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QiJAiN's ANATOMY





E. A.SCHAFER & G.D. THANE



VOL. I. P T - II.

GENERAL ANATOMY OR HISTOLOGY



E. A. SCHAFER



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3 SAN FRANCISCO fc



MEDICAL -fr LIBRAKV ^




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MEDICAL
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John Marshall Williamson
Memorial



QUAIN'S

ELEMENTS OF ANATOMY



EDITED BY



EDWARD ALBERT SCHAFER, F.R.S.

PROFESSOR OF PHYSIOLOGY AND HISTOLOOY IN UNIVERSITY COLLEGE, LONDON,



GEORGE DANCER THANE,

PROFESSOR OF ANATOMY IN UNIVERSITY COLLEGE, LONDON.
>

IN THREE VOLUMES.



VOL. L PART II.

GENERAL ANATOMY OR HISTOLOGY
By PROFESSOR SCHAFER.

ILLUSTRATED BY NEARLY 500 ENGRAVINGS, MANY OF WHICH ARE COLOURED

Cent!) ^Uttiou.



LONDON :
LONGMANS, GREEN, AND CO.,

AND NEW YORK : 15 EAST 16th STREET,

1891.

All rights reserved










LOS DON :
BRADBURY, AOKBW, & CO. LIMD., PRINTKRS, WHITEFRIARS.









1355 /



CONTENTS OF PAKT II.



GENERAL ANATOMY, OB HISTOLOGY.



PAGE

GENERAL ANATOMY, DEFINITION OF .171

Structural elements 172

Intercellular substance . . . .172

THE ANIMAL CELL 172

Structure of the cell-protoplasm . .172
Chemical nature of protoplasm . . 174

Vital phenomena of protoplasm . . 1 74
Amoeboid movements . . . . 175
Inception of foreign particles . . .176
Conditions influencing the contractile

manifestations of protoplasm . . .176
Effect of electrical and other stimuli upon

protoplasm . . . . . 177

Further considerations regarding the

structure of protoplasm , . . .177

THE NUCLEUS OF THE CELL . . . 179

Structure of the nucleus . . ..179

Chemical nature of the nucleus . . 182

Functions of the nucleus . . . . 182

THE DIVISION AND MULTIPLICATION OF

CELLS AND NUCLEI . . . . 183
Direct and indirect division of nuclei . 183
Karyokinesis, indirect or mitotic division

of the nucleus . . . . .185

Historical. ... . . 191

Recent literature . . . . .192
THE EPITHELIAL TISSUES

Structure of epithelial tissue in general . 194
Classification of epithelia . . . . 195

Stratified epithelium . . . . 195

Intercellular bridges and channels . . 196
Transitional epithelium . . . 197
Pavement epithelium . . . . 198

Columnar epithelium .... 198

Glandular epithelium . . . . 201

Ciliated epithelium .... 201

Structure of ciliated epithelium . . . 202
Nature of ciliary movement and influence

of varying conditions and reagents . . 204
Historical ...... 205

Recent literature ..... 206

THE CONNECTIVE TISSUES . . . 207

THE BLOOD 207

Red corpuscles of the blood . . . 208

Structure of the coloured blood-cor-
puscle ...... 210

Structure of the nucleated red corpuscles

of the lower vertebrata . . .212
Colourless corpuscles ot the blood . .213

General characters . . . .213

Action of reagents on . . . . 214

Other microscopic elements in blood . 215
Corpuscles of the lymph and chyle . . 216

