38), Schwann would seem
to have admitted also, the formation of cells by di-
vision, though withsome hesitation. Thushe writes:t+

* Schwann, op. citut., p. 37, a. f.
t Schwann, op. citat., Introduction, p. 4.

THE CELL DOCTRINE. 43

“ A mode of formation of new cells, different from
the above described, is exhibited in the multiplica-
tion of cells by division of the existing ones; in this
case, partition walls grow across the old cell, if, as
Schleiden supposes, this be not an illusion, inasmuch
as the young cells might escape observation in con-
sequence of their transparency, and at a later stage,
their line of contact would be regarded as the parti-
tion wall of the parent cell.”
Schwann believed that the cell-wall was the most
active constituent of the cell, that it possessed the
power not only of producing physical and chemical
changes in its own substance and the cell contents,
but of secreting materials from the surrounding sub-
stance, and depositing them in its interior, explain-
ing in this manner the secretions of glands, the for-
mation of fat in some cells, pigment in others, ete.
It would be easy to point out other defects in the
theory of Schleiden and Schwann, when it is tested
by comparison with the more accurate observation
of the last twenty-five years, none of which should
be permitted to detract from the credit which at-
taches to the originators of this conception. It must
not be forgotten, that it is no less true of science than
of art, that great and important truths in their en-
tirety are gradually developed, and that no single
mind is capable of elaborating them from their in-
cipiency to their complete expression. And as many
clever people had daily noticed the rising of steam
from the boiling kettle without thinking of utilizing
its principles of expansion, so also, many careful ob-
servers had time and again witnessed the cellular or

44 THER CELL DOCTRINE.

vesicular composition of plants, and yet failed to ap-
preciate the importance of the-nucleated cell, and to
deduce from it a law of development applicable to
all organic forms. Again, asthe engine of Watt was
far different from the beautiful and powerful crea-
tion of the mechanic of the present day, so the cell
theory, as developed by Schleiden and Schwann, has
been further evolved by later histologists, We
may therefore truthfully reiterate, with Prof. Hux-
ley, that “whatever cavillers may say, it is certain
that histology before 1838, and histology since then,
are two different sciences—in scope, in purpose, and
in dignity—and the eminent men to whom we allude,
may safely answer all detraction by a proud ‘ circum-
spice,’ ”*

_ According to these observers, then, a perfectly
formed cell would be defined as a closed vesicle, with
certain contents, among which were essentially a nu-
cleolus and nucleus.

—

HENLE, BERGMANN, REICHERT, AND OTHERS, 1840-46.

It is not consistent with our object to include all
of the numerous observations which were multiplied
after this period, incited by the researches of Schlei-
den and Schwann. It is simply to point out the
salient features of those results which point towards
and have culminated in accepted views. It has been
stated that previous to Schleiden’s researches, in
' 1838, the formation of cells by division had been as-
serted as one mode of origin, that Schleiden had de-
clared this an error of observation, and that Schwann

* Huxley, op. citat., p. 290.

THE CELL DOCTRINE. 45

had hesitatingly, if at all, accepted it as a rare
method of cell formation.

Henle,* who, in general, adopted the view of
Schwann as to the primary origin of cells, though he
made exception to its universality of application,
says that cells multiply in three ways:

1, By budding (durch Sprossen), as in certain lower
plants.

2. By endogenous cell development (durch endo-
gene Zeugung), where the cell contents of the mother
cell become the cytoblastema of the daughter cells,
as originally given by Schleiden and Schwann.

3. By division or segmentation (durch Theilung),
of which he says, however, no examples are found
among animals; though he also states in the para-
grapht immediately following, “ We would, with
Schwann, consider cell formation in the yolk, by
‘furrowing,’ an analogous process, if we may con-
sider the yolk as a simple cell.” He then proceeds
to describe how, by a constriction of the surface, the
yolk is divided into two equal parts, these into four,
and so on until the entire yolk becomes a mulberry
mass, made up of little round bodies. This segmen-
tation of the ovum already observed in the yolks of
frogs, fish, molluses, and meduse, Henle says at
this time (1841), has perhaps merely escaped noticet
in the case of the higher animals, as plausibly sus-
pected by Bergmann,§ a suspicion which we need

* Henle, Allgemeine Anatomie. Leipzig, 1841, p. 172 e¢ seg.

+ Henle, op. cit., p. 176. { Henle, op. cit., p. 177.

2 Bergmann, Miiller’s Archiv, 1841. Bergmann also in this
paper objected to the existence of a cell membrane, and correctly

46 THE CELL DOCTRINE.

scarcely say was amply confirmed a little later. But
Henle also states, in the same connection, that certam
cases arise in which perfect cells are developed in a
cytoblastema, in a manner which is inexplicable, and
that from these cells tissues are finally developed.*
Whence the undetermined state of the question at
that time may be easily inferred. Nor is mention
“ here made by Henle of the nucleus of the cell as the
primary seat of thesegmentation. The surface of the
cell is said to be “ constricted ” or “ furrowed,” deeper
and deeper, until the division takes place. This de-
scription is still adhered to by many physiologists of
the present day, who consider that there is a simple
disappearance of the germinal vesicle or nucleus of the
ovum after fecundation, rather than a division of it
into two, and substitution of these for the original one.
While endeavoring to trace out the steps by which the
present most generally accepted views with regard to
the origin of cells were arrived at, it must not be for-
gotten that other dissenting views were also ad-
vanced, though tending differently from those incor-
porated in the text, where it is desired more particu-
larly to trace those culminating in existing doctrines.
Thus did Reichertt+ early (1840), dissent from Schwann,
since he failed to find the nucleus universally present
in the yolk, and he was the first to defend the view

maintained that the spheres of segmentation are cells which are at
first destitute of a cell membrane, though they become invested by
one at a subsequent period.

* Henle, op. cit., p. 177.

+ Reichert, Das Entwickelungsleben im Wirbelthierreich. Ber-
lin, 1840, pp. 6, 98.

THE CELL DOCTRINE. 47

that the segments into which the egg breaks up are
cells. Karsten* (1843), published a dissertation upon
the cell, in which he stated that cells originate with-
out a pre-existing nucleus, and by the expansion of
amorphous granules of organic matter; and more
recently (1863), the same author practically reiterates
this view, since he says that all “ cells of vegetables
originate as minute free vesicles in the ‘fluid contents
of previously existing cells,” and regards the nucleus
as a “small tertiary cell, retarded in its develop-
ment.”+ Again, “when the nucleus is present, the
origin of new cells is quite independent of it.”{ Jn
addition to the statement already given, Henle also
(1843), alleged that some of the so-called fibrous tis-
sues were “formed by the aggregation of granules
in a certain way without the intervention of true
nucleated cells.Ӥ Kdlliker,|| one of the foremost
exponents of the cell doctrine of the present day, in
1844 expressed his dissent from the idea of unity in
the mode of cell formation, and states that if there
is a single method of cell formation which is in-
variable, it remains to be discovered, although he
interpreted the segmentation of the germ of ce-
phalopods in the same manner as Bergniann. Mr.

* Karsten, De Cella vitale Dissertatio. Berlin, 1848.

+ Karsten, Ann. and Mag. of Nat. History, vol. xiii, p. 268.
London, 1864.

{ Karsten, Ann. and Mag. Nat. History, vol. xiii, p. 281.

% Henle, Traité d’Anatomie Générale. ‘Trad. d’Allemand,
par A. J., Jourdan, 2 vol., Paris, 1843, tom. 1, p. 374.

|| Kélliker, Entwickelungsgeschichte der Cephalapoden. Zurich,
1844.

48 THE CELL DOCTRINE.

Paget,* so well known from his Lectures on Surgical
Pathology, suggested in 1846, that a cell might arise
in some other way than from a nucleus, since he had
met morbid growths composed entirely of fibres, in
which not a nucleated cell was present. Most of these
statements are, however, reconciled by the informa-
tion which has since been added to our knowledge of
the subject. -

MARTIN BARRY, 1840.

It was in his first series of embryological researches,
published in Part II of the “ Philosophical Transac-
tions” of the Royal Society of London, for 1838, p.
310, that Dr. Martin Barry declared “that the ger-
minal vesicle (which he regarded as the nucleus), and
ita contents constitute throughout the animal king-
dom the most primitive portion of the ovum.” In his
second series, Part II, 1839, in stating that the ger-
minal vesicle returns to the centre of the cell, post
coitum, he first pointed out that'the nucleus does not
always accompany the cell through the whole vital
process at the periphery (the original position accord-
ing to Schleiden and Schwann), but that it also passes
to the centre, as we now well know. Here, also, he
declares, but in his third series, Part IJ, 1840, he
demonstrates, that there arise in the parent vesicle, two
or more infant vesicles, the parent vesicle disappear-
ing by liquefaction. And in his third series, p. 529,
he says, “ The germinal vesicle does not burst, or dis-
solve away, or become flattened, on or before the fecun-

* Paget, Report on the Progress of Anatomy and Physiology,
Br. and For. Med. Rev., July, 1846.

THE CELL DOCTRINE. 49

dation of the ovum as hitherto supposed. It ceases
to be pellucid.” And on page 531, “The germinal
vesicle fills with cells, and these become filled with
the foundations of other cells; so that the germinal
vesicle is gradually rendered opaque.”

He also describes in this series, in great detail, the
mode in which these cells are produced from’ the
germinal spot, which he considers in the light of a
nucleus to the germinal vesicle. Part II, 1839, p.
360. And though the minute details may not pre-
cisely accord with those of the most recent observa-
tions, the correct idea is clearly grasped. In fact, it
may be said that in minuteness of detail alone does
he differ from later observers, and had he simply
stated that the young cells arise from the nucleus or
nucleolus of the parent cell, he would accord pre-
cisely with the most recent observers. But he is, if
possible, even more explicit when he says, “ The pro-
cess inherited from the germinal vesicle by its off
spring, reappears in the descendants of these. Every
cell, whatever its minuteness, if its interior be dis-
cerned, is filled with the foundations of new. cells,
into which its nucleus has been resolved.” Again
he says,* “ Schleiden has seen the nucleus undergoing
such changes (division), but failed to recognize them.”
And finally, in “ Philosophical Transactions ” for 1841,
pp. 207-8, we have the following striking paragraphs,
which would seem also to correct some previous
errors:

Ҥ 77. I am very much inclined to believe, that

* Barry, Philosophical Transac., 1840, p. 348, 3 385.
5 .

50 THE CELL DOCTRINE.

in the many instances in which authors on ‘cells’
have -described and figured more than one nucleolus
in a nucleus, there has been either an incipient divi-
sion of the nucleus into discs, or the nucleus has
consisted of two or more discs ; the nucleoli of those
authors having been the minute and highly refract-
ing cavities or depressions in the discs. If this has
really been the case, it affords additional evidence, I
think, that the reproduction of cells by the process I have
described—namely, division of the nucleus of the parent
cell—is universal—so numerous have been the in-
stances in question. I may refer to the figures given
by Schwann, who examined nearly every tissue, and
to those of Schleiden, whose observations have been
so extensive on plants. I think, indeed, that many
‘of the figures of Schwann afford evidence of the
division in question having taken place. It is to be
recognized in his delineation of the cells of cartilage,
cellular tissue, middle coat of the aorta, muscle, ten-
don, feather, etc. The same remark is applicable to
a figure given by Reichert of ciliated epithelium
cells. Dr. Henle found that in the layers of his
‘ pflaster-epithelium ’ cells, the nucleus, very distinct
in the lower cells, had almost disappeared in those
situated in the upper part. From this observation,
and from the presence of two nucleoli in some of the
nuclei figured by this observer, as well as from the
nucleus becoming more granular, I think it extreme] y
probable that these cells (including those of the epi-
— are reproduced by the process just referred
0,—division of the nucleus ; additions being no doubt
seieecads made at the lower part of the layer, by
‘which cells previously there are pushed farther out.’

THE CELL DOCTRINE. 51

“883. The nuclei which various observers have
found lying among the fibres of various tissues, have
been considered by them as the ‘ remains of cells.’
This may have been the case, but so far from think-
ing with those observers, that the nuclei in question
were ‘ destined to be absorbed,’ I am disposed to con-
sider that they were sources from which there would
have arisen new cells.”

Without doubt, we can say, as did Goodsir,* in the
above by Martin Barry, we have the “ first consistent
account of the development of cells from a parent
centre, and more especially of the appearance of
centres within the original sphere.” Nothing more
definite, or directly tovwthe point, could be desired,
and we think it may be justly said of Barry, that
he completed the expression of the cell theory in-
augurated by Schleiden and Schwann, in modifying
the mode of origin to conform to most recent obser-
vation.

PROF. JOHN GoopsiIR, 18-45.

In 1845, Prof. John Goodsir published his paper on
“Centres of Nutrition,’t in “ Anatomical and Patho-
logical Observations,” in which he clearly grasped the
two important principles of the modern Cellular Pa-
thology ; first, the activity of these centres (nuclei), their
power “to draw from the capillary vessels, or from
other sources, the materials of nutrition, and to distrib-
ute them by development to each organ or texture after
its kind ;” second, the origin of such centres or nuclei

* Goodsir, Turner’s Edition of Anatomical Memoirs. Edin-
burgh, 1868. Note on p. 390.
+ Goodsir, op. citat., p. 389.

52 THE CELL DOCTRINE.

from previously existing nuclei. In this short paper of
three pages, are contained, as stated, the essentials of
the cell doctrine of Virchow, and as it has recently as-
sumed additional interest on controversial* grounds,
it may be well to introduce as much as bears directly
upon the subject. “The centre of nutrition with
which we are most familiar, is that from which the
‘whole organism derives its origin,—the germinal
spot of the ovum. From this, all the other centres
are derived, either mediately or immediately; and
in directions, numbers, and arrangements, which
induce the configuration and structure of the being.
As the entire organism is formed at first, not by si-
multaneous formation of its parts, but by the suc-
cessive development of these from one centre, so the
various parts arise each from its own centre, this
being the original source of all the centres with
which the part is ultimately supplied.

“ From this it follows, not only that the entire or-
ganism, as has been stated by the authors of the
cellular theory, consists of simple or developed cells,
each having a peculiar independant vitality, but
that there is in addition, a division of the whole into
departments, each containing a certain number of de-
veloped cells, all of which hold certain relations to one
central or capital cell, around which they are grouped.
It would appear that from this central cell, all the
other cells of its department derive their origin. It
is the mother of all those within its own territory.

* Edinburgh Monthly Medical Journal, February and April,
1869, pp. 766 and 959.

THE CELL DOCTRINE. 58

It has absorbed materials of nourishment for them
while in a state of development, and has passed
them off after they have been fully formed, or have
arrived at a stage of growth when they can be de-
veloped by their own powers.

“Centres of nutrition are of two kinds,—those
which are peculiar to the textures, and those which
belong to the organs. The nutritive centres of the
textures are in general permanent. Those of the
organs are in most instances peculiar to their em-
bryonic stage, and either disappear ultimately or
break up into the various centres of the textures of
which the organs are composed.

“ A nutritive centre, anatomically considered, is merely
a cell, the nucleus of which is the permanent source of
successive broods of young cells, which from time to time
fill the cavity of their parent, pass off in certain
directions and under various forins, according to the
texture or organ of which their parent forms a part.”

Prof. Goodsir does not fail to state in the first para-
graph of his paper, that with many of these centres
anatomists have been for some time familiar, but
further remarks, that with few exceptions they have
looked upon them as embryonic structures. He al-
ludes in a note to the observations of Bowman and
Barry, the former on “ Muscle,” and the latter “On
the Corpuscles of the Blood,” in Philosophical Trans-
actions, respectively, of 1840 and 1841, and states in
a second note that “for the first consistent account
of the development of cells from a parent centre, and
more especially the appearance of new centres within
the original sphere, we are indebted to .Martin

5*

54 THE CELL DOCTRINE.

Barry.”* We have carefully read the references in
each instance. In Bowman’s papert we can recog-
nize a brief reference to a possible influence of the
cell upon nutrition, but none as to its origin, in the
following sentence: “It is, however, not impossible,
that in all these cases, there may be during develop-
ment, and subsequently, a further and successive de-
posit of corpuscles (nuclei) from which both growth
and nutrition may take their source.” That Dr.
Barry’s paper is more explicit has been shown.

REMAK, 1853-55.

Remakt defended most effectually the view that
cells originate from previously existing cells by divi-
sion, and that at least in the early stages of the de-
velopment of the embryo, no other mode of cell devel-
opment occurs than by division. Remak also con-
tended for, and according to Stricker,§ established
the same law in respect to the pathological develop-
ment of cells, although Stricker admits also that
Virchow played an iniportant part in the extension
of our knowledge in this direction.

HUXLEY, 1853.||
Allusion has already been made to Prof. Huxley

* Goodsir, Anatomical. Memoirs, vol. ii, p. 889, and note on
pp. 890-91.

+ Bowman, ‘ Muscle,” Philos. Transac., 1840, pt. i, p. 485.

{ Remak, Untersuchung iiber die Entwickelung der Wirtel-
thiere. Berlin, 1852-55.

@ Stricker, Manual of Human and Corporative Histology. New
Syd. Soc. Transl., 1870, p. 84.

|| We resume it will scarcely be inferred by any reader, that

THE CELL DOCTRINE. 55

in connection with Wolff, of whose theory he has
been the able exponent. In the same paper* he has
given us his own views—“‘ conceived in the spirit,
and not unfrequently borrowing the phraseology,
of Wolff and Von Baer.” We present them, as
far as may be consistent with brevity, in his own
words:

“Vitality, the faculty, that is, of exhibiting defi-
nite cycles of change in form and composition, is a
property inherent in certain kinds of matter. There
is a condition of all kinds of living matter in which
it is an amorphous germ—that is, in which its exter-
nal form depends merely on ordinary physical laws,
and in which it possesses no internal structure. Now,
according to the nature of certain previous condi-
tions, the character of the changes undergone, or the
different states exhibited—or, in other words, the
successive differentiations of the amorphous mass
will be different.

“The morphological differentiation may be of two

the views of Prof. Huxley here presented are brought forward as
those now entertained by him, and with which the public have
been made so generally familiar through his lecture on ‘ Proto-
plasm,” or the ‘' Physical Basis of Life,” delivered at Edinburgh,
November 18th, 1868, and originally published in the “ Fort-
nightly Review” for February, 1869; but also largely republished
in numerous English and American periodicals, as well as in a
separate pamphlet, to be had of the publishers of the Yale College
Courant, New Haven, Conn. To one closely observing, however,
we think that these latter views will appear to be foreshadowed
in the theory here given, and which we think of sufficient his-
torical importance to justify its presentation here.

* Huxley, Review of the Cell Theory. Br. and For. Med.
Chir. Rev., October, 1853, p. 305.

56° THE CELL DOCTRINE.

kinds. In the lowest animals and plants,—the so-
called unicellular organisms—it may be said to be ex-
ternal, the changes of form being essentially confined
to the outward shape of the germ, and being unac-
companied by the. development of any internal struc-
ture.

“But in all other animals and plants, an internal
morphological differentiation precedes or accompa-
nies the external, and the homogeneous germ becomes
separated into a certain central portion, which we
have called the endoplast, and a peripheral portion,
the periplast. Inasmuch as the separate existence of
the former necessarily implies’ a cavity in which it
lies, the germ in this state constitutes a vesicle with a
central particle, or a ‘nucleated cell.’ There is no
evidence whatever that the molecular forces of the liv-
ing matter (the ‘ vis essentialis’ of Wolff, or the vital
forces of the moderns), are by this act of differentia-
tion localized in the endoplast to the exclusion of
the periplast, or vice versa. Neither is there any evi-
dence that any attraction or other influence is exercised
by the one over the other ; the changes which each sub-
sequently undergoos, though they are in harmony,
having no causal connection with one another, but each
proceeding, as it would seem in accordance with the
general determining laws of the organism. On the
other hand, the ‘vis essentialis’ appears to have es-
sentially different and independent ends in view, in
thus separating the endoplast from the periplast.

“The endoplast (nucleus) grows and divides; but,
except in a few more or less doubtful cases, it would.
seem to undergo no other morphological change. It

THE CELL DOCTRINE. 57

frequently disappears altogether ; but as a rule it
undergoes neither chemical nor morphological meta-
morphosis. So far from being the centre of activity
of the vital actions, it would appear much rather to
be the less important histological element.

