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=Embryology of the Pig=
=Embryology of the Pig=
Frontispiece  
[[File:Bradley M. Patten.jpg|thumb|alt=Bradley M. Patten|link=Embryology History - Bradley Patten|Bradley Patten ( -1971)]]
Frontispiece


Reconstruction (X 17.5) showing the organ systems of a 9.4 mm. pig embryo. For explanation see figures 60 and 66.  
Reconstruction (X 17.5) showing the organ systems of a 9.4 mm. pig embryo. For explanation see figures 60 and 66.


By BRADLEY M. PATTEN
By Bradley M. Patten


Professor of Anatomy in the University of Michigan Medical School  
Professor of Anatomy in the University of Michigan Medical School




THIRD EDITION
Third Edition




WITH COLORED FRONTISPIECE
With Colored Frontispiece


AND 186 ILLUSTRATIONS IN THE TEXT (CONTAINING 412 FIGURES)  
And 186 Illustrations In The Text (Containing 412 Figures) Of Which 6 Are In Color
OF WHICH 6 ARE IN COLOR




Philadelphia : THE BLAKISTON COMPANY : Toronto


Philadelphia : THE BLAKISTON COMPANY : Toronto


Third Edition




Third Edition
Copyright, October 1948, by The Blakiston Company




Copyright, October 1948, by The Blakiston Company
By P. Blakiston's Son & Co.




BY P. Blakiston ’s Son & Co.  
Copyright, 1951, by P Blakiston's Son & Co , Inc.




Copyright, 1951, by P Blakiston’s Son & Co , Inc.
{{Patten1951 TOC}}
 


Preface to Third Edition  
==Preface to Third Edition==


In making the revision for a new edition of this book it did not seem desirable essentially to change its form or scope. Effort has been concentrated on improving the presentation of the original subject matter and bringing it up to date, rather than on its expansion. The entire book has been reset to a greater page width which has permitted enlarging certain of the illustrations that, in earlier editions, had proved to be too greatly reduced. With the generous cooperation of the publishers several of the important plates on the cardiovascular system have been remade to a larger scale and with color. A number of new illustrations have been drawn for sections where experience has shown that students needed additional graphic assistance in interpreting their laboratory material. It is hoped that these changes will all contribute toward making the book as a whole more serviceable.


In making the revision for a new edition of this book it did not
seem desirable essentially to change its form or scope. Effort has been
concentrated on improving the presentation of the original subject
matter and bringing it up to date, rather than on its expansion. The
entire book has been reset to a greater page width which has permitted enlarging certain of the illustrations that, in earlier editions,
had proved to be too greatly reduced. With the generous cooperation
of the publishers several of the important plates on the cardiovascular
system have been remade to a larger scale and with color. A number
of new illustrations have been drawn for sections where experience
has shown that students needed additional graphic assistance in
interpreting their laboratory material. It is hoped that these changes
will all contribute toward making the book as a whole more serviceable.


Bradley M. Patten  
Bradley M. Patten


August 1048  
August 1048




Preface to First Edition  
==Preface to First Edition==


This book represents an endeavor to set forth in brief and simple form the fundamental facts of mammalian development. The thread of the story and the illustrations have been based on pig embryos because of their value and availability as laboratory material. But special stress has been laid on the embryological phenomena involved instead of on the details of specific conditions existing in the pig. Throughout the book, every efTort has been made to present developmental processes as dynamic events with emphasis on their sequence and significance, rather than as a series of still pictures of selected stages.


This book represents an endeavor to set forth in brief and simple
form the fundamental facts of mammalian development. The thread
of the story and the illustrations have been based on pig embryos
because of their value and availability as laboratory material. But
special stress has been laid on the embryological phenomena involved
instead of on the details of specific conditions existing in the pig.
Throughout the book, every efTort has been made to present developmental processes as dynamic events with emphasis on their sequence
and significance, rather than as a series of still pictures of selected
stages.


Obviously no book can deal fully with all phases of development,  
Obviously no book can deal fully with all phases of development, even in a single form, and still remain serviceable as a text. As this book is for the student, it has seemed expedient, for the sake of clearness and simplicity, to omit many things which I should like to have included. My primary aim has been to write an account in which the essentials stand out adequately interpreted and unobscured by a multiplicity of details — to lay a foundation which can be further built upon in accordance with special needs or individual desires.
even in a single form, and still remain serviceable as a text. As this  
book is for the student, it has seemed expedient, for the sake of clearness and simplicity, to omit many things which I should like to have  
included. My primary aim has been to write an account in which the  
essentials stand out adequately interpreted and unobscured by a  
multiplicity of details — to lay a foundation which can be further  
built upon in accordance with special needs or individual desires.  


Bradley M. Patten


January 1927
Bradley M. Patten


January 1927


==Acknowledgment==
==Acknowledgment==
The pleasantest thing about working on this book has been the generous aid I have received from many sources. Throughout the preparation of the initial edition the encouragement, criticism, and suggestions of my colleagues, Dr. F. C. Waite and Dr. S. W. Chase, were of the greatest help. In the preparation of material and in making the illustrations for the first edition the beautifully accurate work of Miss Kathryn Toulmin was of inestimable value. In making the new drawings added in the third edition I was fortunate in securing the unusually able assistance of Mrs. Dorothy Van Eck.




The pleasantest thing about working on this book has been the
Dr. G. L. Streeter and Dr. C. H. Heuser of the Carnegie Institute allowed me free use of their extensive collection of young embryos and generously gave me many photographs made from their material. To Mrs. Charles S, Minot I am indebted for permission to use several figures from the late Professor Minot's works. The accrediting in the figure legends of these and other borrowed illustrations by no means covers my obligation to other writers. Practically the entire bibliography is a statement of indebtedness for information and ideas.
generous aid I have received from many sources. Throughout the
preparation of the initial edition the encouragement, criticism, and
suggestions of my colleagues, Dr. F. C. Waite and Dr. S. W. Chase,
were of the greatest help. In the preparation of material and in making
the illustrations for the first edition the beautifully accurate work of  
Miss Kathryn Toulmin was of inestimable value. In making the new
drawings added in the third edition I was fortunate in securing the
unusually able assistance of Mrs. Dorothy Van Eck.  


Dr. G. L. Streeter and Dr. C. H. Heuser of the Carnegie Institute
allowed me free use of their extensive collection of young embryos
and generously gave me many photographs made from their material.
To Mrs. Charles S, Minot I am indebted for permission to use several
figures from the late Professor Minot’s works. The accrediting in the
figure legends of these and other borrowed illustrations by no means
covers my obligation to other writers. Practically the entire bibliography is a statement of indebtedness for information and ideas.


I wish I might acknowledge individually the helpful services  
I wish I might acknowledge individually the helpful services rendered by many of my students, but they are too numerous. Several reconstructions which I have used directly or indirectly have been largely their work. Of even greater assistance have been their suggestions during the shaping of the work — suggestions of especial value because they were made from a point of view difficult for an instructor to appreciate without such aid.
rendered by many of my students, but they are too numerous. Several  
reconstructions which I have used directly or indirectly have been  
largely their work. Of even greater assistance have been their suggestions during the shaping of the work — suggestions of especial value  
because they were made from a point of view difficult for an instructor  
to appreciate without such aid.  