Lymph 216



PAGE

. 216
. 216
216



Chyle

Development of the blood-corpuscles
Origin of the white blood-corpuscles .
Origin of the nucleated red corpuscles

of the embryo .... 217

t Origin of the red blood-disks. . . 218

Intracellular origin . . . 218

In the marrow of bone . . . 219

From white corpuscles . . . 220

In the spleen . . . . . 220

From blood-platelets . . . 220

Morphology of the red corpuscles . . 220

Historical 221

Recent literature . . . . . 221

CONNECTIVE TISSUE PKOPEK . . . 224

Areolar tissue . . . . 224

Elastic tissue ..... 225

Microscopic structure of connective tissue 226

Ground-substance ..... 226

White fibres . . . . . 226

Elastic fibres .'.... 227

Areolar tissue . . . . 229

Fibrous tissue ..... 230

Elastic tissue 230

Cells or corpuscles of connective tissue 231

Vessels and nerves . . . . 234

Adipose tissue . . . . . 235

Distribution . . . . . 235

Structure 236

Development 237

Pigment 238

Retiform or reticular tissue . . . 239
Development of connective tissue . . 240
Recent literature . . ... 243

CAETILAGE 244

Hyaline cartilage . . ... 244

Structure ...... 244

Articular cartilage . . . . 246

Costal cartilage ..... 247

Elastic or yellow cartilage . . . 248

White fibro-cartilage .... 249

Development of cartilage . . . . 250

Recent literature ..... 253

BONE

Chemical composition . . . . 254

Minute structure of bone . . . 254
Haversian canals . . . . . 256

Lamellae ...... 257

Lacunse and canaliculi . . . . 258

Cells of bone ..... 259

Apertures and decussating fibres of the

lamella} 259

Perforating fibres . . . . . 261
The periosteum ..... 265



IV



CONTENTS OF PART II.



PAGE

The marrow 265

Blood-vessels of bone .... 267

Lymphatics 267

Nerves 268

FORMATION AND GROWTH OF BOM; . . 268
Intramembranous ossification . . 268
Ossification in cartilage . . . . 272
Growth and absorption .... 280
Regeneration. . . . . . . 282

Recent literature 282

MUSCULAR TISSUE 285

Structure of cross-striated or skeletal

muscles

Fasciculi ......

Fibres, their figure and measurement .
Structure of the fibres, sarcolemma .
Muscular substance ....

Structure of the wing-muscles of insects 291
Comparison of the ordinary muscles of
insects and vertebrates with the wing
muscles of insects . . . .

Effect of the sarcoplasmic accumula-
tions upon the cross-striation of the
ordinary muscles ....

Changes in contraction . . . .

Appearances of muscle under polarised

light

Historical . . . .

Muscle-nuclei

Variations of structure in different

muscles, correlated with differences

of function .....

Mode of attachment of muscular fibres ;

ending of muscle in tendon
Blood-vessels. . , . .

Lymphatics

Nerves . ... . . .

Development of voluntary muscular tissue 300
Growth . .
Regeneration ....

Plain or unstriped muscular tissue .
Distribution ....

Development ....

Cardiac muscular tissue

Recent literature ....

NERVOUS SYSTEM ....

Nerve-fibres . . . . . '

Medullated nerve-fibres

Structure of medullated nerve-fibres



285
286
286
287
287



293



293
294

295
296
298



299

299
300

300



302
302
303
303
304
305
307
308
308
309



Axis-cylinder 310

Medullary sheath j 1 1

Nodes and internodes of Ranvier . 311
Medullary segments . . . . 313
Rod-like and reticular structures in the

medullary sheath . . . -313
Sheath of Schwann, primitive sheath or

neurolemma .....
Pale or non-medullated fibres ; fibres

of Remak ..... 315
Nerve-cell . . . . . . 316

- 317
the nerve-



315



Structure .
Sustentacular tissue of

centres ; neuroglia .
Construction of the nerves and nerve

roots

Vessels and lymphatics
Course of the nerve-fibres in the nerve

trunks .