“The periplast, on the other hand, which has
hitherto passed under the name of cell-wall, contents
and intercellular substance, is the subject of all the
most important metamorphic processes, whether mor-
phological or chemical, in the animal and plant. By
its differentiation, every variety of tissue is pro-
duced ; and this differentiation is the result, not of
any metabolic action of the endoplast, which has fre-
quently disappeared before the metamorphosis begins,
but the intimate molecular changes in its substance,
which take place under the guidance of the ‘vis
essentialis,’ or, to use a strictly positive phrase, occur
in a definite order, we know not why.

“ The metamorphoses of the periplastic substance
are twofold,—chemical and structural. The former
(chemical), may be of the nature either of conversion,
—change of cellulose into xylogen, intercellular sub-
stance, etc., of the indifferent tissues of embryos,
into collagen, chondrin, etc.,—or of deposit,—as of
silica in plants, of caleareous salts in animals. The
structural metamorphoses, again, are of two kinds,
vacuolation or the formation of cavities, as in the
intercellular passages of plants, the first vascular
canals of animals; and fibrillation, or the develop-
ment of a tendency to break up in certain definite
lines rather than in others.”

These views he illustrates by examples from vege-

58 THE CELL DOCTRINE.

table life in the sphagnum leaf, and from animal life
in connective tissue and striped muscle.

As characteristic and distinguishing features of
this theory, we desire to point out, first, the substi-
tution of the term “ endoplast” for “ nucleus ;” that
of “ periplast” for “ cell-wall,” and “ intercellular,”
“substance.” Second, the absolutely passive nature
of the “ endoplast,” which is neither itself the author
of changes nor the subject of changes. Third, the
passive nature.as well of the “ periplast,” so far as
it is the author of changes, though it is pre-emi-
nently the subject of changes, the seat in which changes
take place. And herein, we believe Huxley to have
been misinterpreted by some who have presented
his views elsewhere, as Dr. Beale,* who repre-
sents him as believing the periplast active, that it is
the efficient agent, that it sends in partitions, ete.
But that Prof. Huxley considered it passive we be-
lieve may be legitimately inferred from his text.
As the seat of change, however, accomplished not as
“ the result of any metabolic action of the endoplast,
but of intimate molecular changes in its substance,
which take place under the guidance of the vis essen-
tialis,” the periplast is differentiated into every variety
of tissue. Finaily, we have the distinct admission,
as seen in the sentence last quoted, and also through-
out the entire expression of the theory, of a con-
trolling, guiding principle, through which the difter-

* Beale, Microscope in Medicine. Third Edition. London,
1867, page 147. Beale, Structure and Growth of the Tissues,
London, 1865, pp. 9, 10.

THE CELL DOCTRINE. 59

entiationisaccomplished. This principle, which is here
referred to as the “ vis essentialis,” is elsewhere in-
cluded under the expressions “ vitality,” and “ general
determining laws of the organism.” Though this ad-
mission is seemingly so at variance with the views of
the same observer in 1870, when, in common with
other physicists, he emphatically denied the exist-
ence of “vital force,” or even such a thing as life
itself, yet, as already intimated, we deem it possible
to detect a foreshadowing of his more modern views,
in the following paragraph of the paper whence we
have derived our information:

“We have therefore maintained the broad doctrine
established by Wolff, that the vital phenomena are
not necessarily preceded by organization, nor are in
any way the result or effect of formed parts, but
that the faculty of manifesting them resides in the
matter of which living bodies are composed, as such ;
or, to use the language of the day, that the vital
forces are molecular forces.”*

Huxley moreover says that the three botanical
data upon which Schwann’s theory was based, viz. :

1. The anatomical independence of the vegetable
cell as a separate entity.

2. His conception of the structure of the vege-
table cell, and

3. Its mode of development, were all erroneous.

Since first, he (Huxley). considers that the fact
that by certain chemical or mechanical means, a
plant may be broken up into vesicles, corresponding

* Huxley, loc. citat., p. 814.

60 THE CELL DOCTRINE.

with the cavities which previously existed in it, is of
no more value in proving the independence of these
vesicles, than the fact that a rhombohedron of spar,
broken up with the hammer, into minute rhombohe-
drons, is evidence that those minuter ones were once
independent, and formed the larger by their coales-
cence.

Second, Schwann’s view of the anatomy of the cell
was incorrect, since he regarded the nucleus as in-
variably present, whereas in certain vegetable cells
(as in Hydrodictyon, Vaucheria, Caulerpa, Sphag-
num), it is indubitably absent ; and since he did not
include the nitrogenous primordial utricle, discov-
ered by Mohl, in 1844,* as one of the elements of
the cell.

Finally, Schwann’s mode of cell-development is
erronous, having “been long since set aside by the
common consent of all observers ;” cell-development
always occurring by division, except in the embryo
sac of the Phanerogamia, the sporangia of Lichens,
and of some Alge and Fungi; and even the free
cell-development of the latter is quite different from
that of Schleiden and Schwann, being by the develop-
ment of a cellulose membrane (periplast) around a
mass of nitrogenous substance (endoplast), which
may or may not contain a nucleus.

The difference between the views of Schwann and
Huxley are best expressed by the latter in the con-
trast he draws between those of Schwann and Wolff:

* The existence of the primordial utricle is denied by many
botanists of the present day.

THE CELL DOCTRINE. 61

“For Schwann, the organism is a beehive, its action
and forces resulting from the séparate but harmo-
nious action of all its parts. For Wolff (and Hux-
ley), it is a mosaic, every portion of which expresses
only the conditions under which the formative power
acted, and the tendencies by which it was guided.”
The statements of Prof. Huxley with regard -to
cell-development entirely accord with the most recent
observations on the subject, and are quite important to’
us in tracing out the present state of the cell doctrine.

J. HUGHES BENNETT, 1855.*

Dr. Bennett, of Edinburgh, considered that “the
ultimate parts of organization are not cells nor nu-
clei, but the minute molecules from which these are
formed They possess independent physical and vital
properties, which enable them to unite and arrange
themselves so as to produce higher forms. Among
these are nuclei, cells, fibres, and membranes, all of
which may be produced directly from molecules.
The development and growth of organic tissues is
owing to the successive formation of histogenetic
and hystolytic molecules. The breaking down of
one substance is often the necessary step to the for-
mation of another; so that the histolytic or disin-

* Bennett’s Practice of Medicine. Am. Ed. of William Wood
& Co., N. Y., 1866, p. 118. i
Prof. Bennett has further elaborated his views in the Edinburgh
Medical Journal, March, 1868, and The Popular Science Review,
January, 1869, but his conclusions are substantially the same as
quoted.
6

62 THE CELL DOCTRINE.

tegrative molecules of one period become the histo-
genetic or formative molecules of another.”

Again: “As to development, the molecular is the
basis of all the tissues. The first step in the process
of organic formation is the production of an organic
fluid; the second, the precipitation in it of organic
molecules, from which, according to the molecular
law of growth, all other textures are derived either
directly or indirectly.”’*

Figs. 4, 5, 6, 7, illustrate these views amply.

Fie. 4. Fie. 5. Fre. 6. Fic. 7.

< & Legs SPS,

ae,

Fig. 4, Molecular structure of the scum on its first appearance, in a clear ani-
mal infusion. Fig. 5, Molecular structure of the same six hours afterwards.
The molecules are separated, and the long ones (so-called vibriones) in active
movement. Fig. 6, The same on the second day. Fig. 7, Filaments (so-called
spirilla) formed by aggregation of the molccules, in the same scum on the third
and fourth days, allin rapid motion. 800 diam. linear. (From Bennett's Prac-
tice.) :

Prof. Bennett contends, also, that morbid growths
may easily be shown to originate in a molecular
blastema, though not to the exclusion of pre-existing
cells. The accompanying figures are sufficiently ex-
planatory.

It should be stated also that this author, in com-
mon with others not accepting the cell doctrine in
its entirety, admits the production of cells by buds,
division or proliferation, without a new act of gen-
eration, and that “ this fact comprehends most of the
admitted observations having reference to the cell
doctrine.” +

* Op. citat., p. 119. + Op. citat., p. 128.

THE CELL DOCTRINE. 63

We have in the expression of this theory, a prac-
tical admission of the spontaneous origin,of animal
life, of which Dr. Bennett, in the paper referred to
in the Popular Science Review, for January, 1869,

Fic. 9. Fie. 10. Fie. 11.

Fig. 8, Nuclei imbedded in a molecular blastema. Fig. 9, Young fibre-cells
formed by the aggregation of molecules around the nuclei of Fig. 8. Fig. 10,
Cancer cells, one with a double nucleus. Fig. 11, Histolytic or so-called granule-
cells, breaking down from fatty degeneration. 250 diam. linear. (From Ben-
nett’s Practice.)
openly declares himself the advocate, while the views
are in no way essentially different from those of
Schwann.

Closely allied to this theory is the so-called in-
vestment or cluster theory (Umhiillungs-theorie), de-
scribed by Virchow on page 53 of Cellular Pathology
(Am. Ed. of Chance’s Translation): according to
which “originally a number of elementary globules

Fie. 12,
a@
- fo)
08S (} 9927,
0°95 2,2 Sx
Lory lon.
Po, Sas

Diagram of the Investment (cluster) theory. a, Separate elementary granules,
b, Heap of granules (cluster). e¢, Granule-cell, with membrane and nucleus.

existed scattered throughout a fluid, but that under
certain circumstances they gathered together, not in
the form of vesicular membranes, but sv as to consti-
tute a compact heap, a globe (mass, cluster—Kliimp-
chen), and that this globe was the starting-point of

64 THE CELL DOCTRINE:

all further development, a membrane being formed
outside and a nucleus inside, by the differentiation
of the mass, by apposition, or intussusception.”

TODD AND BOWMAN, 1856.

Notwithstanding earlier approximations to the
truth, we find free cell formation stil] admitted by the
eminent authorities, Todd and Bowman, as one mode
of origin of cells, so late as December, 1856, though
the spontaneous origin of organs is spoken of as ex-
ceedingly doubtful. After describing the elements
of the ovum, considered in its entirety as a nucleated
cell, and referring to the period after fecundation, it
is stated, “ At this period the embryo consists of an
aggregate of cells, and its further growth takes place
by the development of new ones. This may be ac-
complished in two ways: first, by the development
of new cells within the old, through the subdivision
of the nucleus into two or more segments, and the
formation of a cell around each, which then becomes
the nucleus of a new cell, and may in its turn be the
parent of other nuclei; and, secondly, by the forma-
tion of a granular deposit between the cells, in which
the development of the new cells takes place. The
granules cohere to each other in separate groups, here and
there, to form nuclei, and around each of these a delicate
membrane is formed, which is the cell membrane. The
nuclei have been named cytoblasts, because they appear
to form the cells; and the granular deposit in which
these changes take place is called the cytoblastema.

“In every part of the embryo the formation of
nuclei and of cells goes on in one or both of the ways

THE CELL DOCTRINE. 65

above mentioned, and, by and by, ulterior changes
take place, for the production of the elementary
parts of the tissues.”*

Thus did physiologists adhere to the original free
cell formation of Schleiden and Schwann. Singu-
larly, Dr. Carpenter,t who expressly states, in his
Manual of Physiology, edition of 1865, that he has
been led to the view of Professor Beale by. compari-
son of the results of the recent inquiries of several
British and Continental histologists with those of
his own studies, says, a few pages further on (p. 150),
“‘ New cells may originate in one of two principal
modes; either directly from a previously existing
cell, or by an entirely new process in the midst of an
organizable blastema.” He then proceeds to give
the two methods in detail, without in any way deny-
ing the latter.

virncHow, 1858.

Less than two years later, August 20th, 1858,
Prof. Virchow published his “ Cellular Pathology,
as based upon Physiological and Pathological His-
tology.” According to him, the cell is the only ,
possible starting-point for all biological doctrines.
This cell can only originate from a previously exist-
ing cell, taking its primary origin from the ovum,
and the Harveian maxim omne vivum ex ovo, becomes
in its special application, omnis cellula e cellulé. This

* Todd and Bowman, The Physiological Anatomy and Physi-
ology of Man. Am. Edit., Philadelphia, 1857, p. 63.
+ Carpenter, Manual of Physiology. London, 1865. Note on
p. 14,
6*

66 THE CELL DOCTRINE,

is true of all physiological and pathological processes
in the vegetable and animal. In all editions of
“QOellular Pathology” which we have met, the
‘typical cell is described as consisting essentially of
“ cell-wall,” “cell contents,” and “nucleus;” the
“nucleolus,” though usually met in fully developed
older forms, is not considered an essential constit-
uent of the cell. The object of the “nucleus,” ac-
cording to Virchow, is entirely connected with the
life of the cell, that which maintains it as an element
and which ‘insures its reproduction. While to the
“cell contents ” over and above the nucleus, that is
the “residual cell contents,” is due the function of
the cell, that to which is due the contractility of
muscle, the neurility and sensation of nerve, and the
secretory office of the gland cell.*

To secure the universal application of the cell
doctrine, it becomes necessary to eliminate from the
‘vegetable cell, the external non-nitrogenous mem-
brane. known as cellulose, and restrict it to the nitro-
genized portion comprised in the primordial utricle
as the proper cell-wall, and in the protoplasmic con-
tents of the cavity as the proper cell contents, which
contain also the nucleus. “It is only when we ad-
here to this view of the matter, when we separate
from the cell all that has been added to it as an after-
development, that we obtain a simple, homogeneous,

* Virchow, Cellular Pathology, as based upon Physiological
and Pathological Histology. Setond Edition. Translated by
Frank Chance, M.B., etc. Am. Edition, Philadelphia, 1868, p.
37.

THE CELL DOCTRINE, 67

extremely monotonous structure, recurring with ex-
traordinary frequency in living organisms.”*

More recently, however, Virchow is reported as
not regarding the “ cell-wall” as an essential part of
the cell, as stated in Cellular. Pathology ; but that a
nucleus surrounded by a molecular blastema was suffi-
cient to constitute a cell ; then he says that the outer
part of this cell blastema consolidates and forms a
cell-wall as Beale has shown, and that this takes
place in the ameeba when placed in water.t

As thus detined, the cell is the seat of pathologi-
cal and physiological processes rather than the blood,
or the nerves. The cell is active—the ultimate mor-
phological element in which there is any manifesta-
tion of life, and beyond which the seat of real ac-
tion cannot be removed. Hence the term Cellular
Pathology rather than humoral, or neural, or solid-
istic. The so-called erudations are not such in the
strict sense of the term, and the cells which they
contain, whether of pus or organizable lymph, are
the result of proliferation of previously existing cells.
Even “ fibrin, wherever it occurs in the body exter-
nal to the blood, is not to be regarded as an excre-
tion from the blood, but as a local production,” re-
sulting from the activity of the cells of the tissue
in which it is found, and conveyed to the surface by
the transudation of the serous fluids alone.t In the

* Op. cit., pp. 31, 84.

+ Letter from Berlin, in Edinburgh Medical Journal, February,
1865.

} Virchow, op. cit., pp. 485-6,

sy

68 THE CELL DOCTRINE.

above statements we have the first distinctive feature
of Virchow’s theory.

Again, since every organized body is usually made
up of a number of these cells, each independent in
itself, yet combined and arranged for the attainment
of a special end, and therefore mutually dependent,
there result certain communities or cell territories
into which the body is portioned out by Virchow.
But not only is the relation of these cells to each
and to the central cell whence they took their origin
mutually dependent, but in many animal tissues, at
least, we have the so-called intercellular substance, in
a certain definite manner dependent upon the cell
or cells which it surrounds, “so that certain districts
belong to one cell and certain others to another.”
Especially is this the case in pathological processes,
where sharp boundaries may often be drawn between

_cell territories. Herein have we the second dis-

tinguishing character of Virchow’s theory.

_ There are also a third and fourth distinctive fea-
tures. It has already been explained that the prin-
ciple of the theory of Schleiden and Schwann lay
in this, that every tissue, healthy or morbid, results
from the apposition of cells, and that this principle
is still observed as correct, the mode of origin of the
primary cell being alone the object of dispute. Ac-
cording to Virchow, however, it is a special cell
which becomes the starting-point of physiological
and pathological processes, and by its various meta-
morphoses constitutes the healthy or morbid tissue,
excepting epithelial formations. This cell is the so-
called connective tissue corpuscle, or cell of the con-

THE CELL DOCTRINE. 69

nective tissue, which, according to Virchow, is a cell
with all its essential constituents (cell-wall, cel] con-
tents, and nucleus), and not a nucleus alone, as origi-
nally described by Schwann, and later by Henle* and
Landois.t| From the well-known universal preva-
lence of connective tissue, this view receives support.
Thus, it is from the connective tissue corpuscles of
the soft, silk-like connective tissue, so universally
present in muscle, that the muscular fasciculi are
primarily developed. It is from these that nerve-

Fie. 18. Fie. 14.

Fig. 18. Purulent granulation from the subeutaneous tissue of a rabbit, round
about a ligature. a, Connective tissue corpuscles. 6, Enlargement of the cor-
puscles with division of the nuclei. c¢, Division of the cells (granulations). d,

Develop t of the p' ‘puscl 300. (From Virchow.)
Fie, 14. Interstitial purulent inflammation of muscle in a puerperal woman.
mm, Primitive muscular fibres. 77, Develop t of pi puseles by means

of the proliferation of the corpuscles of the interstitial connective tissue. ><280.
(From Virchow.) ,

fibres take their origin. It is by the rapid prolifera-
tion of these corpuscles that pus is formed (Figs.
13 and 14); it is from the perverted growth and de-

* Henle, Bericht tiber die Fortschritte d. Physiol., 1859; 1866,
p. 41.
¢ Landois, Zeits. f. wiss. Zool., Bd. xvi, p. 1.

70 THE CELL DOCTRINE.

velopment of these that tubercle and cancer arise
(Fig. 15), and similarly all pathological new forma-
tions. None of these products are exudations from

Development of cancer from connective tissue in carcinoma of the breast. a,
Connective tissue corpuscles. 6, Division of the nuclei. ¢, Division of the cells.
d, Accumulation of the cells in rows. e, Enlargement of the young cells and
formation of the groups of cells (foci, Zellenheerde), which fill the alveoli of
cancer. J, Further enlargement of cells and groups. g, The same development
seen in transverse section. (From Virchow.)

the blood, according to Virchow. They are entirely
local in their origin. In these views he is supported
by the majority of German observers.

Another mode of formation of pus is however ad-
mitted by Virchow, in the growth and development
of new cells in epithelium, whether in cuticle or mu-
cous membranes. Whether forms of suppuration
exist which may be referred to muscular, nervous,
and capillary elements, he considers doubtful.
~ A fourth and final distinctive feature of Virchow’s
views, concerning which there is less unanimity, even
among German histologists, is his peculiar system of
canals or tubes, produced by the anastomosis of one

f THE CELL DOCTRINE. 7L

cell with another, and which he considers must be
classed with the great canalicular system of the body,
as forming a supplement to the blood and lymphatic
vessels, and as filling up the vacancy left by the old
vasa serosa, which do not exist.* (See Fig. 16.) Of

Fia. 16.

Connective tissue from the embryo of a pig after long-continued ‘boiling.
Large spindle-shaped cells, connective tissue corpuscles (Bindegewebeskérper-
chen), some isolated and some still imbedded in their basis substance, and anas-
tomosing one with the other. Large nuclei with their membrane detached;
cell contents in some cases shrunken. 350. (From Virchow.)

this system he also considers the cordlike fibres of
yellow elastic tissue as forming a part.t These he
considers, with Donders,t as originating by a trans-

* Virchow, op. citat., p. 76.
+ Virchow, op. citat., p. 133, a. f.
t Donders, Siebold und K@lliker’s Zeitschrift, Bd. iif.

72 THE CELL DOCTRINE.

formation of the connective tissue corpuscles them-
selves. He says, “ The transformation of these latter
into the former, can gradually be traced with such
distinctness, that there’ remains no doubt, that even
the coarser elastic fibres directly result from a chemi-
cal change and condensation of the walls them-
selves.* Where originally there lay a cell, provided
with a delicate membrane and elongated processes,
there we see the membrane gradually increasing in

Fic. 17,

Elastic networks and fibres from the subcutaneous tissue of the abdomen of
a woman. au, Large elastic bodies (cell bodies), with numerous anastomosing
processes. bb, Dense elastic bands of fibres on the border of larger meshes.
¢ ¢, Moderately thick fibres spirally coiled upat the end. d d, Finer elastic fibres,
at e with more minute spiral coils. 300. (From Virchow.)

thickness and refracting the light more strongly,
whilst the proper cell contents continually decrease
and finally disappear.