To The Blakiston Company, I am indebted for much helpful
cooperation and especially for their liberality with regard to the
number and quality of the illustrations.


No person other than my wife could have deciphered and put
To The Blakiston Company, I am indebted for much helpful cooperation and especially for their liberality with regard to the number and quality of the illustrations.
into usable form manuscript of the character I frequently turned over
to her for revision and typing. Without her generous help the preparation of the text would have been much more arduous and long
delayed.  


Bradley M. Patten


No person other than my wife could have deciphered and put into usable form manuscript of the character I frequently turned over to her for revision and typing. Without her generous help the preparation of the text would have been much more arduous and long delayed.




==Chapter  12==
Bradley M. Patten




Tke Histogenesis of Bone and tlie
Development of tke Skeletal System


{{Footer}}


I. Histogenesis of Bone
{{Patten1951 TOC}}
 
Histologically bone belongs to the group of tissues known as the
connective and supporting tissues. In spite of their widely varying
adult conditions these.tissues are all similar in that the secreted parts,
rather than the cells themselves, carry out the functional role characteristic of the tissues. It is the secreted, fibrous portion of the binding
connective tissues which ties together various other tissues and
organs ; it is the secreted matrix of cartilage and of bone which affords
rigid support and protection to soft parts and furnishes a lever system
on which the muscles may be brought into play.
 
The cellular elements of these tissues must not be overlooked,
however, in emphasizing the functional importance of the cell
products. The cells are, so to speak, the power behind, in that they
extract the appropriate raw materials from the circulation, elaborate
them within their cytoplasm, and deposit the characteristic secretion
as an end-product. Moreover after the fiber is formed or the matrix
is laid down, it is dependent on the cells for maintenance in a healthy
active conditidn.
 
Embryologically the entire connective-tissue group arises from
mesenchymal cells. It is not surprising, in view of their closely related
functions and their derivation from a common type of ancestral cell,
that one type of connective tissue may be converted into or replaced
by another. This facility for changing the type of specialization is
sometimes referred to as plasticity.
 
The plasticity of the connective-tissue series is well exemplified
in the development of bone. Bone does not form in vacant spaces.
It is always laid down in an area already occupied by some less highly
specialized member of the connective-tissue family. The formation of
some bones begins in areas already occupied by connective tissue —
such bones are said to be intramembranous in origin, or are spoken
 
271
 
 
 
272 HISTOGENESIS OF BONE AND DEVELOPMENT OF SKELETAL SYSTEM
 
 
of as membrane bones. Other bones are laid down in areas already
occupied by cartilage. In this case they are said to be pndochondral in
origin, or, are called cartilage bones. It should be clearly borne in mind
that these terms apply solely to the method by which a bone develops
and do not imply any differences in histological structure, once the
bone is fully formed.
 
Likewise we should know at the outset what histologists mean
when they speak of cancellous bone and compact bone. These terms
refer not to the method of origin of the bone but to its density when
fully formed. Developmen tally all bone goes through the spongy or
cancellous stage. Some bones later become compact, others remain
cancellous. Most bones are compact in some areas and cancellous
in others.
 
The subject of bone development can be presented more simply
if we take up first the formation of primary cancellous bone intramembranously ; then the method by which this same type of spongy
bone is formed within cartilage, and finally the changes by which
cancellous bone, formed in either of the above ways, may become
secondarily compact.
 
Intramembranous Formation of Primary Cancellous Bone. In
 
an area where intramembranous bone formation is about to begin
we find an abundance of mesenchymal cells congregated and numerous small blood vessels present. The mesenchymal cells soon exhibit a
tendency to cluster together in more or less elongated groups here
and there throughout the area. If we study a group of this type which
has been aggregated for a short time we can make out the beginning
of a definite plan of organization. Near the axis of the cord delicate
fibers appear, produced by the secretory activity of the cells. As this
fibrous strand becomes more definite, the cells tend to become ranged
against it (Fig. 151, A). In so doing they retract the cytoplasmic
processes which are so characteristic of undifferentiated mesenchymal
cells and become rounded. In this stage we have essentially a connective tissue in which the fibrous strands are for the most part rather
widely separated from one another, and in which each strand has,
lined up against it^ the cells responsible for its production.
 
The actual deposition of bone matrix begins very soon after the
establishment of these primordial strands of mesenchymal cells and
fibers. In fact one usually finds the formation of bone beginning on
the older part of a strand while the strand itself is still being extended
at one end by the aggregation of more mesenchymal cells (Fig. 151,
 
 
 
HISTOGENESIS OF BONE
 
 
273
 
 
 
I'k;. 151, Formation of trabeculae of membrane bone. Projection drawings from the mandible of a pig embryo 130 mm. in length (cf. Fig. 182).
 
Abbreviations: Matrix cal., ossein matrix impregnated with calcium
salts; Matrix oss., ossein matrix not yet impregnated with calcium salts.
 
A). When the mesenchymal cells ranged against the fibrous axis of
such a strand become active in the secretion of calcareous material
they are spoken of as osteoblasts. We should not lose sight of the fact
that they are the same cells which formed the fibrous axis of the
original strands, given a new name in deference to their further
specialization and altered internal chemistry.
 
In studying the deposition of bone matrix one must bear in mind
its dual nature. The matrix consists of an organic fibrous framework
which is impregnated by a subsequent deposit of inorganic calcium
compounds. We may liken the matrix of bone to reinforced concrete*
In the making of a road or a wall, a meshwork of steel is first placed in
the forms and concrete is then poured in. The steel gives the finished
structure tensile strength and a certain amount of elasticity, the concrete gives form and hardness. So in bone the organic fibers {ossein
fibers) impart strength and resilience, while the calcium salts whh
which the fibers are impregnated give to the completed matrix body
and rigidity.
 
 
 
274 HISTOGENESIS OF BONE AND DEVELOPMENT OF SKELETAL SYSTEM
 
 
The two steps in the deposition of bone matrix may be demonstrated readily in areas where active bone forma^tion is going on,
owing to the fact that the presence of calcium compounds- in a tissue
markedly increases its affinity for stains. Even after most of the calcium salts have been removed from the ossein framework by treatment
of the tissue with acids (decalcification) to permit the making of
sections, the staining reaction is still apparent. This indicates that
the ossein fibers in which calcium has once been deposited are more
or less permanently changed chemically even though all the calcium
possible is subsequently removed.
 