Origins or roots of the nerves
Construction of the ganglia



322

324
326

326
327
331



PAGE

Situation 331

Termination or peripheral distribution of

nerves 333

Termination of sensory nerves . . . 334

Tactile cells 335

Tactile corpuscles or touch-bodies . . 335

End -bulbs 337

Tactile end-organs of birds . . . 340
Corpuscles of Herbst .... 340

Corpuscles of Vater or Paciuian bodies 341
Other modes of ending of sensory
nerves ...... 341

Nerve-endings m tendons . . . 347

Termination of motor nerves . . . 348

In the involuntary muscles . . . 348

In the cardiac muscular tissue . . 348

Nerves of voluntary muscles . . . 349

Development of the nerves . . . 352

Degeneration and regeneration . . 353

Recent literature . . . . 356

BLOOD-VESSELS ...... 360

Arteries ...... 360

Mode of distribution . . . . 360

Structure . . . . . .361

Internal coat ...... 362

Middle coat 363

External coat ... . . 364
Vessels and nerves of arteries . . 365

Veins 366

Mode of distribution .... 366

Structure . . . . . . 366

Internal coat ..... 366

Middle coat . . . . . . 366

External coat ..... 367

Valves . 367

Smaller arteries and veins and capillaries 368

Structure of the capillaries . . . 370

Structure of the small arteries and veins 370

Development of the blood-vessels . . 372

Recent literature 375

LYMPHATIC SYSTEM . . . . . 376
Distribution of lymphatic; vessels . . 376
Origin . . . . . . . 376

Structure 378

Valves 378

Relation of the lymphatics of origin to
the cells and cell-spaces of the connec-
tive tissue ...... 379

Termination of lymphatics . . . 381

Lymphatic hearts 381

Development of lymphatic vessels . . 382
Lymphatic glands ..... 382

Other organs composed of lymphoid
tissue . . . . . . 387

Serous membranes ... . . 388

Form and arrangement .... 388

Structure and properties . . . . 389

Blood vessels, lymphatics and nerves . 391
Synovial membranes . .... 392

Articular . . . . . . 392

Vaginal . . . . . . . 392

Vesicular or bursal .... 393

Structure . 393

Vessels and nerves .... 394

Development . . . ... 395

Recent literature of lymphatic system . 395

SECRETING GLANDS 396

Changes in the cells during activity . 396
Modifications in form of the secreting
surface 398



CONTENTS OF PART II.



The acinous, racemose or acino-tiibular
compound glands .... 400

Mechanism of the discharge of secretion
from a gland ..... 403

Recent literature ..... 403

Mucous MEMBRANES . . . . 405

Structure ....... 406

The epithelium ..... 406

The coriurn . . . , . . 406
Vessels and nerves . . . 408

Papillae and villi . . . . . 408

Glands . . . . . . . 409

THE SKIN 410

The epidermis, cuticle or scarf-skin . 410
Stnicture . . . . . . 410

Development of the epidermis . .412
The true skin, cutis vera, derma or

corium ...... 413

Structure .'.... 414



Papillae

Blood vessels and lymphatics

Nerves .....
Nails and hairs ....

Nails

Formation and growth of the nails
Hairs ......

Root of the hair

Structure of the hair-follicle

Nerves of the hairs

Muscles of the hairs .

Development of hair in the foetus
The sebaceous glands

Development of the sebaceous glands . 427
The sudoriferous glands or sweat glands. 427

Distribution ..... 429

Size . . . . . . . 429

Development . . ... . 429

Recent literature . .... 430



PAGE

415

415
. 416

417
. 417

419
. 420
. 422
. 422

423

424

424
426



VOL. I.



GENEEAL ANATOMY, OR HISTOLOGY.

By E. A. SCHXFER.