“The whole structure becomes in this way more

* Virchow, op. citat., p. 188,

THE CELL DOCTRINE. 73

homogeneous, and to a certain extent sclerotic, and
acquires an incredible power, of resisting the influ-
ence of reagents, so that it is only after long-con-
tinued action that even the strongest caustic sub-
stances are able to destroy it, whilst it completely
resists the caustic alkalies and acids in the degree of
concentration usually employed in microscopical in-
vestigation. The farther this change advances, the
more does the elasticity of the parts increase, and in
sections we usually find these fibres, not straight or
elongated, but tortuous, curled up, spirally coiled, or
forming little zigzags (Fig. 17, ¢,e). These are the
elements which by virtue of their great elasticity,
cause retraction in those parts in which they are
found in considerable quantity, as, for example, in the
arteries. The fine elastic fibres, which are those
which possess the greatest extensibility, are usually
distinguished from the broader ones, which certainly
do not present themselves in tortuous forms. As re-
gards their origin, however, there seems to be no
difference between the two kinds: both are derived
from the connective tissue cells, and their subsequent
arrangement is only a reproduction of the original
plan. In the place of a tissue consisting of a basis
substance and anastomosing reticulated cells, there
afterward arises a tissue with its basis substance
mapped out by long elastic networks with extremely
compact and tough fibres.” This may be looked
upon as the least well determined of the important
points of Virchow’s doctrine, though most German
histologists also favor it. Among these may be

y

‘

74 THE CELL DOCTRINE.

classed Kolliker,* C. O. Weber,t Leydig,t Fried-
reich,§ His,| Donders,] Wittich,** Bottcher,tt+ Bill-
roth,tt and Stricker. They are opposed by Schwann,
Reichert, and Henle, and find little favor among
English and American histologists.

A part of this system, also, according to Virchow,
are the so-called dentinal tubules, the Jacune and
canaliculi of bone, even the continuity traced by
Gerlach,§§ between the ciliated cells of the aqueduct
of Fallopius; that by Heidenhain||| and Briicke{ 4
between the lacteals and cylinder epithelium of the
intestinal villi of the rabbit, by means of corpuscles
of connective tissue ; in the epithelium of the endo-
cardium by Luschka;*** and the results of similar

* Kélliker, Manual of Human Microscopic Anatomy, p. 41,
1860. Also recent paper, in which he completely assents to Vir-
chow’s views, according to N. Y. Quart. J. Pschy. Med., July,
1869.

} Weber, C. O., Virchow’s Archiv, Bnd. xiii-xv.

{ Leydig, Handbuch der Histologie, 1856.

9 Friedreich, Virchow’s Archiv, Bd. xv.

|| His, Beitrige zur Normalen und Pathol. Histol. d. Cornea.
Basel, 1856.

{ Donders, loc. citat.

** Wittich, Virchow’s Archiv, Bd. ix.

+t Béttcher, Virchow’s Archiv, Bd. xiii.

tt Billroth, in Beitrage zur Pathol. Histol., 1858, admits all
but the tubular nature of the processes,

88 Gerlach, Mikrosk. Studien, 1858.

|||| Heidenhain, Moleschott’s Untersuchungen, Bd. iv, 1858, p.
261.

{{ Briicke, Moleschott’s Untersuchungen, Bd. viii, 1862, p. 496.
*%* Luschka, Virchow’s Archiv, Bd. ix, p. 569.

THE CELL DOCTRINE. 75

observations by Eckhart,* Billroth,t and Fried-
reich.{

The other fibrous element of areolar or connective
tissue, which forms the mass of its bulk, the pure
white fibrous or waving, does not, according to Vir-
chow, have its origin in cells, but is a modification
of a previously homogeneous intercellular substance,
deposited between the cells; a view which in its glar-
ing departure from the primary proposition that the
cell is the starting-point, and that every tissue is
composed of cells or some modification of cell forms,
presents one of the few inconsistencies traceable in
the theory of Virchow.

We think it proper, in a historical memoir of this
kind, to refer to some severe critical remarks which
appeared in the Edinburgh Medical Journal, of Feb-
‘ruary and April, 1869, in which Prof. Virchow is ac-
cused of appropriating the observations of Prof.
Goodsir as his own. That there are points in com-
mon, it will be recollected, and, also, that these are
Ist, the invariable origin of cells from previously ex-
isting cells, and 2d, the division of the tissues into
cell territories. Now on the one hand we deem that
the dedication of Virchow’s volume to Prof. Goodsir
is as handsome an accredit as could possibly be given
for whatever of common there may be in the writings
of the two authors, and on the other hand we have
seen that Martin Barry is acknowledged even by

* Eckhart, Beitrage Anat. und Physiol., 1855.
+ Billroth, Miiller’s Archiv, 1858.
{ Friedreich, loc. citat., p. 538.

76 THE CELL DOCTRINE.

Goodsir, to be the author of the “ first consistent
account of the development of cells from a parent
centre.” The idea of cell territories seems, however,
to have originated with Goodsir, nor do we believe,
for the reason stated, that Virchow intended to usurp
his prerogative. The merit of Virchow consists in
his application by actual demonstration of the first
of these points to so large a variety of physiological
and pathological processes, to which is added original
conception in the prominence given to the. connec-
tive tissué corpuscle and the canalicular system,
whatever may be the truth with regard to, either.

SARCODE OF DUJARDIN—PROTOPLASM OF MAX SCHULTZE.
1835-61.-

Dujardin* had, in 1835, discovered in the lower
animals a living, moving, contractile substance,
which he called sarcode. The peculiar appearances of
this substance attracted the attention of many ob-
servers, among whom were Kiihne, Reichert, Ecker,
Henle, Meyen, Huxley, Max Schultze, John Miller,
. and others.. It was thought peculiar to the lower
“animals, and there was assigned to it a property of

“irritability without nerves.” °

The observation of Siebold,+ that the yolk glob-
ules (vitelline spheres of the egg) of Planaria exhibit
contractions and expansions, which with suitable
care continue for hours, and the discoveries which fol-

* Dujardin, Ann. 4. Sciences Nat., tom. iii et v.
¢ Schultze, Max, Organis. d. Polythalamien, 1854,
{ Siebold, Froriep's Notizen, Nr. 380, p. 85.

THE CELL DOCTRINE. 77

lowed of similar movements and changes in form, in
colorless blood-corpuscles, pigment cells, and else-
where, led Kélliker* to express the conjecture that
the contents of all cells are contractile. Virchowt
attributed the ciliary movement to a contractile sub-
stance. Leydigt and Ecker considered the move-
ments of the yolk spherules as phenomena of life,
and Kiihne§ had studied physiologically and chemi-
cally, sarcode and muscular tissue, and pointed out
the similarity of the phenomena presented in the act
of dying, by both. But all considered sarcode as
something different from the animal cell, as a body
sui generis. .
According to Heeckel|| the protoplasm or sarcode
theory, that is, the theory that the albuminous con-
tents of animal and vegetable ceils as well as the
freely moving sarcode of Rhizopoda, Myxomycete, .
etc., are identical, and that in both cases this -albu-
minous material is the original active substratum of
all vital phenomena, was brought forward in its ele-
mentary form by F. Cohn§ in 1850, and by Unger in
1855.** Heeckel says also that it may be considered
one of the greatest achievements in modern biology
and one of the richest in results. It was further de-

* Kolliker, Wurzb. Verh., Bad. viii.

+ Virchow, Archiv, Band v., 1858.

t Leydig, Handbuch der Histologie, 1856.

@ Kiihne, Miill. Archiv, 1859, p. 817.

|| Quart. J. Mic. Sc. July, 1869, p. 228.

q F. Cohn, Nachtrige zur Naturgeschichte des protococcus plu-

vialis, Nova Acta Ac. Leop. Carol., vol. xxii, pars. 2, 1850, p. 605.

** Unger, Anatomie und Physiologie d. Pflanzen, 18655.

7*

78 THE CELL DOCTRINE.

veloped by Max Schultze in 1858, and finally estab-
lished by him in 1861.* He first showed the anal-
ogy between sarcode and the contents of the animal
cell, and that the entire infusorial world, simple or
compound, is made up of cells, thus extending the
typical formative element of Schwaun to the entire
organized creation.

The comparison between sarcode and the proto-
plasm of plants on the one hand, and that of animal
cells on the other, was also made by Pringsheim,+
E. Briicke,t E. Heeckel,§ and W. Kiibne,|| and by
their efforts, together with those of Max Schultze,
Unger, and Cohn, our knowledge of the indepen-
dent life of the cell was extended, in a very short
space of time, further than in the twenty years pre-
vious. J

The name protoplasm for a portion of the contents
of the animal cell had already been brought into use
by Remak, who extended it from the layer which
bore that name in the vegetable cell to the analogous
element in the animal cell. si

* Schultze, Max, Miill. Archiv, 1861, p. 17.

+ Pringsheim, Untersuchungen iiber d. Bau. u. d. Bildung 4d.
Pflanzenzellen, 1854.

t Briicke, E., Elementar-organismen, Wien. Sitzungsb., 1861.

3 Heckel, H., Die Radiolaren, 1862.

|| Kithne, W., Protoplasm und die Contractilitat. Lpzg.,-1864.

{ Stricker, S., Handbuch der Lehre von den Geweben des
Menschen und der Thiere. Leipzig, 1868, p. 8, German Ed.

** Sterling, J. H., As regards Protoplasm in relation to Prof.
Huxley’s Essay on the Physical Basis of Life. Edinburgh, 1869,
p. 14. Conf. also McNab, Monthly Microsc. J., No. xvii, vol.
iii, 1870, p. 838.

THE CELL DOCTRINE. 79

Pringsheim had also shown, in 1854, that no such
membrane as a primordial utricle existed, but that
all. within the cellulose wall of the living vegetable
cell was protoplasm and cell fluid, however complex
its composition.

‘“‘ He admitted that in the cortical layer of the pro-
toplasma a distinct arrangement into layers often
occurred, and these he distinguished as the cutaneous
and granular layers of the protoplasma, but he de-
nied that the primordial utricle could ‘be differen-
tiated as a membrane from the subjacent protoplasm.
If, in animal cells, partly from their relatively small
size, and partly from their greater average wealth in
protoplasma, it is more rarely possible to make a
sharp demarcation between a cortical layer of pro-
toplasm and a cell fluid, there nevertheless exists a
difference in the constitution of the former, such
that a cutaneous layer, destitute of or scantily sup-
plied with granules, incloses the remaining more
granular material. The white blood cell may serve
asanexample. This is, however, very different from
@ proper membrane.”*

Unger} (1855), had been struck with the close
similarity of the mobile phenomena of the Polytha-
lamize with those of the processes of protoplasm
stretched across the, cavity of many vegetable cells.
Although he had not personally investigated the for-
mer, he became convinced from Schultze’s description
that a resemblance amounting to identity existed

* Duffin, A. B., On Protoplasm. Quart. Jour. Mic. Sci., N.
S., vol. iii, 1868, p. 252.
+ Unger, op. citat., p. 280.

80 THE CELL DOCTRINE.
between their movementsand the protoplasm streams
of vegetable cells.* .

Leydig,t in 1856, claimed for the contents of the
cell a higher dignity than for the membrane or cell-
wall. He claimed that a cell was but protoplasm
(klumpchen-substanz) inclosing a nucleus. The cell
membrane, according to him, was simply the hard-
ened periphery of the substance of the cell.

To Max Schultze, however, as already stated, be-
longs the credit of having fully overturned the vesic-
ular idea of cells. Jn 1861,t he insisted upon some
modification of prevailing views, respecting the rela-
tion of cell-wall to cell contents, and contended for a
higher position for that part of the cell correspond-
ing to the protoplasm of Von Mohl (that within the
so-called primordial utricle), and showed how a care-
ful study of the phenomena presented by the pseudo-
podia, extended by the various Rhizopods, might aid
in clearing up the life of the elements of the cell.

He also defined the cell as “ protoplasm surround-
ing a nucleus.” The importance of this definition,
as stated by Stricker,§ lay not so rnuch in the fact
that many cells were denied a cell-wall, as that
the so-called cell contents could now be made to har-
monize with the animal primordial substance or sar-
code. Schultze illustrates his definition by the em-

* Duffin, A. B., loc. citat., p. 252.

+ Leydig, op. citat.

{ Schultze, Max, Ueber Muskelkérperchen, in Reichert and Du-
bois Reymond’s Archiv, 1861.

@ Stricker, op. citat., 56. (German Ed.)

THE CELL DOCTRINE. 81

‘
bryo cells resulting from the segmentation of theovum,
as typical cells, which are thus composed of proto-
plasm surrounding a nucleus, which nucleus, as well
as protoplasm, are products of like constituent parts
of another similar cell. ‘The cell leads in itself an
independent life, of which the protoplasm is espe-
cially the seat, although to the nucleus also undoubt-
edly falls a most important, though not yet precisely
determined role. Protoplasm is for the most part
no further distinct than that it will not commingle
with the surrounding medium, and in the peculiarity
that with the nucleus it forms a unit. Upon the
surface of the. protoplasm, there may form a mem-
brane, which, although derived from it, may be chemi-
cally different, and the assertion that it is the begin-
ning of a retrogression may be defended. A cell with
a membrane cannot divide itself, unless the proto-
plasm within the meinbrane divides itself. <A cell
within a membrane chemically different from proto-
plasm, is like an encysted infusorium.”*

Briicket went even further in his definition, and
said that it was not shown that the nucleus even is
an essential element of the cell. In proof of which
he adduces the cells of cryptogams and says: “ We
have no positive knowledge either of the origin or
function of the nucleus, and, indeed, the constaney
of its occurrence seems subject to certain limitations
if we take into consideration the cells of cryptogams,

* Schultze, Max, Protopl. d. Rhizopoden. Leipzig, 1863.
{ Briicke, E., Die Elementar-organismen, p. 18-22. 1861.

82 THE CELL DOCTRINE.

and do not start out with the belief that the nucleus
is there even though we do not sce it.” Facts in
justification of Briicke’s doubt are adduced by
Stricker* in the discovery by Max Schultze,t in the
Adriatic Sea, of a non-nucleated ameba (Amoeba
porrecta), by Heckel,t in the Mediterranean, of a
non-nucleated protozoon (Protogenes primordialis),
and by Cienkowsky§ of two non-nucleated monads,
namely, Monas amyli and Protomonas ainyli. Heee-
kel says of his Protogenes primordialis that it mul-
tiplies by division. Stricker’s|| own observations on
the fecundated egg of the frog, which confirm those
of Von Baer, incline him to adopt the view of Briicke,
and to omit the nucleus in a theory of elementary
organization.J

Such is the history of and such the properties of
the substance known as “ protoplasm.” But of late

* Stricker, op. citat., p. 6. (German Ed.)

+ Schultze, Max, Organis. d. Polythalam. 1854.

{ Heckel, Zeitschr. f. w. Zoolog., 1865, Bd. xv.

@ Cienkowsky, Max Schultze’s Archiv, 1865.

|| Stricker, op. citat., New Sydenham Society’s Translation,
vol. i, p. 8, London, 1870. See also Stricker’s paper On the De-
velopment of the Simple Tissues, in vol. iii, London, 1873.

{| Heckel! also considers that by no phenomena is the correct-
ness of the ‘* protoplasm theory ’’ so thoroughly proved, and at the
same time in so simple and unassailable a manner, as by the vital
phenomena of the Monera, by the processes of their nourishment
and reproduction, sensitiveness and motion, which entirely pro-
ceed from one and the same very simple substance, a true ‘ primi-
tive slime.’

* Heckel, Ernst, Monograph on the Monera, and Remarks on the Protoplasm
Theory. Q. Jour. Mic, Sci., N.S. vol. ix, 1869.

THE CELL DOCTRINE. 83

years another meaning has been given to the term,
an anatomical one, in which the original general ap-
plication has been altogether ignored, and that is
that part of the cell outside of the nucleus without re-
gard to the properties of the matter, corresponding
to the “cell contents ” in cells which have a cell-wall.
Thus we speak of the protoplasm of the colorless
corpuscle or the liver-cell, or of the squamous epithe-
lial cell, whereas the properties of the substance thus
named are vastly different. In the former two in-
stances living growing matter is meant, in the latter
dead formless material, incapable of growth and re-
production through its own inherent properties.
These differences should be always remembered.

With these general considerations in the history
of “ protoplasm,” we are the better prepared to take
up the theory of

DR. BEALE, 1861.

_In April and May, 1861, Prof. Lionel S. Beale de-
livered the lectures before the Royal College of Phy-
sicians of London, in which he promulgated the
views which have since been further elaborated and
become permanently associated with his name. These
views were published in part, in Beale’s “ Archives
of Medicine,” and in September, 1861, in a volume
“On the Structure of the Simple Tissues of the
Human Body,” in the preface to which he says, “ he
thinks it right to state that the conclusions which
have now assumed a definite form have gradually
grown upon him during the course of observations

84 THE CELL DOCTRINE.

extending over a period of several years. In fact
some of the drawings in this volume, and others
which have'been published elsewhere, equally favor-
able to this view, were made long before any specific
theory had been arrived at.”

‘The “cel/,” or “ elementary part,” as Dr. Beale pre-
fers to call it, is composed of matter in two states,
matter which is forming, and matter which is formed ;
matter which has the power of growing by produc-
ing matter like itself out of pabulum or food, and
matter which possesses no such power, but results
from the death of the forming matter. The former
is known as germinal or living matter, the latter as
formed matter. The former, in varying quantity in
different cells, is central in its situation (see frontis-
piece, Fig. 17), and includes what, has been called by
others nucleus, cell contents, protoplasm, endoplast.
The latter, also present in different quantity in differ-
ent cells, is peripheral (frontispiece, Fig. 17), and in-
cludes what is known as cell-wall, periplast, inter-
cellular substance, and products of secretion.

In its structural characters, germinal matter is soft,
transparent, colorless, and as far as can be determined
by the highest powers, structureless, being visible
only through its difference in refracting power as
compared with the menstruum in which it floats, or
by the granular matter it may entangle; and these
characters are the same at every period of its exist-
ence. In the simplest vegetables they may be studied,
in the thallus of the sugar fungus, among the lowest
animals, in the amceba (frontispiece, Fig. 16), and in
higher animals in the mucus-, pus-, or white blood-cor-

THE CELL DOCTRINE, 85

puscles (frontispiece, Fig. 10), all of which are com-
posed almost purely of germinal matter; the very thin
periphery of formed material being scarcely appreci-
able or distinguishable from the diffraction band.

In its endowments and properties, germinal matter
is acting, living, growing, and moving, through some
inherent power of its own. It alone, as stated, is
capable of producing material like itself out of pabu-
lum, and of multiplying by division, or dropping off
of a portion of itself, which portion immediately
assumes an independent existence, and grows, main-
tains, and reproduces itself like the parent germinal
matter. It is also capable of being stained by an
ammoniacal solution of carmine, and the younger it
is, or more recently formed, the deeper is the stain it
assumes. And since the latest formed always ap-
pears in the centre of the mass, successive tints, or
zones of color, will often be produced in the staining
process, growing deeper from without inward, as
seen in Fig. 17 of the frontispiece.

It has been stated that what is called nucleus by
Virchow and others, is included in germinal matter.
This is true, though the nucleus is not always the
whole of the germinal matter. There may be other
older germinal matter beyond the nucleus, on its
way to conversion into formed material, but still
germinal matter, which assumes a tint with carmine,
but not so deep as the nucleus. Thus, the entire mass
of the pus-corpuscle (frontispiece, Fig. 10), except,
perhaps, its extreme periphery, is germinal matter,
yet there is within this another younger portion of
germinal matter, taking a deeper tint with carmine,

8

86 THE CELL DOCTRINE. i

but which alone of the elements of this cell we are in
the habit of calling “ nucleus.” The “ nucleus,” then,
is nothing but a new centre of germinal matter, and
the “nucleolus” is a younger centre. And there may
even be within this a still younger portion of living
matter, taking even a deeper stain, which might be
called a “‘nucleoleolus.” By this staining process may
we distinguish the nucleolus from a minute oil-drop
often mistaken for it, and which will not admit of
being stained.

On the other hand, germinal matter in a compar-
atively quiescent state is often quite destitute of
nuclei. But let the mass be freely supplied with nu-
trient matter, and nuclei and nucleoli rapidly make
their appearance.

So with regard to the “cell contents” over and
above the nucleus, although they may all be germinal
matter, yet this is not necessarily the case. Thus
in the white blood-corpuscle and mucus-corpuscles,
what Virchow would consider cell contents is all ger-
minal matter ; but the superficial epithelial cell lining
the interior of the mouth has its nucleus alone com-
posed of germinal matter, and much that has been
described as cell contents is really formed matter.
(Figs. 5 and 6 of frontispiece.) More nearly does the
germinal matter of Beale* correspond with the “proto-
plasm” of Max Schultze, with which, indeed, it seems
identical, except that the latter observer seems some-
what at a loss how to dispose of the nucleus, of

* Beale, Protoplusm; or, Life, Force, and Matter. London,
1870, p. 38.