If we look at a strand on which the osteoblasts have been active
for a time (Fig. 151, B) we see, next to the osteoblasts, a zone of bone
matrix which takes very little stain. This is the newly deposited
organic portion of the matrix as yet unimpregnated with calcium salts.
It consists of a feltwork of minute fibers so delicate and so closely
matted together that it is very difficult in ordinary preparations to see
the individual fibers at all. Slightly farther from the osteoblasts the
matrix is densely stained (Fig. 151, B). This part of the matrix has
been impregnated with calcium salts, chiefly phosphates and carbonates, and has thereby been converted into true bone matrix. The
calcium utilized by the osteoblasts in this process is brought to them
by the blood stream where it is carried in soluble form, probably in
organic linkage. It is interesting to note in this connection that the
presence of calcium and of phosphates in the blood is not in itself all
that is necessary for this process. There must be present also sufficient
vitamin D, which in some way facilitates the extraction by the osteoblasts of these raw materials from the blood and their deposition in
insoluble form as part of the bone matrix. The absence of vitamin D
from the system results in the formation of bone matrix deficient in
calcium salts and therefore lacking in rigidity — a cohdition not infrequent in pigs. Stock raisers have miscalled this condition rheumatism
but it is really the same condition known medically as rickets.
 
In the deposition of the matrix, the fibrous core of the original
strand serves as a sort of axis on which the first matrix is laid down.
When such a strand is completely invested by bone matrix, it is called
a trabecula (little beam). As the osteoblasts continue to secrete and
thereby thicken the trabecula, the accumulation of their own product
forces them farther and farther away from the axial strand about
which the first of the matrix was formed. The new matrix added is
not laid down uniformly. It is possible to make out in it markings
 
 
 
HISTOGENESIS OF BONE
 
 
275
 
 
which are suggestive of the growth rings of a tree. Apparently the
osteoblasts work more or less in cycles, depositing a succession of thin
layers of matrix. Each of these layers of the matrix is called a lamella
(Fig. 152). As the row of osteoblasts is forced back with the deposit of
each succeeding lamella, not all the cells free themselves from their
secretion. Here and there a cell is left behind. As its former fellows
 
 
Erythroblast extruding nucleus
 
Reticular connective-tissue cell j Erythroblast in mitosis
Young erythroblast
 
Normoblast
Blood vessel .
 
Fat cell --tHemocytoblast —
 
Granuloblast -
Hemocytoblast
in mitosis
 
Polykaryocyte.
 
 
 
:
 
Osteoblast ^ t
 
 
I
 
Bone cell
 
 
Bone
 
lamella
 
 
Fio. 152. A small area of bone and adjacent marrow
as seen in highly magnified decalcified sections. The
drawing has been schematized somewhat to emphasize
the relations of the cytoplasmic processes of the osteoblasts* and the bone cells so important in nutrition. In
the adjacent marrow developmental stages of various
types ofiiiijioQd cells, have been
 
 
continue to pile up new matrix, it becomes completely buried (Fig,
151, B). An osteoblast so caught and buried is called a bone cell
{osteocyte)^ and the space in the matrix which it occupies is called a
lacuna. The bone cells, thus entrapped, of necessity cease to be active
bone formers, but they play a vital part in the maintenance of the
bone already formed. They have delicate cytoplasmic processes
radiating into the surrounding matrix through minute canalidhli.
The processes of one cell come into communication with the processes
of its neighbors (Fig. 152). Thus the bone cells nearer to blood vessels
 
 
 
276 HISTOGENESIS OF BONE AND DEVELOPMENT OF SKELETAL SYSTEM
 
 
 
 
Fig. 153. Diagrams showing stages in establishing of a characteristic area
of primary cancellous bone by extension and coalescence of originally
separate trabeculae.
 
absorb and hand on materials to their more remote fellows which in
turn utilize these materials in maintaining a healthy condition in the
organic part of the bone matrix. It is the senescence of these cells with
the consequent lowering of their efficiency and the resultant deterioration of the ossein component of the matrix which is in part responsible
for the decreased resiliency of the bones in advanced age.
 
As the various trabeculae in an area of developing bone grow,
they inevitably come in contact with each other and fuse. Thus
trabeculae, at first isolated, soon come to constitute a continuous
system (Fig. 153). Because of its resemblance to a latticework (Latin —
cancellus), bone in thb condition, where the trabeculae are slender
and the spaces between them extensive, is known as cancellous bone.
The spaces between the trabeculae are known as marrpw spaces.
 
Endochondral Bone Formation. As the term implies, endochondral bone formation goes on within cartilage. It cannot be stated
too strongly that cartilage does not, in this process, become converted
int# bone. Cartilage is destroyed and bone is formed where the
cartilage used to be. The actual bone formation is essentially the
same as in the case of membrane bone. The phenomena of special
 
 
 
mSTCXJENESIS OF BONE
 
 
277
 
 
interest in connection with this type of bone development are those
involved in the destruction of the cartilage preliminary to the formation of bone.
 
Cartilage Formation. To trace the process logically we must start
back with the formation of cartilage. The first indication of impending
chondrogenesis is the aggregation of an exceedingly dense mass of
mesenchymal cells. This cell mass gradually takes on the shape of the
cartilage to be formed. The histogenetic changes involved are not at
first conspicuous. During the period of preliminary massing the cells
have been migrating in from surrounding regions and also increasing
the local congestion by rapid proliferation. As they are packed in
together they lose their processes and become rounded (Fig. 154, A,
1). When it seems as if no more cells could possibly be crowded in,
the course of events changes. The cells begin to separate from one
another. This is due to the fact that they have become active in
 
 
 
Fig. 154. Photomicrographs of developing cartilage. The areas photographed were from the margins of the paranasal cartilage of pig embryos
between 25 and 30 mm. in length. For location of cartilage in head see
figure 175.
 
A, Early stage showing: at (1) the massing of mesenchymal cells which
were about to be incorporated in the growing margin of the cartilage; and at
(2) an area where matrix formation is already beginning.
 
B, Slightly more advanced stage of the same cartilage showing: at (1)
increase in the amount and density of the matrix in the center of the growing
cartilage; at (2) concentration of the surrounding mesenchyme to form tire
perichondrium; and at (3) the addition of new cartilage matrix peripherally*
 
 
 
278 HISTOGENESIS OF BONE AND DEVELOPMENT OF SKELETAL SYSTEM
 
 
secreting. It is the accumulation of the secretion of the cells which
gradually forces them farther and farther apart unti^ they come to lie
isolated from one another in the matrix they have produced (Fig. 1 54,
A, 2). Such a method of increase in m^ss, where there are many
scattered growth centers contributing independently to the increase
in bulk of the whole, is known as interstitial growth. This interstitial
growth of young cartilage stands in sharp contrast to the appositional
growth of such rigid substances as bone or dentine or enamel where
the matrix is laid down in successive layers one upon another. Obviously interstitial growth implies plasticity of the substance produced.
Were the substance produced unyielding, the very activity of a number of growth centers within it would soon crowd those growth centers
to obliteration.
 
As the cartilage matrix is increased in amount its affinity for basic
stains becomes more marked, due probably to increase in concentration of the characteristic substance in it known chemically as chondrin.
At the same time the matrix becomes more rigid with a resultant
checking of interstitial growth. The cells continue to secrete to a
certain extent, however, as evidenced by the fact that in mature
cartilage the matrix immediately surrounding the cells becomes more
dense than the rest of the matrix. This area of denser matrix around
the lacuna in which the cell lies is known as the capsule. As the cartilage
grows older the capsules become more conspicuous and many of
them come to contain more than one cell. These nests of cells in a
common capsule are the result of cell divisions, following which the
daughter cells are held imprisoned in the original capsule of the
mother cell — further evidence of the loss of plasticity in the matrix.
 