GENERAL ANATOMY, or as it is now more commonly termed HISTOLOGY, is that
branch of Anatomy which treats of the structure of the textures. As has already
been explained in the Introduction to this volume, the body of every one of the
higher animals is made up of organs adapted for the performance of its several
functions, and these organs are themselves composed of various tissues or textures.
In order that the structure of any organ or part of the body should be understood, it
is necessary to study, both together and separately, the several tissues of which it is
composed, so as to ascertain their composition and the manner in which they are
combined to constitute the organ or part in question. This is chiefly effected by
minute dissociations and thin sections, which are observed with the aid of the micro-
scope, and hence the terms "Minute Anatomy" and "Microscopic Anatomy" are
also applied to this branch of the science. It is found that when the body is thus
dissected and analyzed by the aid of the microscope, that the number of distinct
tissues which are met with is comparatively small, and some of these again, although
at first sight to all appearance distinct, yet have so much in common in their struc-
ture and origin one with another (forms of transition also being met with between
them), that the number becomes still further reduced. The elementary tissues which
may be thus enumerated are as follows :

The epithelial tissues.
The connective tissues.
The muscular tissues.
The nervous tissues.

Particles which are met with in the fluids of the body, such as the corpuscles in
the blood and lymph, are also described amongst the elementary tissues.

Many of the organs are formed wholly of a single one of these elementary tissues,
or with a comparatively slight intermixture of others. Thus the muscles are made
up almost entirely of muscular tissue, with but a small intermixture of connective
tissue, blood-vessels and nerves ; whilst the cartilages are composed wholly of a
variety of connective tissue. On the other hand, there are certain organs or parts of
the body not in themselves distinguished by the preponderance of any special tissue,
but compounded of two or more in varying proportion, the structure of which it is
nevertheless convenient to describe along with the tissues, on account of their wide
distribution in the body, and their uniformity of structure in different parts. Such
are :

Blood-vessels.

Lymphatic vessels.

Lymphatic glands and bodies of like structure.

Serous membranes.

Synovial membranes. *

Secreting glands.

Mucous membranes.

Integument.

VOL. i. sr



172 THE ANIMAL CELL.

Every texture taken as a whole was viewed by Bichat (Anatomic generate, 1801)
as constituting a peculiar system, presenting throughout its whole extent in the body
characters either the same, or modified only so far as its local connections and uses
render necessary ; he accordingly used the term " organic systems " to designate the
textures taken in this point of view. Of the organic systems above enumerated some
are found in nearly every organ ; such is the case with the connective tissue, which
serves as a binding material to hold together the other tissues which go to form an
organ ; the vessels, which convey fluids for the nutrition of the other textures, and
the nerves, which establish a mutual dependence among different organs. These
were named by Bichat the " general systems," to distinguish them from others such
as the cartilaginous and osseous, which being confined to a limited number or to a
particular class of organs, he named " particular systems."

Structural elements. When any tissue is separated by the aid of the micro-
scope into the simplest parts which possess assignable form, such parts are termed
the structural elements of the tissue. In almost every tissue, some at least of these
structural elements retain to a certain extent the characters of the elementary
corpuscles of which the whole body is originally composed. These structural
elements are named the " cells " of the tissue. Others again lose for the most part
those characters, and becoming elongated and modified in structure are termed the
fibres of the tissue, whilst in other cases, fibres are formed not from the cells but
between them. Except the epithelium, all the tissues have fibres as characteristic
structural elements, and some, as the connective tissue, fibres of more than one
kind. But structurally and chemically as well as functionally, the fibres of the several
tissues differ widely from one another.

Intercellular substance. In addition to these separable structural elements,
many of the tissues are composed of a homogeneous matrix or ground substance, in
which the structural elements are imbedded. This matrix may exist in considerable
quantity, as in some varieties of connective tissue, or on the other hand it may be
almost imperceptible in amount, serving merely as a cementing material to connect
together the individual tissue-elements, as in epithelium. From its softness, clear-
ness and homogeneity, this ground substance is often apt to escape observation, but
its existence may always be rendered evident in consequence of the property it
exhibits of combining with salts of silver, a brown deposit of metallic silver occurring
in it on subsequent exposure to the light (v. Eecklinghausen).

Since all the animal tissues however diversified they may appear, originate as
collections of the elementary corpuscles or cells above spoken of, and since these cells
remain, many of them, as constituent elements of the formed tissue, we must first of
all consider minutely what it is that constitutes an animal cell, what is its structure,
its chemical composition, its physical and vital properties, and how it becomes
reproduced and multiplied.