THE CELL DOCTRINE. 87

which he does not speak as a new or young centre
of protoplasm.

Formed material, instead of being active, so far as
the vital acts described as characteristic of germinal
matter are concerned, is passive, non-acting, dead, and
can only increase at the expense and death of the
germinal matter, on the periphery of which it is
formed. It differs widely in its appearance, and is
often “structured ” as in muscle and nerve, but not
necessarily so, as is seen inthe intercellular sub-
stance of hyaline cartilage. It possesses also cer-
tain properties, different in different situations, and
widely different also from those of germinal matter.
Thus it is contractile in the sarcous tissue of muscle,
exhibits neurility in the nerve, is protective in epi-
thelium, is diffluent as the formed material of the
milk-cell (milk), and in the formed material of the
liver-cell (bile). Again, it is hard and elastic in the
intercellular substance of cartilage and epidermis,
horn and nails. It does not become stained on being
soaked in weak solution of carmine in ammonia, and
if by reason of the strength of the solution it should
happen to be stained, the color will wash out on
soaking in glycerin, which is not the case with the
coloring of the germinal matter.

The cause of this permanent staining of the ger-
minal matter by an ammoniacal solution of carmine,
is thought by Dr. Beale to be due to an acid reaction
of this matter, in consequence of which the carmine
is precipitated from its alkaline solution. This view
would seem to be confirmed by the researches of:
Ranke on the Reaction of the Tissues.

88 THE CELL DOCTRINE.

The size of the elementary part, as thus composed,
is extremely various. The smallest’ particles of ger-
minal matter, measured by Dr. Beale, are less than
rovrss Of an inch in diameter, and would not be
called cells in the ordinary sense of the word, yet
they are functionally such; that is, they grow, multi-
ply by division, and under appropriate circumstances
assume the characters of fully formed cells. On the
other hand, the largest epithelial cells, including
their germinal matter and formed material, are often
as large as the 31, of an inch in scare or larger ;
cells of morbid growths are sometimes 345, while the
human ovum, whieh is a typical cell, varies from the
ghy to zhy of an inch. -Pure germinal matter is
rarely seen in masses as large as the 3}, of an inch
in diameter, without breaking up into smaller par-
ticles of germinal matter, and as constituting the
nuclei of fully formed cells, is usually from gy\55 to
3005 Of an inch in diameter.

The method of production of formed material is best
studied in the epithelial structures, particularly in
the epithelium lining mucous cavities, of which sec-
tions may be easily made down to the vessels whence
their nourishment is obtained. In the deep layers,
next the nutrient surface, the cells will be found to
consist of almost pure germinal matter (frontispiece,
Fig. 1), imbedded in a soft, mucus-like, yet continuous
formed matter. WThese masses of germinal matter
divide and subdivide, pushing the older masses to-
wards the surface, to make up for those which are
constantly exfoliated. While this is going on, how-
ever, the germinal matter keeps increasing in size

THE CELL DOCTRINE. 89

until the cells arrive half way towards the surface,
by absorption of nutrient pabulum, which has to dif-
fuse itself through any formed material already ex-
isting. At the same time, a portion of the germinal
matter is being converted into formed material,
which accumulates on its surface, within that already
formed, stretching it, and becoming more or less in-
corporated with it. Thus, both constituents of the cell
increase up to a certain point, the cells constantly growing
in consequence. As new cells are, however, produced
from below, the older ones are removed farther and
farther away, the formed matter becoming more and
more impervious to nutrient pabulum. At length a
point is attained when the entire cell ceases to in-
crease in size, since no pabulum reaches the masses
of germinal matter, though the latter is still being
converted into formed material. Hence, the masses
of germinal matter actually grow smaller as the cell
increases in age; and when the periphery is reached,
there remains but a small nucleus of germinal matter,
with a large quantity of formed material. Thus, we
are enabled to judge of the age of the cell by the
relative quantity of germinal matter and formed
material ; if the former be large, and the latter small,
the cell is young, whereas, if the opposite relation
exists, the cell is old and almost ready to exfoliate.
But exfoliation in health probably does not take
place until the last particle of germinal matter dies,
and the entire cell becomes a mass of passive, life-
less, formed material.

The production of formed material from germinal
matter may also be studied in the conversion of the

8*

90 THE CELL DOCTRINE.

white blood-corpuscle into the red. In the spring of
the year many white corpuscles can be found in the
blood of the frog and newt, undergoing conversion
into formed material at their edges, where the usual
granular appearance is being substituted by the
smooth and slightly colored. This goes on until all
except the nucleus is thus converted. In mammalia
this change goes on until the whole white corpuscle
is thus converted into the red.

Secondary Formed Material.—There are certain
kinds of formed material to which this term is ap-
plied by Dr. Beale. These are the oil of the fat-cell
or vesicle and the starch-granule of the vegetable
cell. It results, as does all formed material, by a con-
version of the germinal matter into this special form.
The young fat-cell, as all young cells, is almost pure
germinal matter; as it grows older, however, and is
exposed to oxidizing influences, the conversion of
germinal matter takes place, partly into the cell-wall
proper of the fat-vesicle, and partly into the second-
ary formed material or oil, until finally it becomes a
mere dot on the inner surface of the cell-wall, or dis-
appears altogether.

The increase of cells, according to Beale, takes place
in several ways; every cell coming from a pre-existing
cell, but the germinal matter is always the portion
in which it originates.

There is not generally a symmetrical division of
the nucleus into two, and these into four, as is so
often described, and as is often seen in the vegetable
cell, but there is rather a budding, and subsequent
dropping off of the portions of germinal matter

THE CELL DOCTRINE. 91

‘which is to produce the new cell, and which almost
always assumes the spherical form when allowed to
float freely. (See Figure 10 of frontispiece.) The
formed material is never active, according to Beale,
but entirely passive in the process of cell multiplica-
tion.

Nutrition of Cells.—So, too, in the nutrition of the
cell, the germinal matter is the sole active agent.
The formed material may act as a filter to the nu-
trient matter, but is quite passive. The pabulum,
which is coursing through the bloodvessels, becomes
converted into germinal matter, which in turn be-
comes formed material, aud so long as this is kept
up, the cell continues to grow. The course taken by
the pabulum, and the order of conversion, is shown
by the arrows, in Figure 17, of frontispiece, and will
be readily understood by reference to the explanation.
Occasionally, and especially in disease, the formed
material may become the pabulum for rapidly mul-
tiplying cells, and thus be consumed.

Intercellular substance has already been spoken of
as formed material. We have it most strikingly
present in the white fibrous tissue, or tissue of ten-
dons, and in hyaline cartilage. If the former be
stained by carmine, and examined in thin section
under the microscope, it will be found composed of
beautiful’ bands of gently waving fibrous tissue, or
tissue exhibiting a fibrous appearance, at varying in-
tervals in which are noted nuclear masses of germinal
matter, which have assumed the tint of carmine. Or,
if dilute acetic acid be added to the specimen, the
fibrous appearance will at once become homogeneous,

92 THE CELL DOCTRINE.

while the nuclei will be rendered distinct, and clearly
visible. In young tendon (frontispiece, Fig. 11), the
masses of germinal matter will be found very abun-
dant, and the intercellular fibroussubstance in smaller
quantity than in old tendon where the masses of ger-
minal matter occur only at long intervals. These
masses of germinal matter, or connective tissue cor-
puscles, it will be recollected, are considered by Vir-
chow as perfect cells, presenting cell-wall, cell con-
tents, and nucleus, and the fibrous intercellular sub-
stance as a subsequént modification of a homogene-
ous matrix, deposited between the cells by the blood-
vessels. These connective tissue corpuscles are re-
garded by Beale as simple masses of germinal matter,
the conversion of which into formed material pro-
duces the fibrous intercellular substance, as seen in
Fig. 11, frontispiece, and between which and the
intercellular substance immediately adjoining, there
is no line of separation, constituting a cell-wall.

As the tendon grows older, the masses of germinal
matter become less abundant, because a larger num-
ber have been totally converted into formed material ;
and the bands of indestructible material which some-
times join them, and which are considered by Virchow
as a part of his canalicular system, are, according to
Beale, nothing but imperfectly converted formed material,
or rather germinal matter, which has not been con-
verted. While the twisted and curling cord-like
fibres of the so-called yellow elastic tissue, also con-
sidered by Virchow as a part of his canalicular sys-
tem, are thought by Beale to be composed in part of
true yellow elastic tissue, such as is found in the

THE CELL DOCTRINE. 93

ligamentum nuche, and likewise formed from nuclei
(frontispiece, Fig. 14}, but in part also of the remains
of nerves and vessels, which were active at an earlier
period of life.*

So, also, with hyaline cartilage. According to Beale,
the intercellular substance of cartilage results from
the conversion of the so-called cartilage-corpuscles or
cells into formed material, and here also the germinal
matter is directly continuous with the matrix, no
proper cell-wall intervening.

Cartilage is not to be considered as a distinct class
of tissue from epithelium, nor can the latter, in all
cases, be distinguished from cartilage by the exist-
ence of separate cells, since in many forms of epi-
thelium, at an early period of existence, the formed
material corresponding to the masses of germinal
matter is continuous throughout, and presents no
indication of division into cells.t A “cell,” or “ele-
mentary part,” then, of fully formed tendon or carti-
lage, would consist of a portion of germinal matter,
with a proportion of formed material about it, ex-
tending to a line midway between that mass of ger-
minal matter and the masses immediately adjacent,
of which the cartilage or tendon is composed; and
such a line would correspond to the outer part of
the surface of an epithelial cell.t In very young car-
tilages, as in very young epithelium, the cells consist

* Beale, On the Structure and Growth of the Tissues, and on
Life. London, 1865, pp. 95, 96, and 101.

+ Beale, Protoplasm; or, Life, Force, and Matter. London,
1870, p. 51.

t Beale, Protoplasm, pp. 51-2.

94 THE CELL DOCTRINE. :

of germinal matter only, with a small quantity of
soft formed material intervening ; and to understand
the true relation of the cells to the intercellular sub-
stunce, the tissue should be studied at different pe-
riods of its growth.

So, too, a “cell” or elementary part of muscle or
nerve, would consist of a mass of germinal matter
(the eo-called nucleus), with a portion of muscular or
nervous tissue corresponding with it, and with which
it is uninterruptedly continuous.

In the formation of the contractile tissue or muscle,
the germinal matter seems to move onward, under-
going conversion at its posterior part, into the mus-
cular tissue, while it maintains itself by absorbing
and converting pabulum. This will be understood by
reference to Fig. 18, of frontispiece. The fibres of
yellow elastic tissue are formed in precisely the same
manner. (See frontispiece, Fig. 14.) Nerve fibres,
which in their completed state consist almost wholly
of formed material, are similarly produced. In the
yount state, the fibre is composed of masses of germi-
nal matter, linearly arranged, and in close proximity.
As the conversion takes place and the fibre is pro-
duced, these become more widely separated, and the
tissue resulting from such conversion is nerve (fron-
tispiece, Fig. 15).

The “ Cell” or * Elementary Part” in Disease.

Here, as in normal nutrition, the germinal matter
is alone active. It is impossible to state precisely
every instance, but it is probable that in the majority

THE CELL DOCTRINE. 95

of cases of disease, the morbid state consists essen-
tially inca modification of the healthy nutrition of
the cell, that is, the cell is made to grow more or
less rapidly, or is perverted in its mode of growth,
though it is likely that within certain limits, the
conditions under which cells ordinarily live may be
modified without deviation from health. But in in-
flammatory processes attended by local products, as
pus or lymph, and in the production of tubercle and
cancer we see the results of excessive multiplication
and perversion of germinal matter consequent upon
the appropriation of an excess of nutrient pabulum.
In other instances, as cirrhosis, where there is shrink-
ing, and hardening, and wasting, we see the effects
of a diminished supply of pabulum, either through
a diminution in the quantity supplied, or an imper-
meability in the septum through which it is com-
pelled to pass.

An increased supply of pabulum may be admitted
to germinal matter, either in consequence of the
removal of barriers through which it is ordinarily
compelled to pass, or in consequence of the nature
of the fluids by which it is bathed. A simple illus-
tration is seen in suppuration in epithelium, or the
germinal matter of any tissue; for, according to
Beale, suppuration and morbid processes generally,
are not restricted to any one kind of germinal
matter, as the connective tissue corpuscle, but may
occur in all germinal matter to which the conditions
are supplied. Using epithelium by way of illustration,
as the result of the increased supply of pabulum, the
germinal matter first grows, as seen in the frontispiece,

96 THE CELL DOCTRINE.

Figs. 7 and 8, then in the luxuriance of its growth,
even at the expense of the formed matter, sends out
buds or processes, which soon drop off and become
separate pus-corpuscles. (Figs. 9 and 10.) These
are produced so rapidly that there is not time for
formed material to form upon their surface in any
quantity, and they have not time, therefore, to pass
on into epithelium. Hence pus-corpuscles are almost
pure germinal matter. So soon as the process ceases,
in consequence of the supply of pabulum being di-
minished, the germinal matter multiplies less rapidly ;
opportunity is permitted for the production of formed
material on its periphery, and the cell now passes
through the different grades of epithelium, as de-
scribed on pages 88, 89, and 90. The pus-corpuscles
are analogous to the deepest layers of epithelial cells
there referred to, which deep cells are in fact the
** mucus-corpuscles,” so-called, well known to be mor-
phologically identical with pus-corpuscles; the for-
mer being simply the young epithelial cell on its way
to become perfect epithelium, while the latter is the
same also, though never allowed to pass into the
perfectly formed state.

Again, in pneumonia, and here we note where the
paths of Virchow and Beale separate more widely, the
so-called “ exudation,” or product which fills up the
vesicular portion of the lung, is regarded by Beale as
the result of a proliferation of minute particles of
germinal matter (very much smaller than white blood-
corpuscles), which have passed out through the capil-
lary walls with the liquor sanguinis.

In all inflammatory processes and fevers, this is

THE CELL DOCTRINE. 97

believed by Dr. Beale to take place to a greater or
less extent, the little masses of germinal matter or
nuclei in the capillary walls also taking part, often
increasing in size to such degree that they materi-
ally obstruct the passage of the blood, and by drop-
ping off portions give rise to bodies floating in the
blood precisely similar to white blood-corpuscles, or
pus-corpuscles; indeed, Dr. Beale considers that this
may be one of the sources of origin of the white
blood-corpuscle.*

So, also, tubercle is believed by Dr. Beale to result
either from the multiplication of masses of germinal
matter which have passed through the capillary
walls from the blood, or from the masses of germinal
matter usually termed nuclei, in connection with
the capillary walls. He says, in illustration: “In a
case of tubercle, which was very rapidly developed
upon the surface of the pia mater, in a man of tuber-
cular constitution, I proved most distinctly that the
tubercles were connected with the vascular walls,
and that if the nuclei had not given origin to them,
they were certainly implicated. My own opinion is,
that these nuclei gave origin to the tubercle-cor-
puscles, in consequence of receiving from the blood
peculiar nutrient matter. In the lung I have seen
appearances which point to a similar conclusion.”{
Would not these views arise from appearances pre-

* Beale, Microscope in Clinical Medicine, third ed. London,
1867, p. 166.
+ Microscope in Clinical Medicine, third ed., 1867, p. 205.
9

98 THE CELL DOCTRINE.

cisely analogous to those represented as giving sup-
port to the view, that tubercle originates in the
perivascular sheaths of bloodvessels? The views of
Beale, H. Charlton Bastian,* and Cornil,+ would
then constitute simply different modes of expression
of the same truths.

ROBIN,t 1867.

Robin, who may be considered the mouthpiece of
the French school of histologists, reduces the human
body to elementary parts, usually microscopic, which
he calls anatomical elements. The forms he makes
threefold,—fibres, tubes, and cells.

The fibres are generally of considerable length,
sometimes extending from the lower part of the
spinal cord to the extremity of the foot. Their di-
ameter is, however, small, often not exceeding .001
millimeter, or .00003987 of an inch.

The tubes offer as objects of study the walls and
the cavity.

* Bastian, H. C., Tuberc. Meningitis, Edinb. Med Jour., 1867,
p. 875.

+ Cornil, Tubercle in Connection with the Vessels, Archiv. de
Phys. Norm. et Path., Jan. et Fev., 1868.

t Our information with regard to M. Robin’s views, is derived
from an admirable exposition of them published in vol. iv, 1867,
of the New York Medical Journal, by Dr. William T. Lusk, who
there states that he has them mainly from a course of familiar
and private instruction, furnished to him by M. C. H. Georges
Pouchet, assistant to M. Robin, Lecturer upon Anatomy and His-
tology to the Ecole Pratique, author of ‘* Un Précis d’Histologie,”
ete., and son of the eminent physiologist, Prof. F. A. Pouchet; so
thut they may be said to be the views also of the elder Pouchet.

THE CELL DOCTRINE. 99

The cells of vegetables have a wall, a cavity, and
contents (air, oil, etc.). The cells of animals, on the
coutrary, are, as a rule, homogeneous. Animal cells
containing a cavity are only found exceptionally.
The substance of cells is ordinarily granular. Most
cells contain an ovoid nucleus more granular than
the substance itself.

In all cells the nuclei afford different chemical
reactions from those of the substance of the element.
Each cell is an independent organism, passing through
various stages of development, from birth to death.

The birth (origin) of the elements takes place by
ist, segmentation ; 2d, genesis; 3d, epigenesis; 4th,
germination.

Ist. SegmentationThe human ovum is a small
hollow sphere, containing in its interior the vitellus
or yolk, which consists of granular matter in a hya-
line substance. At the end of a certain time par-
ticles of the granular matter approximate, unite, and
form a nucleus in the vitellus. Next, the nucleus
elongates, takes an hour-glass form (biscuit), then .
divides. The division of the yolk occurs simulta-
neously. In the same way, the division takes place
into 4, 8, 16,and more parts. ‘These divisions of the
vitellus have received the name vitelline globules.
Their mode of formation is called segmentation.

2d. Genesis.—When the vitelline globes have be.
come very small by successive segmentation (diame-
ter .008 millimeter, 00031396 of an inch), these little
bodies take the name of embryonic cells.

According to M. Robin, these cells dissolve. From
the fusion results a blastema, in the midst of which

100 THE CELL DOCTRINE.

nuclei make their appearance. This is known as
genesis. It is the second and most frequent mode of
the formation of anatomical elements. It is char-
acterized by.the appearance of an anatomical ele-
ment in ‘a fluid termed blastema, in which the
element did not previously exist.

3d. Epigenesis.—When the embryonic cells dis-
solve, the embryo-plastic nuclei are produced by
genesis in the blastema which results from their
fusion. Then little cone-like prolongations of trans-
parent matter are observed at the extremities of the
nuclei, giving rise to the fusiform bodies, which are
the connective tissue corpuscles. This mode of forma-
tion by growth upon another element is known as
epigenesis, and is the mode in which connective tissue
is developed. The prolongations of these fusiform
bodies constitute the non-elastic fibres or white
fibrous tissue element of connective tissue. Some-
‘times the substance deposited by epigenesis upon the
nucleus has several prolongations, forming a stellate
cell or connective tissue corpuscle. These fusiform
and stellate cells are likewise known as embryo-plastic
or fibro-plastic bodies, and this latter term is a most
common one in French histology.

The elastic fibres of connective tissue are likewise
formed by epigenesis, but upon special nuclei, and
the prolongations are insoluble in acetic acid.

There is an early period of fetal life, previous to
the formation of connective tissue, in which we tind
only embryo-plastic nuclei and fusiform bodies in
amorphous matter. This is called embryo-plastic
tissue. Growth at this epoch is most rapid, the

THE CELL DOCTRINE. . 101

fostus reaching in a short space of time the dimen-
sion of .030 millimeter (.0118 of an inch).

4th. Germination.—This is very frequent in vege-
tables, but in animals only one example is known,
viz., at a period previous to the fecundation of the
guna Before segmentation takes place the vitellus
is observed to retract. The hyaline substance pushes
out a prolongation, which becomes round, separates,
and constitutes, an independent muatomical element
exterior to the vitellus, and bearing no part in the
future development of the ovum.