The formation of a matrix so rigid that interstitial growth is
checked, takes place first centrally in an area of developing cartilage.
When the center has become too rigid for interstitial growth to continue, appositional growth begins to take place peripherally. While
the cartilage has been increasing in mass it has been acquiring a
peripheral investment of compacted mesenchyme. This investing
layer of mesenchyme soon becomes specialized into a connectivetissue covering called the perichondrium. The layer of the perichondrium next to the cartilage is less fibrous than the outer layer and the
cells in it continue to proliferate rapidly and become active in the
secretion of cartilage matrix. For this reason it is known as the
chondrogenetic layer of the perichondrium. It is through the activity
(rf the chondrogenetic layer that the cartilage continues to grow
 
 
 
Cartilage
 
ceil
 
 
Cart. trab. 1
 
 
 
Osteoblast
 
 
— Bone
ntotrix
 
 
Bone trabecula
 
 
Periosteum
 
 
Fig. 155. Drawing showing periosteal bud and an area of endochondral
bone formation from the radius of a 125 mm. sheep embryo. The small sketch
indicates the location of the area drawn in detail.
 
Abbreviations: Cart, eros., area from which cartilage has recently been
eroded; Cart, pre-eros., area with cartilage cells enlarged and arranged in
rows presaging erosion; Cart, trab,, remnant of cartilage matrix which has
become calcified and serves as an axis or core about which bone lamellae are
deposited to form a bone trabecula; Mes., mesenchymal cell.
 
279
 
 
 
280 HISTOGENESIS OF BONE AND DEVEI.OPMENT OF SKELETAL SYSTEM
 
 
peripherally, by apposition, long after interstitial growth has ceased
in the matrix first formed. ^
 
Cartilage Erosion. When a mass of cartilage is about to be replaced by bone, very striking changes in its structure take place. The
cells which have hitherto been secreting cartilage matrix begin to
destroy the matrix. The lacunae become enlarged and a curious arrangement of the cartilage cells becomes evident. The cells erode the
cartilage in such a manner that they become lined up in rows (Fig.
155). This process of destruction continues until the cartilage is extensively honeycombed. Meanwhile the tissue of the perichondrium overlying the area of cartilage erosion becomes exceedingly active. There is
rapid cell proliferation and the new cells, carrying blood vessels with
them, begin to invade the honeycombed cartilage (Fig. 155).
 
The Deposition of Bone. It is a striking fact that during its growth
cartilage is devoid of blood vessels, the nearest vessels to it being those
in the perichondrium. The invasion of cartilage by blood vessels
definitely determines its disintegration as cartilage, and at the same
time is the initial step in the formation of bone. For this reason the
enveloping layer of connective tissue, up to this time called perichondrium because of its relation to the cartilage, is now called
periosteum because of the relations it will directly acquire to the bone
about to be formed. This change will not be confusing if we stop to
think that both these terms are merely ones of relation, which translated mean, respectively, that tissue which surrounds cartilage, and that
tissue which surrounds bone. The important fact to bear in mind is that
this enveloping layer of tissue is of mesenchymal origin and therefore
contains cells of the stock that may develop into any of the connectivetissue family to which bone as well as cartilage belongs. When,
therefore, a mass of periosteal tissue {periosteal bud, Fig. 155) grows
into an area of honeycombed cartilage it carries in potentially boneforming cells. These cells come to lie along the strand-like remnants
of cartilage, just as in membrane bone formation osteoblasts ranged
themselves along fibrous strands. The actual deposition of bone proceeds in the same manner endochondrally as it does intramembranously. The only difference is that in one case a strand-like remnant of
cartilage serves as an axis for the trabecula, whereas in the other case
deposition begins on a fibrous strand. Extensions and fusions of the
growing trabeculae soon result in the establishment of typical cancellous bone similar to that formed intramembranously.
 
 
 
HISTOGENESIS OF BONE
 
 
281
 
 
The Formation of G)mpact Bone from Primary Cancellous
Bone. The difference between cancellous bone and compact bone is
architectural rather than histological. The fundamental composition
of the bone matrix, its lamellation, and the relations of the bone cells to
the matrix, are the same in both cases. It is the way in which the
lamellae are arranged that distinguishes these two types of bone from
each other. In cancellous bone the disposition of lamellae is such that
it leaves large marrow spaces between the trabeculae. In compact
bone there has been a secondary deposit of concentrically arranged
lamellae in the marrow spaces which greatly increases the density of
the bone as a whole.
 
The essential differences between the two, and the way in which
cancellous bone may become converted into compact bone, can be
illustrated by a simple schematic diagram. Figure 156, 1, shows the
arrangement of lamellae and marrow spaces in primary cancellous
bone. The osteoblasts which have formed the trabeculae still lie along
them on the surface toward the marrow cavity. If such an area is to
IxTome compact, these osteoblasts enter on a period of renewed
activity and deposit a series of concentric lamellae in the marrow
cavity. Frequently if the marrow spaces are irregular there is a
preliminary rounding out of them by local resorption of the bone
already formed (Fig. 156, 2). This is then followed by the deposition
 
 
 
Fig. 156. Diagram showing transformation of cancellous to compact bone.
The solid lines indicate the lamellae of primary cancellous bone; the dotted
lines show the subsequently added concentric (Haversian) lamellae which
nearly obliterate the marrow spaces of cancellous bone. The sequence of
events is indicated by the numbers. Note that irregularly shaped spaces in
the cancellous bone may be rounded out by absorption before the concentric
lamellae are laid down.
 
 
 
282 HISTOGENESIS OF BONE AND DEVELOPMENT OF SKELETAL SYSTEM
 
 
of the concentrically arranged lamellae, sometimes called Haversian
lamellae after the man who first described them in detail (Fig. 156, 3 ).
In this process the original marrow spaces are reduced to small canals
{Haversian canals^ into which have been crowded the blood vessels
which formerly lay in the marrow cavities (Fig. 156, 4 ). These canals
maintain intercommunication with each other in the substance of the
bone, constituting a network of pathways over which the bone receives
its vascular supply. As compared with the marrow spaces of cancellous
bone, however, they are very small; and the gross appearance of a
bone which has undergone this secondary deposit of concentric
lamellae amply justifies characterizing it as ‘'compact.’’
 
II. The Development of the Skeletal System
 
In dealing with the development of the skeletal system we must
recognize at the outset that the subject is far too extensive to be
covered here with anything like completeness. It is not difficult, however, to become acquainted with the outstanding features in the
development of two or three characteristic bones, as, for example:
the sequence of events in the formation of a flat bone; the steps involved in the establishment and growth of a long bone; the way
separate ossification centers appear in a common primordial cartilage
mass and give rise to the various parts of a vertebra. Familiarity with
such type processes gives one an understanding of the factors operative in the development of the skeleton as a whole and a background
sufficient to permit ready and intelligent following up of developmental details in specific bones in which one may become interested.
 