THE ANIMAL CELL.

An animal cell is a corpuscle of microscopic dimensions, the cells of the human
body seldom exceeding -g^th of an inch in diameter, and many being as small as one-
tenth of this or even less. But whether small or large, every cell consists of two dis-
tinct parts : of the main substance of the cell, which has received the name of proto-
plasm (fig. 201,^?), and of a minute vesicular structure, generally placed near the
centre of the cell, and termed its nucleus (n).

THE PROTOPLASM OF THE CELL.

Structure of the cell-protoplasm. Until comparatively recent years it was
universally held that the principal or living substance of the cell to which the name



STRUCTURE OF CELL-PROTOPLASM.



173



of protoplasm is applied is a completely homogeneous material, which, although it
might contain granules of solid matter or globules of watery fluid (vacuoles) im-
bedded in it, is nevertheless itself entirely devoid of structure (fig. 201, A). It is
possible that this view may still hold good for some cells both animal and vegetable,
but in most cells, especially those which are " fixed " or non-amoeboid, it is found that a
differentiation of the protoplasm has occurred in such a manner that a part of it
appears under high powers of the microscope in the form of a network or spongework,





__ -p.



Fig. '201. A, DIAGRAM OF A CELL, THE PROTOPLASM OF WHICH APPEARS STRUCTURELESS, BUT is OCCUPIED

BY VACUOLES AND GRANULES. B, DIAGRAM OF A CELL, THE PROTOPLASM OF WHICH IS COMPOSED OF
SPONGIOPLASM AND HYALOPLASM.

p, protoplasm ; n, nucleus; ', nucleolus.

Fig. 202. CELL FROM THE EPIDERMIS OF AN EMBRYO OF SALAMANDRA, TREATED WITH 1 PER CENT.
HYDROCHLORIC ACID. (After Kolliker.)

7i, nucleus, with network of chromoplasm ; p, protoplasm, showing a reticulum of plastin.

whilst the rest of the protoplasm occupies the meshes of this network (figs. 201, B, and
202). The network is known as the reticulum or spongioplasm ; while the substance
which occupies its meshes may be designated the enchylema (Carnoy) or hyaloplasm.



Fig. 203. CELL WITH RADIALLY DISPOSED

RETICULUM FROM THE INTESTINAL EPI-
THELIUM OF A WORM. (Carnoy.)

me, membrane of the cell ; pc, pro-
toplasm of the cell ; mn, membrane of
the nucleus ; pn, achromatic substance
of the nucleus, with the convoluted chro-
inatin filament, bn, contracted into the
centre.



me



pc-




The proportion which these bear
to one another varies in different
cells, but it may be stated that as
a general rule the amount of
spongioplasm becomes augmented
as the development of the cell
proceeds, and that the younger a
cell the greater is the relative
amount of hyaloplasm.

The shape and size of the meshes of the reticulum also vary in different cells,
and even in different parts of the protoplasm of the same cell. Thus in some cells
the constituent fibrils or cords of the spongioplasm are disposed radially from the nu-
cleus to the periphery (fig. 203), and the meshes are radially elongated ; in others they
are disposed evenly in all directions, whilst in some a concentric disposition of the
fibrils can be determined. The fibrils are invariably united together (as is implied by
the use of the term reticulum) into a network, and at the nodes of the network where

N 2



174 CHEMICAL NATURE OF PROTOPLASM.

they join one another they may be slightly enlarged. These enlargements at the
junctions of the fibrils as well as the optical sections of those fibrils which are coursing
in a direction more or less parallel to the visual axis of the observer appear, when
the horizontally running fibrils are not clearly seen, as distinct points or granules in the
protoplasm ; it is perhaps for this reason that the cell-substance was formerly con-
stantly described as being finely granular.