The following account of certain special elements
illustrates and further explains the views of M.
Robin. Red blood-globules (hematies), diameter, .007
millimeter (3355 of an inch); thickness .002 milli-
meter (;343, of an inch). Blood-globules are elastic,
—a property enabling them to elongate, and pass
through capillaries which have a calibre less than
the diameter of the blood-globule. They are homo-
geneous thronghout—i.e., have no cell-wall. Blood-
globules are formed by genesip in the blood plasma. In
the foetus they make their appearance before the
white blood-globules (leucocytes). In man there are
two kinds of red blood-globules, viz.: first, embry-
onic; second, normal. The embryonic blood-globules
are double the size of the normal ones. They have
a slightly granular nucleus, situated nearly in the
centre, which is insoluble in acetic acid. The nor-
mal blood-globules are not a transformation of the
embryonic. They appear by genesis in the midst of
the blastema of the blood. After the fourth month,
the embryonic globules cease to form, and as the

gx

102 THE CELL DOCTRINE.

mass of the blood increases, the proportionate num-
ber diminishes with great rapidity.

Leucoeytes, or white blood-globules, are found in
many tissues, in the blood, on the surface of mucous
membranes; in a word.they are the pus-corpuscles. In
form, they are round, with pale, well-defined bor-
ders, and contain extremely fine gray granules. They
possess a very thin envelope, and a granular cell con-
tents. The normal diameter is .008 millimeter
(sagy Of an inch). On the addition of water, the
leucocytes swell, the granular particles are agitated
by a peculiar movement (first observed by Brown),
and finally, a considerable number of these particles
unite, so as to form two or three little masses, that have
been mistaken for nuclei. Upon the addition of acetic
acid the same reaction follows, but with greater
rapidity.

The mode of production may be followed, step by
step, upon the surface of wounds, especially little
ones. At first a hyaline liquid appears. At the end
of a couple of hours, this liquid becomes finely gran-
ular, and then all at once, in the midst of the granu-
lations, we perceive small granular bodies analogous
to leucocytes, offering the same chemical reactions,
but measuring only .003 millimeter (.000118 of an
inch) in diameter. They are, iv fact, leucocytes of
young growth. When leucocytes are retained in the
economy, as in shut sacs, they increase in size, and
reach a diameter of .012 millimeter (,;'5y of an inch),
Then they fill with fat-granules, and are known as
corpuscles of inflammation (exudation corpuscles, com-
pound granule-cells). Finally the substance and in-

THE CELL DOCTRINE. 103

vesting membrane of the leucocytes disappear, the
granules dissolve and are reabsorbed.

Capillaries.—The finest capillaries are anatomical
elements of tubular form, with transparent resistant
walls which measure .001 mm. (.00003937 of an inch)
in diameter. These walls contain granular ovoid
nuclei, which project, sometimes exteriorly, some-
times upon the inner surface of the tubes. These
nuclei measure .006 mm. (.00023622 of an inch) in the
transverse, and .008 mm. (.00031596 of an inch) in
the long diameter. Their long axis is parallel to
that of the vessel. The finest capillaries have a
diameter of .007 nm. (.0003756 of an inch), leaving
a calibre (after deducting the walls) of .005 mm.
(.00019685 of an inch), or .002 mm. (.00007874 of an
inch) less than the average diameter of the blood-
globules which traverse them.

They are formed as follows: Ist. In new tissues,
hollow projections push out from contiguous capil-
laries, which meet and unite together. 2d. A solid
filament forms, in which nuclei-make their appear-
ance. Subsequently, the filament becomes hollow,
and its nuclei remain the nuclei of the capillary.

A single perusal of these views as thus illustrated,
will convince the reader that spontaneous formation is
the prevailing mode of origin of the elements of
tissues, according to the French school. Such pe-
rusal cannot fail to convince the reader also of the
accuracy of description of the fully formed elements
described by Robin.

104 THE CELL DOCTRINE.

PROF. HUXLEY,* 1869.

There is one kind of matter which is common to
all living beings, and that matter is “ protoplasm,”
the scientific name for “the physical basis of life.”
In illustration from vegetable life, each stinging
needlé or hair of the common nettle consists of a
very delicate outer case of wood, closely applied to
the inner surface of which is a layer of semifluid mat-
ter, full of innumerable granules of extreme minute-
ness. This semifluid lining is protoplasm, which thus
constitutes a kind of bag, full of a limpid fluid, and .
roughly corresponding in form with the interior of
the hair which it fills. When viewed with a suffi-
ciently bigh magnifying power, the protoplasmic
layer of the nettle hair is seen to be in a condition of
unceasing activity. Local contractions of the whole
thickness of its substance pass slowly and gradually
from point to point, and give rise to the appearance
of progressive waves, just as the bending of successive
stalks of wheat by a breeze produces the apparent
billows in a wheat-field.

But in addition to these movements, and inde-
pendently of them, the granules are driven, in rela-
tively rapid:streams, through channels in the proto-
plasm which seem to have a considerable amount of
persistence. The currents in adjacent parts com-
monly take similar directions, coursing in a general
stream up one side of the hair and down the other,

* Protoplasm ; or, The Physical Basis of Life. A Lecture by
Prof. Huxley, delivered in Edinburgh, November 18th, 1868.

THE CELL DOCTRINE. 105

though partial Nek also exist which take differ-
ent routes; so that sometimes trains of granules
may be seen coursing swiftly in opposite directions,
within a twenty-thousandth of an inch of each other;
and occasionally opposite streams come in direct col-
lision, and after a longer or shorter struggle one pre-
dominates. The cause of these currents seems to lie
in contractions of the protoplasm which bounds the
channels in which they flow, but which are so minute
that the best microscopes show only their effects and
not themselves.

Among the lower plants it is the rule rather than
the exception, that contractility should be still more
openly manifested at some periods of their existence.
The protoplasm of Alge and Fungi becomes, under
many circumstances, partially or completely freed
from its woody case, and exhibits movements of its
whole mass, or is propelled by the contractility of
one or more vibratile cilia.

In illustration of animal protoplasm, Prof. Huxley
adduces the colorless corpuscles of the blood, which,
under the microscope, at the temperature of the body,
exhibit a marvellous activity, changing their forms
with great rapidity, drawing in and thrusting out
prolongations of their substance, and creeping about
as if they were independent organisms. ‘“ The sub-
stance which is thus active is a mass of protoplasm,
and its activity differs in detail rather than in prin-
ciple from that of the protoplasm of the nettle. Under
sundry circumstances the corpuscle dies, and becomes
distended into a round mass, in the midst of which
is seen a smaller spherical body, which existed, but

106 THE CELL DOCTRINE.

was more or less hidden, in the living corpuscle, and
is called its nucleus. Corpuscles of essentially similar
structure are to be found in the skin, in the lining
of the mouth, and scattered through the whole frame-
work of the body. Nay, more, in the earliest condi-
tion of the human organism, in that state in which
it has just become distinguishable from the egg in
which it arises, it is nothing but an aggregation of
such corpuscles, and every organ of the body was,
once, no more than such an aggregation. Thus a
nucleated mass of protoplasm turns out to be what may
be termed the structural unit of the human body. Asa
matter of fact, the body, in its earliest state, is a
mere multiple of such units ; and, in its perfect con-
dition, it is a multiple of such units, variously modi-
fied.” The formula which expresses the essential
structural character of the highest animal, very
nearly covers all the rest, as the statement of its
powers and faculties covered that of all others.
“ Beast and fowl, reptile and fish, mollusk, worm,
and polype, are all composed of structural units of
the same character, namely, masses of protoplasm
with a nucleus. There are sundry very low animals,
each of which, structurally, is a mere colorless blood-
corpuscle, leading an independent life. But, at the
very bottom of the animal scale, even this simplicity
becomes simplified, and all the phenomena of life are
manifested by a particle of protoplasm without a nucleus.

“What has been said of the animal world is no less
true of plants. Imbedded in the protoplasm at the
broad, or attached end of the nettle hair, there lies
a spheroidal nucleus. Careful examination further

THE CELL DOCTRINE. 107

proves that the whole substance of the nettle is made
up of a repetition of such masses of nucleated proto-
plasm, each contained in a wooden case, which is
modified in form, sometimes into a woody fibre,
sometimes into a duct or spiral vessel, sometimes
into a pollen grain, or an ovule. Traced back to its
earliest state, the nettle arises as the man does, in a
particle of nucleated protoplasm. And in the lowest
plants, as in the lowest animals, a single mass of such
protoplasm may constitute the whole plant, or the
protoplasm may exist without a nucleus. Under
these circumstances it may well be asked, how is one
mass of non-nucleated protoplasm to be distinguished
from another? why call one ‘plant,’ and the other
‘animal?’ The only reply is that, so far as form is
concerned, plants and animals are not separable, and
that, in many cases, it is a mere matter of conven-
tion whether we call a given organism an animal or
a plant.”

The researches of the chemist have also shown a
like uniformity of chemical composition in “ proto-
plasm ” or living matter, proving that whatever its
source, it contains carbon, hydrogen, oxygen, and
nitrogen, producing in their combination a complex
substance, which in our ignorance of its more exact
nature, we call proteinaceous or albuminoid matter.

Further, the matter of life is composed of ordi-
nary matter, and again resolved into ordinary matter
when its work is done. Waste is constantly going
on, which must be supplied by food, which is con-
verted into protoplasm. A solution of smelling salts
in water, with an infinitesimal proportion of some

108 THE CELL DOCTRINE.

other saline matters, contains all the elementary
bodies which enter into protoplasm, yet an animal
cannot make protoplasm out of these. And this is
characteristic. It must take it ready made from
some other animal or some plant, the animal’s highest
feat of constructive chemistry being to convert. dead
protoplasm into the living matter of life, which is
appropriate to itself. Therefore, in seeking for the
origin of protoplasm, we must eventually turn to the
vegetable world. The plant, however, takes carbonic
acid, water, and ammonia,'and converts it to the
same stage of living protoplasm with itself, though
some of the fungi need higher compounds to start
with; and no plant can live on the uncompounded
elements of protoplasm, and the absence of any one
of the elements renders the plant unable to manufac-
ture protoplasm. These elements, carbon, hydrogen,
oxygen, and nitrogen, are related to the protoplasm
of the plant as the protoplasm of the plant to the
animal.

Thus far it is plain that the views of Prof. Huxley
accord with those of many eminent histologists and
physiologists, the result of whose observations have
been embodied in these pages, and his descriptions
will be accepted as undoubtedly accurate. More
widely, in common with the school of so-called
“physicists,” of which he is one, does he differ in
his views as to the phenomena exhibited by protoplasm.

According to Huxley, protoplasm once produced, all
the phenomena exhibited by it are simply its properties,
just as the phenomena exhibited by water in its various
states are properties. They do not take place through

THE CELL DOCTRINE. 109

the guidance of any principle called “ vitality,” any
more than the phenomena of water take place by
virtue of “aquosity.” Prof. Huxley can discover no
halting-place between the admission that protoplasm
of one animal or vegetable is essentially identical
with and readily converted into another, and the
further concession that all vitul action may, with
equal propriety, be said to be the result of the molecu-
lar forces of the protoplasm which displays it. The
thoughts to which we give utterance are the expression of
molecular changes in protoplasm. These are admit-
-tedly so-called materialistic terms. Yet Prof. Huxley
says: ‘« Nevertheless, two things are certain: the one,
that I hold the statement (above) to be substantially
correct ; the other, that I, individually, am no material-
ist, but on the contrary believe materialism to involve
grave philosophical errors.” Such union of materialis-
tic terminology with the repudiation of materialistic
philosophy, he believes to be “not only consistent
with, but necessitated by sound logic.” This he pro-
ceeds to show in this manner: If it be supposed that
knowledge is absolute, that we know more of cause
and effect than a certain definite order of succession
of facts, and that we have a knowledge of the ne-
cessity of that succession, then there is no escape
from utter materialism and necessarianism. But it
is impossible to prove that anything whatever may
not be the effect of a material: and necessary cause,
and no act is really spontaneous, since a really spon-
taneous act is one which has no cause. Yet any one
familiar with the history of science will admit that
its object has always meant, and means the exten-,
10

110 THE CELL DOCTRINE.

sion of the province of matter and causation, and the
concomitant gradual banishment from all regions of
human thought, of what we call spirit and sponta-
néity,—that is, the object of all science has been and
is to find out the causes of all phenomena; and there
is no difference between the conception of life as the
product of a certain disposition of material molecules,
and the old notion of an Archeeus governing and di-
recting blind matter within each living body, except
that here, as elsewhere, matter and law have devoured
spirit and spontaneity. And moreover, the physiology
of the future will gradually so extend the realm of
matter and law, until it is coextensive with knowl-
edge, with feeling, and with action. It is this prog-
ress of knowledge, according to Prof. Huxley, which
so many of the best minds conceive to be the progress
of materialism, which they watch with such fear and
powerless anger as a savage feels, when, during an
eclipse, the great shadow creeps over the face of the
sun. We know nothing of this terrible “ matter,”
except as the name for the unknown and hy potheti-
cal cause of states of our own consciousness, and as
little of that “spirit,” except that it is also a name
for an unknown and hypothetical cause of states of
consciousness, that is, matter and spirit are both
names for the imaginary substrata of groups of natu-
ral phenomena. Dire necessity and “iron” law are
gratuitously invented bugbears. If therebean “iron”
law, it is that of gravitation, and if there bea physi-
cal necessity, it is that a stone unsupported will fall
to the ground. We know nothing more of this latter
phenomenon, except that stones always have fallen

THE CELL DOCTRINE. 111

to the ground under these conditions, and that they
will continue to fall to the ground thus unsupported.

It is simply convenient to indicate that,all the con-
ditions of belief in this case have been fulfilled, by
calling the statement that unsupported stones will
fall to the earth a “law of nature.” But when for
will we exchange must, we introduce an idea of neces-
sity which does not lie in the observed facts, and is
not warranted by anything that is discovered else-
where. And with regard to which Prof. Huxley
says: “ For my part, I utterly repudiate and anathe-
matize the intruder. Fact I know,and Law I know ;
but what is this necessity, save an empty shadow of /
my own mind’s throwing? But, if it is certain that
we can have no knowledge of the nature of either
matter or spirit, and that the notion of necessity is
something illegitimately thrust into the perfectly
legitimate couception of law, the materialistic position
that there is nothing in the world but matter, force, and
necessity, ts as utterly devoid of justification as the most
baseless of theological dogmas.

“The fundamental doctrine of materialism, like
those of spiritualism, and most other ‘ isms,’ lie out-
side ‘ the limits of philosophical inquiry,’ and David
Hume’s great service to humanity is his irrefragable
demonstration of what these limits are. Hume
called himself a skeptic, and therefore others cannot
be blamed if they apply the same title to him; but
that does not alter the fact that the name, with its
existing implications, does him gross injustice. If
a man asks me what the politics of the inhabitants
of the moon are, and I reply that I do not know;

’

112 THE CELL DOCTRINE.

that neither I nor any one else have any means of
knowing ; and that, under these circumstances, I de-
cline to trouble myself about the subject at all, I do
not think he has any right to call me a skeptic. On
the contrary, in replying thus, I conceive that I am
simply honest and truthful, and show a proper regard
for the economy of time. So Hume’s strong and
subtle intellect takes up a great many problems
about which we are naturally curious, and shows us
that they are essentially questions of lunar politics,
in their essence incapable of being answered, and
therefore not worth the attention of men who have
work to do in the world.” .

“Tf we find that the ascertainment of the order of
nature is facilitated by using one terminology, or one
set of symbols, rather than another, it is our clear
duty to use the former, and no harm can accrue so
long as we bear in mind that we are dealing merely
with terms and symbols. In itself it is of little mo-
ment whether we express the phenomena of matter

in terms of spirit, or the phenomena of spirit in
terms of matter; matter may be regarded as a form
of thought, thought may be regarded as a property
of matter—each statement has a certain relative
truth. But with a view to the progress of science,
the materialistic terminology is in every way to be
preferred. For it connects thotght with the other
phenomena of the universe, and suggests inquiry
into the nature of those physics conditions, or con-
comitants of thought, which are more or less acces-
sible to us, and a knowledge of which may, in future,
help us to exercise the same kind of control over the

THE CELL DOCTRINE. 113

world of thought as we already possess in respect to
the material world; whereas, the alternative, or
spiritualistic terminology is utterly barren, and leads
to nothing but obscurity and confusion of ideas.
Thus, there can be little doubt that the further
science advances, the more extensively and consist-
ently will all the phenomena of nature be represented
by materialistic formule and symbols. But the man
of science, who, forgetting the limits of philosophical
inquiry, slides from these formule and symbols into
what is commonly understood by materialism, seems
to me to place himself on a level with the mathema-
tician, who should mistake the 2’s and y’s, with
which he works his problems, for real entities, and
with this further disadvantage, as compared with the
mathematician: that the blunders of the latter are of
no practical consequence, while the errors of sys-
tematic materialism may paralyze the energies and
destroy the beauty of a life.”

These are the views of the “ physicists,” so-called,
a school represented. by Prof. Huxley, Prof. Owen,
Herbert Spencer, Mr. Grove, Prof. Tyndall, and
others. Prof. Owen, in the last pages of vol. iii of
The Anatomy of the Vertebrates, declares himself the
champion. of spontaneous generation, and he main-
tains, also, that the formation of living beings out of
inanimate matter by the conversion of physical and
chemical into vital modes of force, is a matter of
daily and hourly occurrence. Mr. Grove says that
“in a voltaic battery and its effects we have the
nearest approach man has made to an experimental
organism,” and that in the human body we have

10*

114 THE CELL DOCTRINE.

chemical action, electricity, magnetism, heat, light,
motion, and possibly other forces “ contributing, in
the most complex manner, to sustain that result of
combined action we call life.”

ADDISON, WALLER, COHNHEIM (1842, 1846, 1867).

At the annual meeting of the Provincial Medical
and Surgical Association of England, held at Exeter,
August 3d and 4th, 1842, William Addison read a
paper entitled “ Experiential and Practical Re-
searches on the Structure and Function of Blood-cor-
puscles, on Inflammation, and on the Origin and Na-
ture of Tuberclesin the Lungs.” In the section “On
Pus-corpuscles,” he says: “‘ The colorless blood-cor-
puscles appear to form pus-corpuscles.” In the section
on “Inflammation,” we find the following: “ The cir-
culation in the web of a frog’s foot was watched at
intervals for half an hour, and only a few lymph-glob-
ules were seen. A crystal of salt was applied, and
the examination continued: its first effect was to
quicken the rate of the circulation ; this soon ceased,
and the blood became stationary, the vessels being
red and congested. In the capillaries contiguous to
the congested vessels, the blood was oscillating to and
fro ; a little further off, the circulation was very quick.
In half an hour the number of lymph-globules had
increased considerably, and the circulation in the
congested vessels was resumed; but the corpuscles
passing through them sometimes oscillated to and
fro, sometimes retrograded, and at others hurried or
darted through the vessels with the utmost velocity.

THE CELL DOCTRINE. 115

The current of the red corpuscles in some of the
veins appeared to be confined to the centre of the
vessel, and not to touch the circumference, which was
occupied by a great many lymph-globules. On the
following morning the whole interior of the inflamed
vessels appeared to be lined with lymph-globules.
By gently altering the focus of the microscope they
were seen below the red current, and many of them
appeared to lie externally to the boundary of the vessels.”
Again, page 258:* “ During some of these experi-
ments the islets of tissue between the capillaries be-
came distinctly cellular, and appeared as if over-
spread with irregular-shaped lymph-globules.”
These various statements, while they permit the
inference that Addison first conceived that the col-
orless blood-corpuscle becomes the pus-corpuscle in
inflammation, do not allow us to infer that he actu-
ally observed its migration through the walls of
vessels. Indeed, he says, page 259: “‘ The phenomena
observed in the foregoing experiments corroborate
the views of those distinguished physiologists who
entertain the opinion that the capillary distribution
of the blood is situated in the channels of the tissue,
and not in vessels with a distinct membranous coat.” +
In 1846 Wallert more correctly appreciated this
migration, but it was not until Cohnheim published
his famous paper on “Inflammation and Suppura-

* The Transactions of the Provincial Medical and Surgical
Association, vol. xi. London, 1848.

+ Miiller’s Physiology, vol. i, p. 229.

{ London, Dublin, and Edinburgh Philosophical Magazine,
vol. xxix, pp. 271 and 398,

116 THE CELL DOCTRINE.

tion,”* in 1867, that the observation attracted the
attention it deserved. Since then it has been con-
firmed again and again, and most conclusively lately
by Arnold, of Heidelberg.t .