Development of Flat Bones. The flat bones, such as the bones
of the cranium and face, are for the most part of intramembranous
origin. We are, therefore, already familiar with the early steps in their
development from our study of the histogenesis of membrane bone
(Figs. 151 and 153). After a mass of primary cancellous bone has
been laid down in a configuration which suggests that of the adult
bone being formed, there appears about this mass a peripheral concentration of mesenchyme (Fig. 157, A). This periosteal concentration
of mesenchymal tissue contains potentially bone-forming cells which
soon become active and lay down a dense layer of parallel lamellae
about the spongy center of the growing bone (Fig. 157, B). Anatomically this dense peripheral portion is known as the outer table of the
bone. The inner portion, which in the flat bones usually remains
cancellous, is called the diploii. The original mesenchymal tissue which
 
 
 
THE DEVELOPMENT OF THE SKELETAL SYSTEM
 
 
283
 
 
Periosteum
Marrow space
 
l^one trabeculae
 
 
A
 
Subperiosteal
bone lamellae
 
 
Bone trabeculae
Marrow space
 
 
Periosteum
 
 
Fig. 157. Diagrams showing the manner in which the dense peripheral
layer of a flat bone is formed by the deposition of subperiosteal lamellae about
an area of primary cancellous bone.
 
I
 
remains in the marrow spaces of the diploe develops into characteristic “red bone marrow” rich in blood-forming elements (Fig. 152).
 
The story of the growth of the mandible, a membrane bone which
starts after the manner of flat bones but which later takes on a very
elaborate shape and finally becomes largely compact, can be gleaned
by a comparative study of figures 178, 180, and 184.
 
Development of Long Bones. The long bones are characteristically of endochondral origin. The cartilage in which they are
performed is a tempiorary miniature of the adult bone. Ordinarily
there are several ossification centers involved in the formation of long
bones. The first one to appiear is that in the shaft or diaphysis. The
location of this center is shown schematically in figure 158, A, Such
 
 
 
B
 
 
 
 
284 HISTOGENESIS OF BONE AND DEVELOPMENT OF SKELETAL SYSTEM
 
 
b
 
A
 
 
Fig. 158, Diagrams showing liie progress of ossification in a long bone.
The stippled areas represent cartilage; the black areas indicate bone.
 
A, Primary ossification center in shaft. B, Primary center plus shell of
subperiosteal bone. C, Entire shaft ossified. D, Ossification centers have
appeared in the epiphyses. E, Entire bone ossified except for the epiphyseal
cartilage plates.
 
details as the cartilage erosion which preceded its appearance and
the manner in which the deposit of bone was initiated have already
been considered (Fig. 155). Our interest now is in the relation of such
an endochondral ossification center to other centers, and to the bone
as a whole.
 
Almost coincidently with the beginning of bone formation within
the cartilage the overlying periosteum begins to add bone externally
(Fig. 158, B). In view of the fact that the bone-forming tissue carried
into the eroded cartilage arose from the periosteum, this activity of
the periosteum itself is not surprising. Moreover we have already
encountered this same phenomenon of periosteal bone formation in
the outer table of flat bones.
 
The formation of bone which starts at about the middle of the
shaft soon extends toward either end until the entire shaft is involved
(Fig. 158, C), leaving the two ends {epiphyses) still cartilage. Toward
the end of fetal life ossification centers appear in the epiphyses. The
number and location of these epiphyseal centers vary in different long
bones. There is always at least one center in each epiphysis and there
may be two or more. Not uncommonly there are two centers in one
epiphysis and one in the other, as illustrated in figure 158, D.
 
Between the bone fojcmed in the diaphysis and that formed in the
epiphysis there persists a mass of cartilage known as the epiphyseal
plaie which is of vital importance in the growth in length of the bone.
 
 
 
 
 
THE DEVELOPMENT OF THE SKELETAL SYSTEM
 
 
285
 
 
We should expect from the rigidity of bone matrix that interstitial
growth could not account for its increase in length. This was long ago
demonstrated experimentally by exposing a developing bone and
driving into it three small silver pegs, two in the shaft and one in the
epiphysis. The distance between the pegs being recorded, the incision
was closed and development allowed to proceed until a marked
increase had occurred in the length of the bone. On again exposing
the pegs, the two in the shaft were found to be exactly the same
distance apart as when they were driven in, but the distance between
the pins in the shaft and that in the epiphysis had increased by an
amount corresponding to the increase in length of the bone. This
indicates clearly that the epiphyseal plates constitute a sort of temporary, plastic union between the parts of the growing bone. Continued
increase in the length of the shaft is accomplished by the addition of
new bone at the cartilage plate. These epiphyseal plates persist during
the entire postnatal growth period. Only when the skeleton has
acquired its adult size do they finally become eroded and replaced
by bone which joins the epiphyses permanently to the diaphysis.
 
As the bone increases in length there is a corresponding increase
in its diameter. The manner in which this takes place is also susceptible of experimental demonstration. If madder leaves, or some of the
alizarin compounds extracted from them, be fed to a growing animal,
the bone formed during the time the feeding is continued is colored
red. If the madder is discontinued, bone of normal color is again
formed ; but the color still remains in the bone laid down while madder was being added to the diet. Thus it is possible, by keeping a record
of alternate f>eriods of feeding and withholding madder and comparing these records with the resulting zones of coloration in a bone, to
obtain very accurate information on the progress of bone growth and
resorption. Applied to the development of long bones this method
shows their increase in diameter to be due to continued appositional
growth beneath the periosteum. As the bone is added to peripherally
there is a corresponding resorption centrally. This central resorption
results in the formation of a cavity in the axis of the long bone which
is called the marrow canal (Fig. 158, C). With the further increase
in the diameter of a bone, its marrow canal becomes correspondingly
enlarged. A significant mechanical fact might be cited in this connection. 'Engineers have determined that the strongest rod which can be
made from a given weight of steel is obtained by molding it into
tubular form. The development of an essentially tubular shaft by
 
 
 
286 HISTOGENESIS OF BONE AND DEVELOPMENT OF SKELETAL SYSTEM
 
 
progressive increase in the size of the marrow cavity gjivcs a long bone
maximum strength with minimum weight.
 
The Formation of the Vertebrae. The development of the vertebrae is of interest to the student primarily because it exemplifies so
excellently a fundamental embryological phenomenon — the origin
of separate parts from an undifferentiated primordial tissue mass,
and the subsequent association of these parts to form an organized
structure. In studying young embryos we traced the history of the
mesodermic somites through their early differentiation. It will be
recalled that from the ventro-mesial face of each somite there arises
a group of mesenchymal cells called collectively a sclerotome (Fig.
42). These cells migrate from either side toward the mid-line and
become aggregated about the notochord. From these masses of cells
the entire vertebral column is destined to arise.
 