The external layer of the protoplasm of many cells, especially that of fixed cells
like those which constitute the varieties of epithelium, and in a notable degree the
cells of the higher plants, may become altered by the deposition in or upon it of
chemical substances which render it firmer and more resistant than the rest of the
cell. In this way a cell-membrane (fig. 203, me) may be formed, either complete or,
more commonly, furnished with minute pores which allow of the passage of fluid or
the connection of the cell with its neighbours by filaments of protoplasm which
traverse the pores. In those cases where a cell-membrane is formed the peripheral layer
of the protoplasm is usually furnished with a denser reticulum than the rest, but this
is not the case in cells like the white corpuscles of the blood and the amoeba which have
no cell-membrane. In these the exterior of the protoplasm may be clearer than the
interior, indeed a reticulated appearance may be altogether wanting in it, or may be re-
placed by a few radial striae. Such clearer peripheral protoplasm is sometimes known as
the " ectoplasm " to distinguish it from the granular-looking " endoplasm " around
the nucleus.

Chemical nature of protoplasm. Protoplasm may contain imbedded in its
substance (generally in the hyaloplasm) various adventitious materials, such as
granules of proteid or starch, globules of fat, crystals of various kinds, pigment-
granules and globules of watery fluid (vacuoles). These may have been formed
by the cell either from its own substance or directly from materials imbibed in a
fluid form, or they may have been bodily incepted in the solid form, but they are not
essential to its composition, although no doubt when they occur they play an
important part in relation to its vital activities. 1 Apart from these the protoplasm
of all vegetable and animal cells, since it is similar in its functional manifestations,
is also in all probability similar in chemical constitution. So far as the
proteids are concerned, it would appear that there is a preponderance of the less
stable members of that group, such as globulins and albumoses as distinguished
from the more stable albumins, and that these cell-proteids appear to be associated
with ferment-like reactions like those which produce digestive or coagulative
changes. The chief proteid found is a muco-globulin, containing phosphorus, to
which the name plastin has been applied. It would further seem that the presence
of certain inorganic substances, and especially calcium, is essential to the
life and therefore to the functions of protoplasm, but in what manner the
lime may be combined with the organic basis of the living material, remains
as yet quite undetermined. It must further not be forgotten that water, enters
very largely into the composition of all living material. 2



CHEMICAL AND PHYSICAL CHANGES. AMOEBOID MOVEMENTS.

Vital phenomena of protoplasm. During the life of a cell its protoplasm' is
constantly undergoing chemical and physical changes. The chemical changes are in
some measure determinable by comparing the products which are given off by the

1 The terms " deutoplasm " and " paraplasm " (Kupffer) have sometimes been applied to materials
contained within a cell, which are not considered to constitute a part of the actual protoplasm.

2 For an account of the present state of knowledge regarding the chemistry of protoplasm and of the
cell generally, consult Halliburton, "A Textbook of Chemical Physiology," 1891, pp. 190 to 211.



VITAL PHENOMENA OF PROTOPLASM. 175

cells of a tissue with the nutritive material which they absorb. In all the higher
.animals this nutritive material is the blood or lymph, but the products which are
formed are not entirely the same for all cells, since they vary in some measure with
the specific activity of the cell ; thus the cells of the salivary glands yield the saliva,
those of the mammary gland milk, and those of the liver form, besides other substances,
glycogen. But all protoplasm, whatever may be its specific function, has this in
common, viz., that it absorbs and combines with oxygen, and yields carbon dioxide
and other products of oxidation, and as a result of these processes of oxidation heat
and other kinds of energy are produced. These chemical changes are always more
marked as the functional activity of the cell becomes increased, and accordingly any
circumstances which tend to promote the activity of protoplasm, such as warmth,
electrical or other stimulation, the action of certain drugs, tend proportionally to
increase the activity of its chemical processes. One general chemical property
of living protoplasm is that by virtue of which it is able to assimilate and eventually
to convert into its own substance non-living proteid material. In this manner the
protoplasm of a cell may increase in amount, or in other words the cell may grow ;



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