The theory which grew out of Cohnheim’s obser-
vation was, that it is the colorless corpuscle of the blood,
rather than the connective tissue corpuscle, which is
the starting-point of all new formations, healthy and
morbid. It is the accumulation of these corpuscles
outside of the bloodvessels, in the interspaces of the
fibrillar connective tissue, which forms the beadlike
rows of cells figured by Virchow and others. Itis these
which, accumulated in larger numbers, forms the pus
of an abscess. It is these which, perverted in the
direction of their development, produce the tissue of
tubercle, cancer, sarcoma, and other forms of morbid
growth. It is these, also, which become the medium
of repair of all injured or destroyed tissues which are
capable of regeneration, whatever their complexity
or simplicity.

That the colorless corpuscle plays a most important
role in the production of new formations, healthy
and morbid, few now deny; but that it is the sole
morphological element, exclusive of the connective
tissue corpuscle, thus active, many will not admit.
Conspicuous among those who have combated this
exclusive view are Stricker and Norris,t whose ob-

* Ueber Entziindung und Hiterung, Virchow’s Archiv, Bd. x1,
1867, p. 1.

+ Ueber Diapedesis, Virchow’s Archiv, vol. 58, 1873.

{ Versuche uber Hornhaut Entziindung, Studien aus dem In-
stitute fiir Experimentelle Pathologie in Wien, aus dem Jahre
1869.

THE CELL DOCTRINE. 117

servations and experiments in the production of
inflammation in the corneee of frogs and rabbits have
attracted much attention. Their object was to show
that the connective tissue corpuscles (the corneal cor-
puscles here) shared in the formation of the products
of inflammation by their proliferation, and that the
colorless corpuscles themselves, after their migration
beyond the walls of the bloodvessels, also underwent
cell division, and thus contributed in a second man-
ner to the formation of pus. They argued, also, that
the enormous accumulations of pus in large abscesses
could not be reasonably accounted for on the ground
that the only source of the pus-corpuscle is the wan-
dered-out colorless corpuscle.

These observations of Stricker and Norris are gen-
erally acknowledged as having settled the question
in favor of the view, that there are two elements of
organization,—the colorless corpuscle and the connec-
tive tissue corpuscle, either of which may also become
the starting-point of pathological new formations.

NEW VIEWS ON THE STRUCTURE OF CELLS AND
NUCLEI.—1877-78.

As early as 1867, C. Frommann* published a pam-
phlet on the “Normal and Pathological Anatomy of
the Spinal Cord,” in which he states that the pres-
ence of fibrils in the nuclei and nucleoli of cells, first
observed in ganglion-cells, was demonstrated by him
also in the cells of connective tissue, cartilage, and

* Untersuchungen iiber die normale und pathologische Anato-
mie des Riickenmarks, 2 Theil. Jena, 4. Mit 6 Taf. p.17. For
abstract of, see Henle and Meissner’s Bericht fiber die Forschritte
der Anatomie und Physiologie, in Jahre 1867, Leipzig, 1868.

118 THE CELL DOCTRINE.

bone-cells, and in the epithelium of the mouth and
capillary vessels. He observed clear shining fibrils pro-
ceeding from the nucleolus joining themselves to others
proceeding from the nucleus and protoplasm. From
the nucleoli of connective tissue arise one, two, and,
more seldom, three fibrils. These sometimes dwindle
away in the nucleus itself; at others, after a straight
or curved course, leave the cell and lose themselves
in the neighborhood. Repeatedly was observed the
entrance of a nucleolus-fibre into a cell process, and
if two cells were united by a process from each, the
nucleolus-fibre was seen to pass from one cell to
another. So also fibres originating in the nucleus,
sometimes as many as six in number, could be fol-
lowed into the protoplasm, and more seldom beyond
the cell. Certain fibres appear to cease in the nu-
‘cleus by a free extremity, as though cut off. Others
have attached to them glistening granules, which in
fresh, as well as hardened preparations, are con-
tained in variable number in the nucleus. Frommann
believes that the granules of the nucleus and proto-
plasm are the nodal points of a very fine fibrous net-
work, from which fibrils go off and leave the cell.
These are found in the cells of all the tissues named
above. In consequence of this complicated structure
of the nucleus, Frommann thinks it improbable that
it multiplies by division; he believes in a free new
formation of the nucleus in the protoplasm, where,
alongside of nuclei of ordinary appearance, smaller
homogeneous ones make their appearance.

In .1873,* Heitzmann asserted that the substance

* Untersuchungen iiber das’ Protoplasma, Sitzungsber., d. k.
Akad, d. Wiss. zu Wien, Bd. Ixvii und Ixviii, Abth. iii, 1873.

*

THE CELL DOCTRINE. 119

of various cells, amcebee, blood-corpuscles, cartilage-
cells, bone-cells, epithelial cells, etc., contain net-
works of minute fibrils, into which pass fibrils radi-
ating from the interior of the nuclei of these cells.

In 1875, Frommann* again described, in accordance
with Heitzmann, a minute network of fibrils in the
nuclei of blood-corpuscles of Astucus fluviatilis, which
passed through the nuclear membrane into a similar
network in the substance of the blood-corpuscles.

Schwalbe,t in 1875, found the nucleoli in the nuclei
of ganglion-cells in the retina often possessed of mi-
nute filamentous prolongations, and the uuclear
membrane showing prominences on its inner surface.
Schwalbe designates as “ nucleolarsubstanz,” the nu-
cleolus and filaments, nuclear membrane and its
prominences, to distinguish it from the rest of nu-
clear matter, which he calls “ kernsaft”’ or “ nuclear
juice.”

Kupffer,t in 1875, maintained that the substance
of the lower cells of the frog, the odontoblasts, the
epithelial cells of the salivary gland of Periplaneta
orientalis, is composed of a hyaline (non-fuid) ground

See also a paper by Heitzmann on the same subject, in the New
York Medical Journal for 1877. For the historical facts from this
date in the development of these new views, I am indebted to the
very valuable paper of Dr. Klein in the Quarterly Journal of Mi-
croscopical Science for July, 1878.

* Frommann, Zur Lehre von der Structur der Zellen, Jenaische
Zeitschrift f. Naturw., Bd. ix, 1875, p. 280.

+ Bemerkungen iiber d. Kerne d. Ganglionzellen, Jenaische
Zeitschrift f. Naturw., Bd. x, 1875, p. 25.

{ Ueber Differenzirung d. Protoplasma an den Zellen thierescher
Gew., Schriften des Naturw. Vereins f. Schleswig-Holstein, Heft
iii; and Beitr. z. Anat. u. Physiol., Festgabe f. Carl Ludwig, 1875.

120 THE CELL DOCTRINE.

substance, “ Paraplasma,” and of a granular fibrillar
contractile “Protoplasma,” imbedded in the former.
The relation and distribution of the protoplasmic
fibrillar substance varies in cells of different kinds.

Strassburger,* in 1876, observed that in developing
cells of Phaseolus multifiorus, a network of fibrils is
present radiating from the nucleolus, and permeating
the interior of the nucleus in connection with a simi-
lar network of the cell substance.

In 1876, Bitschlit observed in the nuclei of colored
blood-corpuscles of the frog and newt, minute fibres
with granular thickenings, but no nucleolus as as-
serted by Ranvier.

Moyzel,t in 1875, found in the epithelium of the
cornea of the frog, rabbit, and cat, during regenera-
tion, large round nuclei, in which he observed filamen-
tous masses, either convoluted or radiating from a
central point. He regarded these forms as due to a
particular stage of division, as described by Biitschli
and Strassburger.

O. Hertwig,§ in 1876, also distinguished a “nuclear
substance,” from a “ nuclear juice,” and Biitschli,|| in
the same year, a “nuclear matter” from a “ nuclear
fluid.” The “nuclear matter” of the latter com-

* Ueber Zellbildung und Zelltheilung, Jena, 1875. Abstract in
Quart. Jour. Mic. Sci. +, vol. xvi, 1876, p. 188.

¢ Studien iiber die ersten Entwickclungserscheinungen der
Eizelle, Abhand]. d. Senkenbergischen Naturf. Gesellsch., Bd. x,
1876.

{ Ueber eigenthumliche Vorgiinge bei der Theilung der Kerne
in Epithelialzellen, Centralbl. f. Medic. Wissensch., No. 50, 1875.

% Beitrage zu einer einheitlichen Auffassung der verschiedenen
Kernformen, Morphol. Jahrbuch, 1876, Bd. 2, H. 1, 1876, p. 73.

|| Loc. citat., vol. x.

THE CELL DOCTRINE. 121

prises the nuclear membrane, the nucleolus, and a
fibrillar stroma, which latter, in some instances, ex-
tends in a radial manner from the nucleolus.

E. Van Beneden* saw a fine protoplasmic reticu-
lum in the large axial entoderm cell of Dicyema,
which (reticulum) exhibited slow spontaneous move-
ments. In the nucleus of the ripe ovum of Astero-
canthion rubens he also observed within the nuclear
membrane and beside the nucleolus a delicate net-
work of finely granular substance, which he calls
“nucleoplasma,” including several “ pseudo-nucleoli.”
According to him the germinal vesicle of the ripe
ovum of the rabbit also contains a minute network.

Arndt,} in 1876, distinguished in the nucleus a
homogeneous ground substance and elementary glob- -
ules; the former possesses a vesicular structure, and
incloses in its meshes the latter.

W. Flemming,t in observations on the structure
of nuclei found in the membrane of the urinary blad-
der of Salamandra maculata, in 1876, saw a very deli-
cate and dense network of fibres uniformly pervading
the interior of the nucleus, and attached to the
nuclear membrane. This network, “ Geriistformige

* La maturation de l’euf, la fécondation et les premiétres phases
du développement embryonaire des Mammiferes. Bul. de l’Acad.
Royale de Belgique, 2 Ser., t. 40, 1875; also Contributions 4 ]’his-
toire de la vesicule germinative et du premier noyau embryonaire,
in the same journal, January, 1876.

+ Uber den Zellkern, Sitzungsb., d. Medicin. Vereins zu Greifs-
wald, Nov., 1876. .

' £ Beobachtungen tiber die Beschaffenheit des Zellkerns, Archiv
fiir Mikrosk. Anat., Bd. xiii, 1876, p. 693, and following.

1

122 THE CELL DOCTRINE.

Structur,” was seen by Flemming in the nucleus of
all cellular elements of the bladder of Salamandra,
epithelial cells, connective tissue cells, migratory
cells, unstriped muscle-cells, nerve-cells, endothelial
cells, and blood-corpuscles. With regard to Flem-
ming’s observations, Klein* says their clearness and
extent leave no doubt that the network in the nu-
cleus represents a definite and pre-existing structure.
This has been to a certain extent questioned by Lang-
hanst as regards the fresh cells of the human decidua
serotina, but, Klein again observes, “ Flemming’s
assertions cannot be in the least shaken, considering
that he observed the above structure, not only after
the use of reagents, e. g., acetic acid, chromate of
potash, alcohol, chromic acid, with or without sub-
sequent staining in carmine or hematoxylin, but also
in the absolutely uninjured bladder, i. e., while this
organ was being observed in the living curarized
animal.” And, in a later note, Flemming} states
that he observed the same network in the nuclei of
various cells, also in the living and perfectly unin-
jured larva of Salamandra.

Eberth§ noticed nuclei containing anastomosing
filaments in their interior in the epithelium of the
cornea and the endothelium of the membrana Des-
cemeti under normal conditions. He regards them

* Loc. citat.

+ Zur Lehre von Zusammensetz. des Kerns, Centralblatt f.
Medic. Wissenschaft, 1876, N. 50.

{ Zur Kenntniss des Zellkerns, in Centralbl. f. Medic. Wiss.,
1877, No. 20.

@ Ueber Kern und Zelltheilung, Virchow’s Archiv, Bd. 67,
1876.

THE CELL DOCTRINE. 123

as peculiar forms in the development and division of
nuclei.

Eimer* found in numerous nuclei that the gran-
ules of the “ granular zone” surrounding his “hya-
loid” are due to protoplasmic filaments, which per-
meate the interior of the nucleus, and anastomose
with each other 80 as to form a network, which ex-
tends to the membrane of the nucleus, and also sends
radiating fibrils through the hyaloid into the nu-
cleus. The network of fibrils of the nucleus is, in
some instances, also in connection with fibrils and
networks of the same belonging to the cell substance
itself.

The most recent contribution on this subject is the
valuable paper by Dr. E. Klein, already alluded to,
and published in the Quarterly Journal of Microscopical
Science, for July, 1878. In this paper Klein shows
by observations} on the ordinary freshly killed newt

* Weitere Nachrichten iiber den Bau des Zellkerns, etc., Archiv
f. Mikrosk. Anat., Bd. xiv, 1877, p. 94. Also, ‘‘ Notes and Mem-
oranda’? of the April number of the Quart. Jour. Microsc. Sci., °
1878.

+ The following is the method pursued by Klein: The stomach
of a freshly killed newt is eut open and placed into a 5 per cent.
solution of chromate of ammonia in a closed vessel, where it is kept.
for about twenty-four hours. It is then washed in water for about
half an hour, and placed ‘after this in a dilute solution of picro-
carmine, where it is left till it assumes a deep pinkish-yellow tint.
It is now washed in water, and microscopic specimens are pre-
pared inthismanner. The mucous surface of the organ is scraped
with a small scalpel, whereby smaller or larger flakes may be easily
removed; they are placed in a very tiny droplet of glycerin on a
glass slide ; by slight knocking with the rounded or flat top of any
thin rod or needle-holder these flakes are broken up into micro-

" ™

124 THE CELL DOCTRINE.

(Triton cristatus), that the statements of Flemming
are correct, and presents in addition several observa-
tions he has made with reference to the cell-sub-
stance itself, and the relation of it to the intranuclear
network.

Dr. Klein examined gland-cells, surface epithelial
cells, endothelium, unstriped muscle-fibres, connec-
tive tissue corpuscles, and nerve-fibres. And every
where the nuclei showed an extremely beautiful net-
work of fibrils. This he designates as the intra-
nuclear network, which is imbedded in a homoge-
neous ground-substance. The network does not in
all instances extend up to the nuclear membrane, but
leaves a narrower or broader zone next to the mem-
brane unoccupied. But in all instances the network
is in connection with the limiting membrane by nu-
merous fibrils. The spaces formed by their anasto-
moses are not uniform, being sometimes larger in the
peripheral parts than in the central, and sometimes
the reverse. All forms are found between a network
with fibrils as in a net, and a honeycomb of mem-
branous structure as in a sponge. In almost all in-
stances are observed a smaller or greater number of

scopic fragments, a drop of glycerin is placed on a covering-glass,
and this is inverted over the above specimen.

Examined under a moderately high power, say Hartnack’s 7 or
8, or Zeiss’s D or E, we recognize easily innumerable isolated cells
or groups of epithelial cells, and a great many isolated nuclei or
fragments of nuclei. If the scalpel has been drawn over the sur-
face of the mucous membrane with a little energy, the preparation
contains great numbers of gland-cells, isolated and in continuous
masses, and also other elements belonging to the tissue of the
mucosa,

THE CELL DOCTRINE. 125

minute bright spots, which are shown by careful
focussing to be fibrils of the network seen in optical
transverse section or at the points of anastomosis.
In some nuclei, according to Klein, the more irregu-
larly shaped dots are due to a thickening of fibrils
from place to place, although Flemming attributes
them altogether to the fermen cause, (transverse sec-
tions of fibrils). Whence it is clear, the more shrunk
the intranuclear network, or the more twisted and
convoluted the fibrils, the more does the nucleus
present the appearance of being granular.

The nucleolus receives an entirely peculiar expla-
nation at the hands of Klein, in which he seems to be
sustained by the observations of Van Beneden, 0. -
Hertwig, Flemming himself, Auerbach, Eimer, and
especially of Strassburger, Schwalbe, and Langhans.
With regard to it Klein first says with undoubted
truth: “ Now, to every experienced student of his-
tology, it must have become apparent that if there
is one thing unsatisfactory, unreliable, puzzling, and
inconstant about the nucleus of vast numbers of
cells, it is this very nucleolus.” Second, that after
a very prolonged examination he has arrived at the
conclusion that these large particles (nucleoli) are
due to one of two things: in some instances they are
distinctly thickenings of the network, in others they
appear to be merely due to the shrivelling up and
intimate fusion of a part of the network. The in-
constancy as regards size, shape, and number of the
so-called nucleoli seem to him to point very strongly
in the above direction.

The assertions which have been made as regards

11*

126 THE CELL DOCTRINE.

spontaneous movements of nucleoli by Auerbach,
Brandt, Eimer, Kidd, and others, he regards as quite
compatible with the above view, for Van Beneden
has observed movements in the intranuclear network,
and it is quite possible that the above assertions
might refer to such movements.

With regard to the nuclear membrane already men-
tioned, Klein says that it is composed of an outer
thicker portion, which is the limiting membrane proper
and closely connected with it, an inner, more or less
incomplete layer, which is a peripheral condensation
of the intranuclear network, with which it is connected
by longer or shorter threads. The clear space some-
times observed between the membrane of the nucleus
and the intranuclear network is due to a retraction
of the latter from the former.

Klein has also demonstrated a network of fibrils
in the substance of cells outside of the nucleus, which
he designates as the intracellular network, in contra-
distinction to intranuclear, in the meshes of which
again is the interfibrillar or ground-substance, which
in the case of the columnar epithelial cells on mucus-
surfaces, known as goblet-cells, is mucus,* but finds
them also in the endothelium of the surface, in un-
striped muscle-fibres, connective tissue corpuscles, and
nerve-fibres.

Klein further traces a direct connection of the fibrils

* They are best seen on the slender goblet-cells of the stomach
of the newt, kept for twenty-four hours in Miiller’s fluid, placed
then for half an hour in a mixture of two parts chromic acid (4
per cent.), and one part methylated alcohol, washed after this in
water, and stained with picro-carmine.

THE CELL DOCTRINE. 127

of the intracellular with those of the intranuclear network,
and along with Eberth, Marchi, and Eimer, traces in
the epithelial cells of the foregut of the newt, the
cilia in direct continuation through the cell-cover with
the fibrils of the intracellular network.

The intracellular network is less conspicuous than
the intranuclear, but possesses the same general
characters.

The importance of this subject has seemed so great
that I have added at the end of the volume the essen-
tial portions ofsthe plate from the Quart. Jour. of
Micros. Sci., for July, 1878, which, with its descrip-
tion, will give a very accurate notion of these new
views as confirmed by Dr. Klein, than whom, of
modern observers, I consider none more reliable.

SUMMARY—PRESENT STATE OF THE CELL DOCTRINE—
AUTHOR’S VIEWS.

Minute analysis of the solids of the organism has
long been an object of the histologist and physiolo-
gist, which, resulting first in the partes similares of
Aristotle and Galen, has finally reached the so-called
“cell” or “elementary part” as the ultimate physi-
cal element of organization, out of which all tissues,
healthy or diseased, are formed. Our ideas as to the
exact physical constitution of this elementary part
have undergone considerable change since the first
announcement by Schleiden and Schwann, in 1838, of
its exact physiological position. The most impor-
tant modifications of the original conception of a cell
is that which removes altogether its vesicular char-

128 THE CELL DOCTRINE.

acter. So that the term “cell” is really no longer a
correct one, since the object to which it is applied is,
in its youngest, most active state, at least, a solid
mass or “clump” of living matter without the ves-
tige of a wall or envelope about it. The word has,
however, become so intimately associated with his-
tology that it is doubtful whether it will ever fall
into disuse, nor does it much matter, so long as cor-
rect notions of the elementary part are obtained.
This latter term, “ elementary part,” is, however, to
be preferred.

Consistently with the latest determined facts, the
cell or elementary part is best defined as the smallest
mass of living matter possessing the essential life prop-
erties of reproduction, nutrition, growth, and develop-
ment. Tosuch substance the terms “sarcode,” “ pro-
toplasm,” “ gerininal matter,” and “ bioplasm ” have
been applied, the first by Dujardin, the second by
Max Schultze and Remak, and the third and fourth
by Beale.

The term “sarcode” has nearly gone out of use,
and “ protoplasm ” has come to be largely used for
that part of the cell outside of the nucleus, without
regard to its living properties. The term “ bioplasm,”
on the other hand, has not yet received such exten-
sion of meaning, while it also etymologically defines
the substance it represents, and is therefore much to
be preferred.