The first significant change which takes place in these primordial
masses is the clustering of sclerotomal cells derived in part from each
of the two adjacent somites into groups which are located opposite
the intervals between the myotomes. In studying series of transverse
sections this arrangement is easy to overlook unless the density of the
cells about the notochord is carefully noted in passing from section
to section. It shows very clearly, however, in frontal sections (Fig. 159).
Each of these cell clusters is the primordium of the centrum of a vertebra. Once formed they rapidly become more dense and more definitely
circumscribed (Fig. 160). Soon after the centrum takes shape, paired
mesenchymal concentrations extending dorsally and laterally from
the centrum establish the primordia of the neural arches and of the
ribs (Fig. 161).
 
 
 
Fig. 159. Semi-schematic coronal sections through the dorsal
region of young embryos to show how the vertebrae became intermyotomal in position. Note that the primordium of a centrum is
formed by cells originating from the sclerotomes of both the
adjacent pairs of somites.
 
 
 
THE DEVELOPMENT OF THE SKELETAL SYSTEM
 
 
287
 
 
 
Fig. 160. Transverse section from pig cm])ryo of 17 mm. cut at the level of the
lungs to show the structures in the dorsal body-wall. (After Minot.)
 
 
mantle layer ependymal
 
 
marginal layer
of eord
 
neural arch
 
 
of cord
 
 
layer
 
 
eympathetic ganghon
anterior cardinal vein
 
 
 
left left duct
erihre fetdium vt Ctteter
 
 
Fig. 161 . Transverse section of 20 mm. pig embryo cut at the level of the lungs
to show the developing vertebra and ribs. (After Minot.)
 
 
 
288 HISTOGENESIS OF BONE AND DEVELOPMENT OF SKELETAL SYSTEM
 
 
 
Fig. 162. Transverse section from 40 mm. pig embryo cut at the level of the
lungs to show the developing vertebra and ribs.
 
 
The Stage in w^hich the various parts of the vertebrae arc sketched
in mesenchymal concentrations, is frequently spoken of as the blastemal stage. It is rapidly followed by th^ cartilage stage. Conversion to
cartilage begins in the blastemal mass first in the region of the centrum
and then chondrification centers appear in each neural and each
costal process (Fig. 161). These spread rapidly until all the centers
fuse and the entire mass is involved (Fig. 162). The cartilage miniature of the vertebra thus formed is at first a single piece showing no
lines of demarcation where the original centers of cartilage formation
became confluent, and no foreshadowing of the separate parts of
which it will be made up after the cartilage has been replaced by
bone. Shortly before ossification begins the rib cartilage becomes
separated from the vertebra, but the vertebra itself remains in one
piece throughout the cartilage stage (Fig. 162).
 
The locations of the endochondral ossification centers which
appear in a vertebral cartilage are indicated schematically in figure
163. It readily can be seen how the spreading of these centers of bone
formation will establish the conditions which exist in an adult vertebra. The median ossification center gives rise to the centrum. The
centers in the neural processes extend dorsally to complete the neural
arch. The spinous process in most of the vertebrae is formed by a
prolongation of these same centers to meet dorsal to the neural canal.
 
 
 
 
Fig. 164. Diagram of four types of vertebrae indicating the parts derived
from the different ossification centers shown in figure 163. The part formed by
the median center in centrum is cdncentrically ringed; the parts arising from
the costal centers are stippled; parts derived from the lateral centers in the
neural arches are indicated in line-shading,
 
289
 
 
 
 
 
 
 
290 HISTOGENESIS OF BONE AND DEVELOPMENT OF SKELETAL SYSTEM
 
 
In forms such as the pig the spinous processes of tl>e more anterior
thoracic vertebrae are very long. In these vertebrae additional ossification centers appear in the spinous process and fuse with those in the
 
Scapula Humerus
 
 
 
Radius Mandible
 
& ulna
 
 
Fig. 165. Photograph (X 1/^ showing the
ossification centers which have appeared in pig
embryos of 35 mm. This and the two following
figures were made by photographing in transmitted light embryos in which all the uncalcified
tissues had been rendered transparent by treatment with potassium hydroxide and glycerine.
 
neural processes. The transverse processes with which the tubercles
of the ribs articulate are formed by the lateral extension of the
primary ossification centers in the neural processes. These same centers
extend ventrally also, and meet the centrum (cf. Figs. 163 and 164).
 
 
Scapula
/ Humerus
 
 
 
Fic. 166. Photograph (X IM) showing the progress of ossification in the
skeleton of a 65 mm. pig embryo.
 
 
 
Fig. 167. Photograph {X showing the extent of ossification in the skeleton of a 90 nun. pig embryo.
 
 
 
292 HISTOGENESIS OF BONE AND DEVELOPMENT OF SKELETAL SYSTEM
 
 
The shaft of the rib is formed by extension of its primary ossification center (Fig. 163). After birth, secondary epiphyseal centers
appear in the tubercle and head of the rib. These centers are separated from the shaft by persistent cartilage plates in the manner
described in discussing the development of long bones. Fusion of the
secondary epiphyseal centers with the shaft of the rib does not take
place until the skeleton has acquired its adult dimensions.
 
The foregoing discussion has been based on a thoracic vertebra in
which the relations of the rib to the vertebra show most clearly.
All the vertebrae have the costal element represented, although it is
greatly reduced and modified in other regions than the thoracic. A
study of figure 164, in which the components of vertebrae from the
cervical, thoracic, lumbar, and sacral regions are schematically
indicated, will make the homologies apparent. With these homologies
in mind it is sufficiently evident, without going into further detail,
how all these vertebrae arise by a process similar to that described
for the thoracic vertebrae.
 
The Progress of Ossification in the Skeleton as a Whole. It
 
would carry us beyond the scope of this book to take up the development of specific bones. Each has its own story involving the formation
of the connective tissue or the cartilage mass which precedes it ; local
erosion centers if it be preformed in cartilage; number, location, and
time of appearance of ossification centers; growth in length and
diameter; development of epiphyses; time of fusion of epiphyses and
diaphysis; and finally the development of muscle ridges and articular
facets. Without entering into a discussion of details of this sort, it is
possible nevertheless to follow the general progress of ossification in
the skeletal system as a whole. Embryos which have been treated with
potassium hydroxide and then cleared in glycerine clearly show the
various ossification centers. In such preparations the areas where
calcium salts have been deposited stand out white in reflected light
and opaque in transmitted light. ^Figures 165-167, which are photographs of preparations of this type, can be used to trace the history
of the more important bones. It should perhaps be stated explicitly
that these figures arc included primarily to give a general view of the
progress of ossification and secondarily to afford a readily available
source of reference for following up points of interest that may arise.
It is not a profitable use of the student’s time to attempt to memorize
the ossification centers which have appeared in embryos of any given
age.
 