The Nucleus of the Cell—In the interior of most
cells are found one or more differentiated masses of
bioplasm, usually round or nearly so,and more gran-
ular or darker in hue by tranamitted light than the

THE CELL DOCTRINE. 129

remainder of the cell, which are called nuclei. These
nuclei, although strikingly constant, are not invari-
ably present, as pointed out by Briicke,* in 1861, in
the cells of cryptogams, and confirmed by the earlier
discovery in 1854, of a non-nucleated amceba (Ameeba
porrecta) by Max Schultze,t in the Adriatic Sea ;
also later by Heckelt and Cienkowski§ in their dis-
coveries in 1865, by the former of a non-nucleated
protozoon (Protogenes primordialis) in the Mediterra-
nean, and by the latter of two non-nucleated monads
(Monas amyli and Protomonas amyli). Stricker’s||
observations on the fecundated egg of the frog in-
cline him to adopt the view of Briicke, and omit
the nucleus in a theory of elementary organiza-
tion. Such facts as these prove erroneous the defini-
tion of a cell proposed by Leydig and Max Schultze,
“ protoplasm surrounding a nucleus,” and the defini-
tion more recently accepted by Virchow,{ that a cell
is “a nucleus surrounded by a molecular blastema;”
though if we restrict ourselves to the cells concerned
in the organization of the higher animals, the latter
may be anatomically correct, but the former involves
the confusion already referred to in the use of the
word “ protoplasm,” which is here applied to some-

* Briicke, E., Die Elementar-organismen, p. 18-22, 1861.

+ Schultze Max, Organis. d. Polythalam., 1854.

{ Heckel, Zeitschr. f. w. Zoolog., 1865, Bd. xv.

@ Cienkowsky, Max Schultze’s Archiv, 1865.

| Stricker, S., in vol. i, p. 8, Stricker’s Histology, New Syden-
ham Society’s Translation, London, 1870; also p. 504, vol. iii,
London, 1873.

q Letter from Berlin, in Edinburgh Medical Journal, Febru-
ary, 1865.

‘130 THE CELL DOCTRINE.

thing outside of the nucleus to the exclusion of the
latter, to which, if to anything, should be assigned
the term.

For the nucleus possesses pre-eminently three of the
properties of bioplasm named, nutrition, reproduc-
tion, and growth, and in it usually take place the
first steps towards the production of new cells, in,
first, its increase of size through nutrition, and
second, by fission, which subsequently extends to the
remainder of the cell. Indeed the chief function of
the nucleus has been heretofore considered as that of
the reproduction of the cell. But this is by no means
invariably the case, as indeed it is clear should not
be, when we recall the properties of bioplasm or
germinal matter. For although the nucleus is a sep-
arate differentiated portion of bioplasm, it does not
in every cell comprise the whole of the bioplasm of
that cell. This is perhaps best illustrated by ‘the
pus-cell and colorless biood-corpuscle, which are pure
bioplasm, but which still contain within them sepa-
rate centres of germinal matter or nuclei. Consist-
ently with this fact, the pus-cells or white blood-cells
multiply not necessarily through a primary fission
of the nucleus, but often by separating a portion of
the external bioplasm, which becomes an independent
cell ,with the endowments of its predecessor. See
Fig. 7,8, 9,10, of frontispiece. This fact has further
a very important illustration in the fecundated germ
or ovum of sexual generation. It is now universally
admitted that the non-fecundated germ is a distinctly
nucleated cell, and it is almost as generally acknowl-
edged that immediately after fecundation the nu-

THE CELL DOCTRINE. 1381

cleus disappears, to be replaced, however, by a new
one, under favorable conditions, which appears to
take no part in the subsequent cleavage processes.*
Recent descriptions of the nucleus include in its
anatomy a nuclear membrane, which is described by
Auerbach as a somewhat thick, highly refractile,
doubly contoured membrane, which appears to be
less distinctly separated from the cell protoplasm
than from the exterior of the nucleus. This mem-
brane is regarded by Auerbach as having been formed
about the original droplike nucleus by the differ-
entiation of the inmost layer of the protoplasm into
a species of interior cell membrane. According to
Klein, it is composed of an outer thicker portion,
which is the limiting membrane proper, and closely
connected with it an inner more or less incom-
plete layer, which’ is a peripheral condensation: of
the intranuclear network, with which it is connected
by longer or shorter threads. According to Auer-
bach, the nuclei first appear as clear spaces, vacuoles

* According to Prof. Stricker (‘* Development of the Simple
Tissues,’”’ in vol. iii, of ‘Human and Comparative Histology,”
New Syd. Soc. Ed.), precise statement to the effect that the germi-
nal vesicle is persistent and becomes transformed into the nucleus
of the cleavage cells, has only been made by Johann Miiller in the
case of Entoconcha mirabilis (Monatsberichte der Berliner Akade-
mie, September, 1851). But in recording this fact Stricker seems,
as many more recent writers have done, to have entirely overlooked
the much earlier observation of Martin Barry, who distinctly as-
serts (Philosophical Transactions, London, 1840, p. 529), that “ the
germinal vesicle does not burst, or dissolve away, or become flat-
‘tened, on or before the fecundation of the ovum, as hitherto sup-
posed.”

132 THE CELL DOCTRINE.

filled with a tenacious fluid mass possessing no dis-
tinct wall. Each droplet then acquires a membrane
by differentiation of the inmost layer of cell proto-
plasm and nuclei, and intermediary granules after-
wards make their appearance. Once differentiated,
the nuclear membrane is an integral part of the nu-
cleus, constituting the latter a true vesicle, isolable
as a whole by mechanical means.*

The Nucleolus.—Very commonly also, though not
invariably attending it, a second differentiated mass
of matter is found within the nucleus, to which the
term nucleolus is applied. According to Beale, it is
possessed of like endowments, and is supposed to be
the most recently formed bioplasm, the non-vitalized
circulating albumen of the blood being converted
through the agency of pre-existing bioplasm into
the latter substance. If the cell normally develop,
the nucleolus growing becomes the nucleus, the lat-
ter being gradually oxidized upon its exterior and
converted into the cell-contents and cell-wall, while
a younger centre of germinal matter takes the place
of the original nucleolus. This living matter exhibits
the important property of being stained by weak so-
lutions of various coloring matters, as carmine, anilin,
etc., the younger matter taking on the deeper stain,
and by these means its demonstration is rendered
strikingly easy. In rapidly growing cells, a still
younger centre of bioplasm may sometimes be demon-
strated within the nucleolus, while the nucleus re-
maius a distinctly differentiated mass capable of also

* Auerbach, Organologische Studien, Breslau, 1878-4.

THE CELL DOCTRINE. 133

being stained. To this youngest centre of bioplasm,
the term ‘“nucleoleolus” might not be inappropriately
applied.

Klein has announced in his recent paper “ On the
Structure of Cells and Nuclei” (Quart. Journ. Mi-
croscop. Sci., July, 1878), that after very prolonged
examination he has arrived at the conclusion that
these large particles (nucleoli) are due to one of two
things: in some instances they are distinctly thick-
enings of the nuclear network of fibrille demonstra-
ted by Frommann and Heitzmann, and confirmed by
Flemming, Hertwig, E. Van Beneden, Klein himself,
and others, or they are merely due to the shrivelling
up and intimate fusion of a part of this network.

Auerbach supposes that the nucleolus is formed
by the aggregation about a centre, of nucleolar sub-
stance, which is derived either from the periphery of
the nucleus itself or from the inmost layer of cell
protoplasm. According to him, nucleoli may also
multiply by fission. This division is associated with
a movement of the new nucleoli through the nu-
clear ground-substance, the cause of which is not
understood. “The reverse, or the fusion of several
nucleoli into one, also occurs. Auerbach says also
that many facts speak for the identity of nucleolar
’ substance and cell protoplasm. In optical appearance,
nucleoli and the substance of young cells agree, and
vacuolation may occur in either, large nucleoli being
seldom free from clear spaces. Both are amceboid,
the movements in nucleoli having been described by
many observers. Lastly, nucleoli have, as Auerbach
himself shows, that characteristic of vital protoplasm,

12

134 THE CELL DOCTRINE.

the power of multiplication by division. That is,
nucleoli have all the capabilities of elementary or-
ganisms, and are in truth cytodes. Regarded in
this light, they are real daughter cells, which have
arisen by an endogenous process, and the nucleus is
the chamber in which they develop. It is now
merely necessary for them to find a way out through
the body of the mother cell, in order to begin life as
independent beings. That this method of increase
has become obsolete in the cases of a majority of cells
of higher adult animals, or perhaps only occurs in
pathological processes, may be quite true, according
to Auerbach, without offering any obstacle in the
way of such a hypothesis. In the specialization of
function which appears as we ascend the animal
series new means of gaining the same end present
themselves, while the old vanish or are made use of for
other purposes. The phases in the act of propaga-
tion of a unicellular organism may be less significant,
and have a far different outcome in a unit cell of a
more complex creature.*

The fact of the total or partial disappearance of
the nucleus or germinal vesicle shortly before or im-
mediately after fecundation, has been amply con-
firmed by recent observers, although there are some
slight differences in the exact mode of its acceptance.
The most important recent contributions on the sub-

* Auerbach, Organologische Studien, Breslau, 1873-4, or see
abstract of Auerbach’s paper, by Priestley, in vol. xvi, Quart.
Jour. Microsc. Sci., 1876, whence I have obtained the above views
of Auerbach with regard to nuclei and nucleoli.

THE CELL DOCTRINE. 135

ject have been by Auerbach,* Strassburger,t+ Hert-
wig,t and Van Beneden,§ to whom may be added
Biitschli,|| Oellacher,{ Kleinenberg,** and Balfour.tt+

‘Under the influence of the fertilizing act the ovum
again becomes nucleated, the nucleus arising in a
fusion of pronuclear bodies, one of which may pos-
sibly be derived from the fertilizing element (sper-
matozoid). The nucleus then begins series of changes,
about which most observers are agreed, but for the
details of which the reader is referred to the sources
named in the footnote.

* Organologische Studien. Breslau, 1878-4.

+ Ueber Zellbildung und Zelltheilung, Jena, 1875.

t Beitrage zur Kenntniss der Bildung, Befruchtung, und Thei-
lung des thiereschen Hies, Morpholog. Jahrbuch, 1, 1875.

@ La Maturation de l’euf, la Fecondation et les Premiéres
Phases du Développement Embryonaire des Mammiferes, d’aprés
des Recherches faites chez le Lapin. Bruxelles, 1875.

Excellent abstracts of these four papers, by John Priestly, are
found in the Quart. Jour. of Microsc. Sci., for July, 1878, vol.
xvi, new series. In the same number will be found an original
paper by Van Beneden, entitled ‘‘ Contributions to the History of
the Germinal Vesicle, and of the first Embryonic Nucleus,” in
which he compares the views of Hertwig with his own.

|| Vorlaufige Mittheilung iiber Untersuchungen betreffend die
Ersten Entwickelungsvorginge in Befruchteten Ei von Nemato-
den und Schnecken, Zeitsch. fiir Wiss. Zoologie, Bd. xxv; and
“ Vorliufige Mittheilung Einiger Resultate von Studien iiber die
Conjugation der Infusorien und die Zelltheilung, Zeitsch. fiir
Wiss. Zoologie, Bd. xxv, 1875.

{ Beitrige zur Geschichte des Keimblaschens im Wirbelthierei,
Archiv fiir Mikrosk. Anat., Bd. viii, 1872.

** On the Anatomy and Development of the Fresh-water Hydra.
Leipzig, 1872.

+t Balfour, Developmental History of Hlasmobranch Fishes, in
the Journal of. Anatomy and Physiology, January, 1876.

1386 THE CELL DOCTRINE.

As already stated, previous to the researches of
Miller alluded to, no one doubted the disappearance
of the germinal vesicle after fecundation. This was
thought to have been established by Purkinje* and
Von Baer.t But immediately after Miiller made his
announcement, others followed in its confirmation.
Among these were Leydig,t Gegenbauer,§ Fol,|| Leuc-
kart,J Pagenstecher,** Mecznikow,}t Heckel,{{Kol-
liker,§§ and Van Beneden,||| as the result of his earlier
researches. Still later, however, of the above-named

* Purkinje, Symbole ad ovi avium historiam ante incuba-
tionem, 1825.

t Baer, C. E. v., Untersuch. u. die Entwicklungsgesch. der
Fische, 1835; and Untersuch. u. die Entwicklungsgesch. der
Thiere, Bd. ii, Konigsberg, 1828.

{ Leydig, Ueber den Bau und die Systematische Stellung der
Roderthiere, Zeit. fiir Wiss. Zool., Bd. vi, p. 102, 1856.

@ Gegenbauer, Beitrige zur Naheren Kenntniss der Siphono-
phoren, Zeit. fiir Wiss. Zool., Bd. v; Zur Lehre vom Genera-
tionswechsel bei Medusen und Polypen, p. 24; Untersuch. iiber
Pteropoden und Heteropoden, Leipzig, 1855. Ueber die Ent-
wickelung der Sagitta, Halle, 1856.

|| Fol, Die erste Entwickelung des Geryonideneies, Jenaischi
Zeitsch., Bd. vii, 1878, p. 474.

{ Leuckart, Die Menschlichen Parasiten, Bd. ii, 2te Lief. p.
822, 1868.

** Pagenstecher, Die Wochenen, Leipzig, 1865.

tt Mecznikow, Embryologische Studien an Insecten Zeitsch.
fiir Wiss. Zovlogie, Bd. xvi, p. 484, 1866.

t¢ Heckel, Zur Entwickelungsgeschichte der Siphonophoren,
Utrecht, 1869.

@@ Kolliker, Die Schwinimpolypen von Messina, Leipzig, 1853.

||| Van Beneden, Recherches sur la Composition et la Significa-
tion de lceuf basées sur 1’Etude de son mode de Formation et des
Premiéres Phénoménes Embryonaires, Mem. Couronné de ]'Acad.
Roy. de Belgique, 1. xxxiv.

THE CELL DOCTRINE. 137

more recent observers, Auerbach, Strassburger,
Oellacher, Kleinenberg, Biitschli, Balfour, and Van
Beneden, in accordance with fresh researches made
on account of the doubts which those of Oellacher
and Kleinenberg had created in his mind, all agreed
that the germinal vesicle disappeared entirely either
before or during fertilization. Hertwig and Van
Beneden both furnish a carefully detailed account of
the steps in the destruction of the germinal vesicle,
the former in the unripe ova of Toxopneustes lividus,
and the latter in the ovum of the rabbit. While
agreeing in many particulars, they differ in their ac-
count of the final dissolution. According to Hert-
wig, the whole structure, after having become periph-
eral, disappears, leaving behind, of its contents, the
germinal spot, or nucleolus, which becomes the nucleus
of the ovurn ready for the changes of fertilization.
Van Beneden, on the other hand, believes that all the
constituents of the germinal spot disappear in toto.
It seems also that the view of Hertwig, that a part
of the germinal vesicle remains to form the nucleus
of the ovum, was previously suggested by Derbés and
Von Baer. The former, in 1847,* described the
ovarian ova as consisting of three zones, the germi-
nal spot, the germinal vesicle, and the yolk, of which
the middle one only disappears. Von Baerf{ also
states that in the case of echinoderms, the germinal

* Derbés, Observ. sur le Mecanisme et le Phén. qui accomp. la
Formation de l’Embryon chez l’Oursin Comestible, Ann. des Sc.
Nat., Zoologie, 1847, vol. viii.

+ CO. E. v. Baer, Neue Untersuch., iiber die Entwickel. der
Thiere, Froriep’s Neue Notizen, vol. xxxix, 1846.

12*

138 THE CELL DOCTRINE.

spot remains as a nucleus of the ovum when the ger-
minal vesicle is no longer seen.

Similar statements were made by Leydig,* and
Bischoff,} the former with regard to Piscicola, and
the latter in the case of mammals.

The Cell-contents and Cell-wal!.—As cells grow older
and are farther removed from the bloodvessels which
nourish them, there is found to be, exterior to the
nucleus, a portion which no longer admits of staining
by carmine solutions of the strength which will tinge
the nucleus. Further, the extreme periphery of this
is often found condensed so as to present under the
microscope a double contour, and to form an actual
limitary membrane to the cell. To the former the
name “ cell-contents ” has come to be applied, and to
the latter, “ cell-wall.”

With regard to the “ cell-contents,” it is plain that
a strict adherence to the term would require to be
included the nucleus and nucleolus when present,
and such was the original scope of the term when a
cell was defined as a “ closed vesicle or bag with cer-
tain ‘contents,’ among which is essentially a nu-
cleus,” a definition now very properly rejected. It
is to this portion of the cell also, that the German
histologists of the present day apply the term “ proto-
plasm,” instead of to the nucleus and nucleolus, which
in many cells, at least, alone possess the endowments
suggested by the word protoplasm (zpoto-, first, tAacpza,

* Leydig, Zur Anatomie von Piscicola Geometrica, Zeitsch. f.
Wiss. Zoologie, vol. i, 1849.
¢ Bischoff, Entwickelungsgeschichte des Kaninchen eies, 1842.

THE CELL DOCTRINE. 1389

a thing formed). It is for this reason that I would
prefer to reject the term altogether, but it has come
to be so closely associated with this portion of the
cell, that such omission would now seem impossible.

Itis at least probable that this “ protoplasm ” or cell-
contents, the portion of the cell exterior to the nu-
cleus, is derived from the latter by a change in its
structure and composition, a change which in the
absence of more accurate knowledge we may char-
acterize as of the nature of oxidation. But what-
ever this change is, its effect is to alter entirely the
properties of the matter in which it has taken place.
So that it no longer possesses the power of growth
through active efforts of nutrition, that is, by con-
verting the pabulum of the blood into material like
itself, but continues to grow at the expense of the
nucleus or bioplasm, which is gradually converted
into it. It is this to which Beale has applied the
name of “dead” or “formed” material, but for
which I prefer the term non-germinal, since this ac-
curately marks the property which it has lost, and
it is not always lifeless in any other sénse, because
in it still reside properties which are inconsistent
with the state known as death. Thus the function
of the tissue of which the cell forms a part commonly
resides in the cell-contents or non-germinal matter,
as that of “contractility” in muscle and “ neu-
ridity ” in nervous tissues. In some situations it is
truly dead, as where it becomes the secretion of glands,
as milk and bile.

It has already been stated that the “ cell-wall”
when present is simply condensed periphery of cell-

140 THE CELL DOCTRINE.

contents, and is therefore a part of the formed or
non germinal matter, and is in no way essential to
the constitution of the cell*

In structure the cell-wall when present is usually
homogeneous, transparent, in a word structureless.
Recent observations have, however, shown this to be
not invariable. Thus,it is not difficult to demon-
strate with the higher powers certain lines or striz
in the outer thickened edge of the columnar cells
capping the villi of the small intestine, which are
perhaps correctly interpreted by Funke, Kélliker,
aud others, as porous canals in the cell-wall, through
which the fine particles of the emulsified fats are
absorbed. These may at least be discovered occupy-
ing corresponding situations with great distinctness.
Schroen has also described similar markings in the
thickened cell-wall of the rete Malpighii of the
human skin.

Intercellular Substance.—Closely allied to the cell-
wall is the so-called intercellular substance of tissues.
Although it is true that all tissues originate from
cells, yet there are comparatively few which in their
perfect state are composed purely of cells, but the
latter are more or less separated by a substance be-
tween them. Thus cartilage or “gristle” is com-
posed of cells and an intercellular substance, which
is either hyaline or fibrous, and all the connective
tissues, under which are included bone, cartilage,
already alluded to, white fibrous tissue, yellow elastic
tissue, and even teeth, are similarly composed of
cellular elements, and a something between them,
which is either homogeneous or structured.

THE CELL DOCTRINE. 141

The mode of origin of this intercellular substance
has given rise to considerable discussion. Schwann
assumed that there originally existed spindle-cells, the
caudate corpuscles, and that out of these cells, fas-
ciculi of connective tissue were developed by split-
ting up of the body of the cell.

Fie, 18.

A

“tg
al al \

A, Bundle of common, wavy, connective ‘tissue (intercellular substance) split-
ting at its end into fine fibrils. B, Diagram of the development of connective
tissue, according to Schwann. a, Spindle-shaped cell (caudate corpuscle, fibro-
plastic corpuscle of Lebert), with nucleus and uucleolus. d, Cleavage of the body
of the cell into fibrils. C, Diagram of the development of connective tissue,
according to Henle. a, Hyaline matrix (blastema) with nucleolated nuclei regu-
larly distributed through it. b, Fibrillation of blastema (direct formation of
fibrils), and transformation of the nuclei into nucleus-fibres.

Henle thought that originally there were no cells,
but nuclei only were developed in the blastema at
certain intervals, while the fibres which afterwards
appeared were produced by a direct fibrillation of
the blastema, and that while the intermediate sub-
stance was thus being differentiated into fibres, the
nuclei gradually became elongated, so as at length to

142 THE CELL DOCTRINE.

run into one another, and thus give rise to peculiar
longitudinal fibres (nucleus-fibres or kernfasern).

Reichert contended that there were cells in great
abundance between which was deposited intercellular
substance, with which the membrane of the cells be-
came subsequently blended, reaching thus a stage in
which there was no longer any boundary between
the cells and the intermediate substance. The nuclei
he thought also disappeared in some instances. ‘The
intercellular fibres he said were a false interpretation
of an optical image.

Virchow, with Schwann and against Henle, be-

Fie, 19.

a

Diagram of the development of connective tissue, according to Virchow's in-
vestigations. A, Earliest stage, hyaline basis (intercellular) substance, with
largish cells (connective tissue corpuscles) ; the latter drawn upin rows at regular
intervals; at first, separated, spindle-shaped and simple; at a later period, anas-
tomosing and branched. B, More advanced stage; at a, the basis-substance which
has become striated (fibrillated) presents a fasciculated appearance on account of
the cells imbedded in it in rows, the cells becoming narrower and smaller; at 5,
the striation of the basis-substance has disappeared under the influence of acetic
acid, and the fine and long-anastomosing fibre-cells (connective tissue corpuscles),
still retaining their nuclei, are seen.

—_—

lieves that spindle-shaped cells indisputably exist,
and with Henle and Reichert and against Schwann,

THE CELL DOCTRINE. 1438

a splitting up of the cells into fibres does not occur,
but that a previously homogeneous intercellular sub-
stance becomes fibrillated, and that the cells them-
selves preserve their integrity. In young connective
tissue, cells undoubtedly exist, with at least their
more important parts as nuclei and protoplasm, but
in fully formed fibrous tissue the only portion of the
cell which I have been able to satisfy myself, in an
almost daily study of healthy and diseased struc-
tures, is constantly present, is the nucleus.

Max Schultze, in Germany, Beale, in England, and
Leidy, in this country, believe the intercellular sub-
stance to originate as the cell-contents and cell-wall
by a conversion of the nucleus or bioplasm at its
periphery, and a pushing off of this converted
matter by the deposition of new bioplasrn within the
nucleus. In fact, according to them, the intercellu-
lar substance is simply the oldest cell-wall or formed
material, under which term Beale consistently in-
cludes all intercellular substance. See Figs. 11, 12,
frontispiece.

This view seems to me also to be most consistent
with observation, and until something more reason-
able is offered I shall adopt it. One of the most
potent arguments in its favor is the fact that the
cell-wall and intercellular substance are often so
closely contintous that they cannot be separated,
either visually or by dissection. On this account
Beale has also described a “cell” or “elementary
part” of cartilage and similar structures, as includ-
ing in its formed material the intercellular substance
extending to a point midway between it and its
neighbor.

144 THE CELL DOCTRINE.

The Structure of Cells.—As to structure, cells have
been heretofore described as structureless, or vari-
ously granular, and with powers not exceeding 250
diameters, they may still be so described. Few cells
even with these powers are structureless, while on
the other hand, a highly granular condition of a cell
is considered indicative of some pathological change,
except in the case of nerve-cells, which in many
situations are admittedly highly granular in the
normal state. The same may be said of the nucleus.
Occasionally, however, even previous to 1867, certain
nerve-cells were described as striated in appearance.
Examples of structureless cells are most striking in
the unicellular organism, as the ameba (Fig. 16, front-
ispiece), in which we have a structureless nucleated
mass, which owes whatever of structure it possesses
to foreign particles, which it takes up as food. So far
as the structure of the elementary part of the more
complex organisms, however, is concerned, the recent
discoveries, already described (see p. 117) as discernible
by a power as low as 300 diameters, demand a total
change in the description of the structure of cells.
Henceforward we must describe not only the nucleus
but also the cellular substance (protoplast) as fibrillar
in structure, made up of a network of delicate fibres
the meshes of which are filled with an “ interfibril-
lar ” or “ ground substance,” which is structureless,
and that the fibrille of the intracellular and intra-
nuclear networks ate continuous. And if Klein be
correct we must define the nucleoli as merely local
thickenings, natural or artificial, of the intranuclear
network. The intranuclear networks are much more

THE CELL DOCTRINE, 145

distinct than the intracellular. See plate at end of
volume.

The Shape of Cells.—The shape of cells is extremely
varied. In its fully developed state each tissue is com-
posed of cells which may be said to be characteristic of
it. First, the typical form of a cell may be said to be
spherical, and this is the shape of all young cells in
whatever situation found. See Fig. 10, frontispiece.
Second, in glandular tissue, where perhaps cells are
changing more rapidly than in any other situation
in health, their deviation from the round shape is
only such as results from mutual compression, forming
more or less polygonal cells, as seen in Fig. 3, p. 39.
Third, in epithelial structures we have upon the ex-
treme periphery either flat scales containing a small
proportion of bioplasm (nucleus) in proportion to the
. non-germinal matter (cell-contents), and of irregular
outline, forming the so-called squamous epithelium,
Figs. 3, 4, 5, 6, frontispiece ; or fourth, similar scales,
of regular outline, many-sided and with their edges
so adapted as to form a pavement-like structure,
forming tessellated epithelium ; or ji/th, elongated nu-
cleated cells, which from their shape are called colum-
nar cells. Sixth, in certain situations, as the respira-
tory passages, these columnar cells are further provid-
ed with hairlike prolongations, known as cilia, which
in health exhibit a constant waving motion, whence
the cells are called ciliated cells. Wherever columnar
cells are present in successive layers (as is usually the
case), they lose their characteristic shape and ap-
proach more and more the spherical outline as we
recede from the surface, until the deepest and there-

18

146 THE CELL DOCTRINE.

fore youngest cells are again spherical. Seventh, in
nervous tissues again, we meet cells which from their
prolongations, which may be one or many, are called
polar cells, and unipolar, bipolar, or multipolar, ac-
cording to the number of processes they possess.
Highth, in the so-called connective tissue again, we
have a variety in the shape of the cells. Thus in
areolar tissue or connective tissue proper, we have
young round cells composed almost wholly of germi-
nal matter, exhibiting amceboid movements and all
the characters of the leucocyte or colorless corpuscle
(Fig. 10, frontispiece), as well as the elongated spin-
dle shaped nucleated cell, 80 characteristic as to have
long ago received the name connective tissue corpuscle
(Fig. 16, frontispiece, Fig. 18, B.). These latter cells
also possess prolongations, which unite with those
extending from adjacent cells, and being hollow thus
form a canalicular system, of exceeding fineness,
which is believed to be capable of conveying nutrient
juices in the absence of bloodvessels of sufficient size
to conduct the corpuscles of the blood. The appear-
ance of this system is shown in Figs. 16 and 19, from
Virchow’s Cellular Pathology. Ninth, the cells of
striated muscular tissue exhibit a prolonged oval shape,
sometimes resembling that of the connective tissue
corpuscle, while the cells of unstriped muscular tissue
are typical spindle-cells, with bellied centres and
staff-shaped nuclei.

Again, in cartilage we have cells exhibiting various
modifications of the spherical shape, while the cells
of bone give us a tenth form. These are contained
in correspondingly irregular cavities in its substance,

THE CELL DOCTRINE. 147

and are almost fantastic in their irregularity, ex-
hibiting also prolongations which unite with those
of neighboring bone-cells, and with the Haversian
canals which conduct the bloodvessels through bone.
The shape of the cells through whose agency the
teeth are formed is also seen to be peculiar, those
whence the dentine is developed being provided with
a single long process, giving an eleventh form of cell.
Those producing the enamel are columnar, while the
crusta petrosa, or portion of bone-like substance cover-
ing the fang, contains cells similar to those of bone.

Finally, the cells of adipose or fatty tissue require
allusion. They are spherical or compressed vesicles
or sacs of considerable size, are filled with oil, and
exhibit in consequence a brilliant, highly refracting
character, under the microscope, indicated by a broad
dark border and a transparent centre. In the begin-
ning they are in no way different from other young
cells, but, according to Beale, their bioplasm, instead
of being converted into the ordinary non-germinal
matter of albuminous composition, undergoes a fatty
conversion, in the course of which it gradually di-
minishes in size and is thrust towards the wall of
the cell, where it may sometimes be demonstrated
by staining as a small flattened nucleus. The oil
thus produced is termed by Beale secondary formed
material, to distinguish it from the albuminous non-
germinal matter. The same term is applied by him
to the secondary product, starch, found in the shape
of concentrically laminated granules in many vegeta-
ble cells. According to most other histologists, how-
ever, the fatty contents of the fat-vesicles are an infil-

148 THE CELL DOCTRINE.

tration of fat from the blood, superadded to the pro-
toplasm or cell-contents, pushing the latter with the
nucleus to one side, without substituting it or being
derived directly from it as is alleged by Beale.

In pathological formations all the different forms of
cells here alluded to are met with, and there is now
no special type of cell which is known by its shape
to have a pathological impression. It is rather by
the rapidity of growth of cells, their arrangement
and relation to the intercellular substance, as well as
peculiarities in the latter substance itself, that we
know a structure to be a pathological formation.
The “ cancer-cell,” which was so long an object of
wonder and fear, and eagerly sought for as such, is
no longer acknowledged to be anythiny peculiar as
to form. At the same time, when cells from a sus-
pected growth are observed to be very large, to con-
tain numerous nuclei or centres of bioplasm, and to.
exhibit great variety in shape, we have evidences of
that rapidity of growth which is more or less char-
acteristic of malignant formations.

The Size of Cells.—This is likewise extremely vari-
ous. They may be particles of such extreme minute-
ness as to be recognized as mere dots by the highest
powers of the microscope, the smallest particles of
germinal matter measured by Dr. Beale being less
than the ; 57555 of an inch (.000024 mm.) in diame-
ter, while the largest epithelial scales are 51, of an
inch (.1 mm.) in diameter, and the cells of morbid
growths often reach the ,4, of an inch (.127 mm.).
The human ovum, which may be regarded as a typi-
cal cell, with nucleus, nucleolus, and contents, varies

THE CELL DOCTRINE. 149

from the ,1, to ;}y of an inch (.105 mm. to .21
mm.). The smallest particles of germinal matter
above alluded to would not be called cells in the or-
dinary sense of the word, but they are such consist-
ently with our definition, and it must moreover be
remembered that the term is ouly applied to such
particles of matter endowed with the life properties
of reproduction, nutrition, growth, and development.
No particle of oil, inorganic or other matter, which
does not possess these properties can be characterized
as a living cell, whatever its size.

Pure germinal matter is rarely seen in masses as
large as the 5}, of an inch (.05 mm.), since it usually
breaks up into smaller masses to form independent
cells before it reaches this size. As constituting the
nuclei of fully formed cells, it is usually from the
svov tO spy Of an inch (004. mm. to 008. mm.) in
diameter. While nuclei exhibit less variation in size
than do the cells, they are also more constant in shape,
being generally round or oval. Sometimes, however,
they also exhibit a stellate shape.

The Origin of Cells.—It is to be regretted that this
roost important question as to the exact origin of
cells is not yet definitely settled. A very short time
ago the proposition omnis cellula e cellula wag thought
to have been abundantly proven, and that all living
things come from previously existing things, was
generally admitted. Butalthough this view has been,
in the conception of some, shaken by more recent ex-
periment, it still remains the only law of cell forma-
tion in the minds of most physiologists, and I be-
lieve there are none who deny that it forms in all

13*

150 THE CELL DOCTRINE.

the higher forms of animal and vegetable life the
only method of cell genesis which is constantly before
us, while the phenomena of spontaneous generation
are confined to the creation of the lowest vegetable
and animal organisms, so that the proposition omnis
cellula e cellul@ may be now considered as generally
accepted. Prof. H. Charlton Bastian remains the
most eminent English supporter of the doctrine of
spontaneous generation, against whonr Huxley, Rob-
erts, Tyndall, Beale and others have directed arti-
cles and experiments.

In the division of cells to produce young cells, it
is usually in the nucleus that the segmentation first
begins, extending thence to the protoplasm outside of
it, although this is not invariable. For any portion
of the bioplasm of a cell may grow, separate and form
a young cell. It is the bioplasm or germinal matter
alone, however, which can give rise to the cell, and
never the non-germinal matter or formed material.
It is seldom, also, that we see in the animal cell, ex-
cept in the segmentation of the fecundated ovum,
that symmetrical division of the nucleus into two,
these into four, these into eight, and so on, as is con-
stantly seen in the vegetable cell, but it is rather a
budding,and subsequent dropping off of portions of
bioplasm which become young cells, and which al-
most always assume the spherical form when allowed
to float freely (see Fig. 10, of frontispiece).

Strasfbunger* in his recent work on Cell Formation
and Cell Division, admits three methods of increase

* Ueber Zellbildung und Zelltheilung, Jena, 1875.

THE CELL DOCTRINE. 151

of cells,—cell division, free cell formation, and re-
newal (vollzellbildung) of-cells. He says that in the
animal kingdom cell division seems to be the only
authenticated mode of increase of cells. Of methods.
of free cell formation, only a few instances are re-
corded in vegetables. The last method, renewal or
rejuvenescence, has reference rather to the complete
formation or development of cells than to their
origin.

Nutrition of Cells.—In the nutrition of the cell, the
pabulum comes to it from the periphery, being
strained through the formed material. The new
germinal matter takes its place in or near the centre
of the original mass, constituting, according to Beale,
a new centre of germinal matter, which may be the
nucleus, if no other circumscribed centre be present,
or the nucleolus, if it be deposited within such a centre.
Other new centres, according to him, may again take
position within these, and assume the relation of nu-
cleolus to the original nucleolus, which now becomes
the nucleus, an older centre of germinal matter ;
while the original nucleus has probably been con-
verted into the second constituent of the cell, the
formed material. If the nucleolus be what Klein con-
siders it, a mere thickening of the intranuclear net-
work referred to, this order of succession in the for-
mation of differentiated centres of germinal matter
is of course impossible, and it can only be said that
the new matter takes its position towards the centre
of the cell mass.

Ameboid Movement (Diapedesis).—Germinal mat-
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152 THE CELL DOCTRINE.

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ce

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te

INDEX.

Addison, 114

Ameeba, 144

Ameeboid movement, 151
Aristotle, 13

Arndt, 121

Arnold, 26

Auerbach, 133

Author’s views, 127

B.

Baumgartner, 26
Barry, Martin, 48
Beale, 83
Beneden, Van, 121
Bennett, 61
Bergmann, 44
Bibliography, 152
Bioplasm or Germinal Matter, 84
Borellus, 16
Bowman, 64
Bricke, 81
Biitschli, 120

s C.

Cell, Leydig’s definition of, 80
M. Schultze’s “ ‘“ 80
Virchow’s nee 5
author’s 8 “198

Cell doctrine, present state of, 127
formation, 150
division, 150
nutrition of, 151
contents, 138
wall, 138
structure of, 140
Cell, or elementary part in dis-
ease, 94
Cells, structure of, 144
shape of, 145
size of, 148
in pathological formations,
148

een of, 149

and nuclei, new views on the
structure of, 117

polygonal, 145

young, 145

ciliated epithelial, ae
columnar “
squamous “ is

tessellated “ 145

of adipose tissue, 147

of bone, 146

of connective tissues, 146

of nervous tissue, 146

of striated muscular tissue,
146

of unstriped muscular tissue,

Cohnheim, 114
Compound microscope, invention
of, 15

198 INDEX.

D. K.
Diapedesis, 151 Klein, 123.
Dujardin, 76 Kupfer, 119
Dutrochet, 29 H L

Eberth, 122
Eimer, 123
Elastic tissue, yellow, 71, 92

F.

Fallopius, 13
Flemming, 121
Formed material, 87
=e secondary, 90
Frommann, 118

G.

Galen, 13

Germinal matter, 84
Globular theory, the, 21
Goodsir, 51

H.

Haeckel, 77

Haller, 17

Heitzmann, 118
Henle, 44

Hertwig, 120
Hodgkin, 28

Home, Sir Everard, 25
Hooke, 17

Huxley, 54 and 104

I.

Lllustrations, list of, xi
Intercellular substance, 91, 140

Leeuwenhoek, 17

M.

baat 16
Mayzel, 120
Meyen, 30

Milne, Edwards, 25

N..

Network, intracellular, 144
intranuclear, 131 and 144
Norris, 116
Nucleus, 128
discovery of, 30
Nucleolus, 132
“new explanation of, 125
Nucleus fibre, 142
Nuclear membrane, 131

Oken, 21
us

Plate, description of, 201
Preface to second edition, vii
to first ik
Prochaska, 24 gy
Protoplasm, 138 /*
of M. Schultze, 76

R.

Raspail, 29
Reichert, 44
Remak, 54
Robin, 98

INDEX. 199

8. Tubercle, 97
Sarcode of Dujardin, 76
Schleiden, 32 Vv.
Schwalbe, 119
Schwann, 32- Virchow, 65

Strassburger, 120
Stricker, 116
Swammerdam, 16

T.
Waller, 114
Tissues, formation of, 91 Wolff, 19

Todd, 64

EXPLANATION OF PLATE. 201

EXPLANATION OF PLATE,

ILLUSTRATING DR.’ KLEIN’S PAPER, ‘‘ OBSERVATIONS ON THE
STRUCTURE OF CELLS AND NUCLEI.”’

(From the Quarterly Journal of Microscopical Science, for July, 1878.)

Fiaures 1-11 (incl.) refer to preparations of stomach, 12-20 to
those of mesentery of newt. For method of preparing see text.
All figures are represented as seen on Hartnack’s small stand with
eye-piece ITI, and Zeiss’s objective F.

Fia. 1 represents a goblet cell; the intranuclear network is well
shown, and also the fibrils passing from this into the upper and
lower part of the cell.

Fig. 2.—Theintranuclear network separated from the membrane
of the nucleus.

Fia. 3.—A goblet-cell (like that of 1) as seen obliquely from
above, showing the intracellular network.

Fig. 4,—The intracellular network looked at vertically from
above. ;

Fia. 5.—Isolated nucleus showing the intranuclear network.

Fie. 6.—A gland-cell, showing the dense network of fine fibrils
of the cell-substance; the nucleus has escaped, but there are still
left a few fibrils, probably connecting the two networks, viz., the
intranuclear and the intracellular.

Fie. 7.—An isolated nucleus of a gland-cell; the wall of the
nucleus is broken at one place and the intranuclear fibrils are seen
passing outwards.

"+ Fie, 8.—A connective tissue corpuscle—endothelial plate—seen
in profile; both the intracellular fibrils and those of the intra-
nuclear network are well shown.

Fig. 9 —A similar cell seen from its broad surface.

Fie. 10 —Portion of acell; the intranuclear network shrunk.

Fie. 11.—An epithelial cell; the intracellular fibrils are well
shown. The top of the cell is seen in an oblique direction, and

202 EXPLANATION OF PLATE.

the network of fibrils is therefore brought into view. Prepara-
tion treated with Miiller’s fluid, and then with mixture of chromic
acid and spirit ; see text.

Fie. 12,—Isolated endotheliul plate of surface of mesentery.
The intracellular and intranuclear networks of fibrils are well
shown.

Fia. 18.—A capillary bloodvessel; the two upper nuclei are
seen from their broad side, the two lower in profile. They all
show the network of fibrils. In the lower portion of the wall of
the vessel an imperfect network of fibrils may be perceived.

Fic. 14, a and b.—Two unstriped muscle-fibres. The intra-
nuclear network of fibrils is well seen; these are in connection
with fibrils of the substance of the muscle-fibre; there are seen
numerous transverse markings along almost the whole length of
the muscle-fibre. That these transverse markings correspond to
tings which constitute the cortical part, 7. e., the sheath, is well
shown in a.

Fig. 15.—A non-medullated nerve-fibre of mesentery of newt;
the nerve-fibre has a delicate sheath, the nuclei of which contain
a distinct network of fibrils.

Fig. 16.—A connective tissue corpuscle seen sideways. The
nucleus contains a network of fibrils, in connection with that of
the cell-substance.

Fras. 17 and 18.—Two migratory cells; 17, a common pale one,
18, « coarsely granular one. Their nuclei show the network of
fibrils very well.

Fie. 19.—A cecal dilatation of a lymphatic vessel of the same
membrane, showing the nuclei of the endothelial cells forming
the wall of the lymphatic, and also nuclei of lymph-corpuscles.

Fig. 20,a and e.—Two connective tissue corpuscles; the dis-
tinction between “ ground-plate ”’ and “ fibrillar substance ”’ is well
shown ; the “ fibrillar substance”’ is in connection with the in-
tranuclear fibrils.

b, a nucleated plate ensheathing a minute non-medullated nerve-
fibre. The intranuclear network is seen in connection with the
processes and fibrils of the connective tissue corpuscle ¢.

WARS
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