 
 
==Chapter  13==
 
 
Tke Development of tke Face and Jaws
and tke Teetk
 
 
I. The Face and Jaws
 
The Stomodaeum. In studying the early development of the
digestive tract we saw that the primitive gut first appeared as a
cavity which was blind at both its anterior and posterior ends (Fig.
37). Its opening in the future oral region is established by the meeting
of an ectodermal depression, the stomodaeum, with the cephalically
growing anterior end of the gut. The stomodaeal depression, even
as late as the time the oral plate ruptures and establishes communication between the anterior end of the gut and the outside world, is very
shallow (Fig. 40). The deep oral cavity characteristic of the adult is
formed by the forward growth of structures about the margins of the
stomodaeum. Some idea of the extent of this forward growth can
be gained from the fact that the tonsillar region of the adult is at
about the level occupied by the stomodaeal plate when it ruptures.
The growth of the structures bordering the stomodaeum, then, not
only gives rise to the superficial parts of the face and jaws, but actually
builds out the walls of the oral cavity itself.
 
The Jaws. Because the face of a young embryo is pressed against
the thorax it is difficult to study unless the entire head is cut off and
mounted separately. Preparations of this kind observed under a dissecting microscope by strong reflected light show the surface configuration of the facial region very clearly. The most conspicuous landmarks
are the stomodaeal depression, which in view of its fate we may now
call the oral cavity, and the olfactory pits. In embryos as small as
7 mm. most of the structures which take part in the formation of the
face and jaws are already clearly distinguishable (Fig. 168). In the
mid-line cephalic to the oral cavity is a rounded overhanging prominence known as the Jrontal process. On either side of the frontal process
are horseshoe-shaped elevations surrounding the olfactory pits. The
 
293
 
 
 
294
 
 
DEVELOPMENT OF FACE, JAWS AND TEETH
 
 
median limbs of these elevations are known as the nap-medial processes
and the lateral limbs are called the naso-lateral processes.
 
Growing toward the mid-line from the cephalo-lateral angles of
the oral cavity are the maxillary processes. In lateral views of the
head (Figs. 31 and 32) it will be seen that the maxillary processes
and the mandibular arch merge with each other at the angles of the
mouth. Thus the structures which border the oral cavity cephalically
are: the unpaired frontal process in the mid-line, the paired nasal
processes on either side of the frontal, and the paired maxillary
 
 
 
Fig. 168. Face of 7 mm. pig embryo photographed
X 15. Note especially the unmistakably paired character of the thickenings which later fuse in the mid-line
to complete the mandibular arch.
 
 
processes at the extreme lateral angles. From these primitive tissue
masses the upper jaw and the nose are derived.
 
The caudal boundary of the oral cavity is less complex, being
constituted by the mandibular arch alone. In very young embryos
(Fig. 168) the origin of the mandibular arch from paired primordia
is still clearly evident. Appearing first on either side of the mid-line
are marked local thickenings due to the rapid proliferation of mesenchymal tissue. Until these thickenings have extended from either side
to meet in the mid-line there remains a conspicuous mesial notch.
With their fusion, the arch of the lower jaw is completed (Figs. 1 69172).
 
 
 
THE FACE AND JAWS
 
 
295
 
 
In 10-12 mm. embryos (Fig. 169) very marked progress can be
seen in the development of the facial region. The maxillary processes
are much more prominent and have grown toward the mid-line,
crowding the nasal processes closer to each other. The nasal processes
have grown so extensively that the frontal process between them is
completely overshadowed (cf. Figs. 168 and 169). The growth of the
 
 
 
Fig. 169. Face of 11 ,5 mm. pig embryo photographed
X 12. Fusion of the right and left components of the
mandibular arch is practically complete. Both the
medial and lateral limbs of the horseshoe-shaped nasal
processes have undergone conspicuous enlargement.
Note especially the approximation of each naso-medial
process to the maxillary process of the same side.
 
 
medial limbs of the nasal processes has been especially marked and
they appear almost in contact with the maxillary processes on either
side.
 
The groundwork for the formation of the upper jaw is now well
laid down. Its arch is completed by the fusion of the two nasomedial processes with each other in the mid-line, and with the
maxillary processes laterally (Fig. 170), The premaxillary bones
carrying the incisor teeth are formed, later, in the part of the upper
jaw which is of naso-medial origin. The maxillary bones, carrying all
 
 
 
296
 
 
DEVELOPMENT OF FACE, JAWS AND TEETH
 
 
the upper teeth posterior to the incisors, are developed in the part of
the arch arising from the maxillary processes.
 
Nasal Chambers. The olfactory pits have by this time become
much deepened, not only by the growth of the nasal processes about
them, but also by extension of the original pits themselves which soon
break through into the oral cavity (Figs. 93 and 97, C). We may now
 
 
lateral
process
 
naso*‘ medial
process
 
tojngue
 
hyomandibular
 
cleft
 
hyoid arch
 
 
Fig. 170. Face of 16 mm. pig embryo photographed X 10. The
naso-medial processes have fused with the maxillary processes on
either side, and with each other in the mid-line, thus completing
the arch of the upper jaw.
 
speak of the external openings of the nasal pits as the nostrils {external
nares) and their new openings into the oral cavity as posterior nares or
nasal choanae. The septum of the nose is formed by fusion in the midline of the original naso-medial processes'; the upper part of the
bridge of the nose is derived from the frontal process; and the alae
of the nose arise from the naso-lateral processes (Fig. 172).
 
Naao-Iacrimal Duct. Where the naso-lateral process and the
maxillary process meet each other there is formed iot a time a well
 
 
 
THE FACE AND JAWS
 
 
297
 
 
 
Fir;. 171. Face of 17.5 mm. pig embryo photographed X 10. The originally separate processes have now largely lost their identity in the series of
fusions which have taken place in the formation of the face.
 
 
marked groove, which extends to the mesial angle of the eye (Fig. 169).
This is known as the naso4acrimal groove. It soon closes over superficially (Fig. 171), and it is usually stated that the deep portion of the
original groove is converted into a tube, the naso-lacrimal duct^ or
tear duct, wjhich drains the fluid from the conjunctival sac of the eye
into the nose. Recently Politzer has maintained that the nasolacrimal duct arises as an independent epithelial downgrowth from
the conjunctival sac which follows closely along the line of closure of
the old naso-optic furrow.
 
Tongue. While these changes are going on externally, the tongue
is being formed in the floor of the mouth. Anatomically the tongue is
usually described as consisting of a freely movable part called its body,
and a less freely movable portion, called its root, by which it is
attached in the oro-pharyngeal floor. The body of the tongue arises
from a small median elevation, the tuberculum impar^ and paired lateral
lingual primordia. These elevations appear very early in development
on the inner face of the first branchial (mandibular) arch (Fig. 173,
B). The tuberculum impar grows slowly and is soon crowded in on
 
 
 
298
 
 
DEVELOPMENT OF FACE, JAWS AND TEETH
 
 
by the more rapidly growing lateral lingual primordia which form
the great bulk of the body of the tongue (Fig. 173, c5.
 
 
 
Fig. 172. Face of 21.5 rum. pig embryo
photographed X 10. The characteristic features
of the adult face are even at this early stage
clearly recognizable. The regions of the upper
jaw and nose which have arisen from originally
distinct primordia are differentiated by shading.
Vertical hatching indicates origin from frontal
process; stippling, from naso-lateral processes;
small crosses, from naso-medial processes; horizontal hatching, from maxillary processes. The
entire lower jaw is derived from the mandibular
arch.
 
 
Arising in the pharyngeal floor at the bases of the second and
third branchial arches is an elevation known as the copula (i.e., yoke)
 
 
THE FACE AND JAWS
 
 
299
 
 
because of the way it joins these arches together (Fig. 173, B). The
copula, supplemented by some tissue from the adjacent basal portions of branchial arches 2, 3, and 4, gives rise to the root of the
tongue.
 
All the various elevations which thus take part in the formation
 
 
- Branchial arch i
 
-Branchial arch 2
 
'Branchial arch 3
.Branchial arch 4
-Glottis
 
 
Lateral lingual anlagt
 
Copula
 
Epiglottis
 
Glottis
 
 
Branchial arch i
Lateral lingual anlage
 
Branchial arch $
 
Branchial arch 4
 
Arytenoid ridge
 
c
 
Fio. 173. Dissections of pig embryos made to expose the floor of the
mouth and show the development of the tongue. (After Prentiss.) A, 7 mm.;
B, 9 mm.; C, 13 mm. (All figures X 12.)
 
 
B
 
 
 
Tuherculum impof
 
Branchial arch 2
 
Epiglottis
 
Glottis
 
 
Lateral lingual anlage
Tuherculum impar
 
Epiglottis
Arytenoid ridge
 
 
Branchial arch i
Tuherculum impar
 
Branchial arch 2
 
Branchial arch 3
Branchial arch 4
 
Arytenoid ridge
 
 
 
300
 
 
DEVELOPMENT OF FACE, JAWS AND TEETH
 
 
 
THE FACE AND JAWS
 
 
301
 
 
of the tongue must be thought of as composed of an outer covering
and the underlying mesodermal tissue which causes the covering to
bulge into the lumen. The covering tissue arises in situ from the lining
of the branchial arches involved. The sensory innervation of the surface of the tongue is, therefore, just what one would expect from the
basic relations of the cranial nerves to the branchial arches. The
epithelium of the body of the tongue gets its sensory supply from the
lingual branch of the mandibular division of the trigeminal (V) nerve
(Fig. 97, A, B), and from the chorda tympani branch of the seventh
nerve. The root of the longue receives its sensory fibers from the glossopharyngeal (Fig. 94) and vagus nerves.
 
The skeletal muscle that makes up the main mass of the tongue
beneath the mucosal covering is derived from mesodermal cell masses
that are believed to migrate into the pharyngeal floor from the myotomes of the occipital somites. Ontogenetically, in mammalian embryos this migration is exceedingly difficult to trace, for the cells of
myotomal origin early mingle indistinguishably with the local mesenchymal cells. Nevertheless the way the hypoglossal nerve (XII),
which is the cranial nerve arising at the level of these occipital myotonies, grows in with the developing lingual muscles (Figs. 93, 94,
and 97, A, B) and innervates them, furnishes strong circumstantial
evidence for this interpretation of tongue muscle origin and migration.
 
Palate. Coincidently also the palatal shelf is being formed in the
upper jaw and separating off the more cephalic portion of the original
stomodaeal chamber. Since it is into this cephalic portion of the cavity
that the nasal pits break through (Figs. 93 and 97, C), the formation
of the palatal shelf in effect prolongs the nasal chambers backwards
so they open eventually into the region where the oral cavity becomes
continuous with the pharynx.
 
The palate as well as the arch of the upper jaw is contributed to
by both the naso-medial processes and the maxillary processes. From
the premaxillary region a small triangular median portion of the
palate is formed (Fig. 174). The main portion of the palate is derived
from the maxillary processes. From them shelf-like outgrowths arise
 
Fig. 1 74. Photographs (X 6) of dissections of pig embryos made to expose
the roof of the mouth and show the development of the palate. A, 20.5 mm.;
B, 25 mm.; C, 26.5 mm.; D, 29.5 mm.
 
The diagrams of transverse sections are set in to show the relations before
(E) and after (F) the retraction of the tongue from between the palatine
processes.
 
 
 
 
302
 
 
DEVELOPMENT OF FACE, JAWS AND TEETH
 
 
on either side and extend toward the mid-line (Fig. 174, A— G). When
these palatal shelves first start to develop, the tongue lies between
them (Fig. 174, E). As development progresses the tongue drops down
(Fig. 174, F); the palatal shelves are extended toward the mid -line
and finally fuse with each other medially and with the premaxillary
process anteriorly to complete the palate (Fig. 174, D). At the same
time the nasal septum grows toward the palate and becomes fused
to its cephalic face (Figs. 174, F, and 178). Thus the separation of
 
 
 
 
Fig. 175. Transverse section of snout of 28 mm. pig
embryo (X 12). The area included in the rectangle is
shown in detail in the following figure.
 
 
right and left nasal chambers from each other is accomplished at the
same time that the nasal region as a whole is separated from the oral.
 
II. The Development of the Teeth
 
The Dental Ledge. Local changes leading toward tooth formation can be made out in the jaws of embryos as small as 1 5 mm. or
even less. By the time a size of 28-30 mm. has been attained, a definite
thickening of the oral epithelium can readily be seen on both the
upper and the lower jaw. This band of epithelial cells which pushes
into the underlying mesenchyme around the entire arc of each jaw
is known as the labi<Hlental ledge {labio-dental lamina) (Figs. 175 and
176). Shortly after its first appearance, cross-sections show this ledge
 
 
 
THE DEVELOPMENT OF THE TEETH
 
 
303
 
 
of epithelial cells to be differentiating into two parts, a more distal
part which by its ingrowth marks off the elevation which is to become
 
 
 
Fig. 176. Drawing (X 130) showing labio-dental ledge of
28 mm. pig embryo. For location of area represented see preceding figure.
 
 
 
Fig. 177. Drawing (X 130) showing differentiation of the
labio-dental ledge into labio-gingival lamina and dental ledge.
 
The ingrowth of the labio-gingival lamina initiates the separation of the lip from the gum (gingiva). From the dental ledge a
series of local bud-like outgrowths are formed, each of which
gives rise to the enamel cap of a tooth.
 
The region shown is the same as that in the preceding figure
but from a slightly older (37 mm.) embryo.
 
the lip from that which is to become the gum, and a more proximal
part which is destined to grow into the gum and give rise to the
enamel-forming organs of the teeth. The part of the original labio
 
 
 
304
 
 
DEVELOPMENT OF FACE, JAWS AND TEETH
 
 
dental ledge which separates the lip from the gum (gingiva) is known
as the labio-gingival lamina^ and the part of the original ledge which is
to take part in tooth formation is known as the denial ledge or dental
lamina (Fig. 177).
 
Enamel Organs. Soon after the dental ledge is established, local
buds arise from it at each point where a tooth is destined to be formed.
Since these cell masses give rise to the enamel crown of the tooth they
are termed enamel organs. As would be expected, the enamel organs
 
 
nasal bone
 
 
cartilage of
 
nasal septum
 
 
nasal
 
chamber
 
 
nasal process
 
of premaxilla
 
 
 
dental papilla
enamel organ
 
 
labiogingival
 
lamina
 
Meckel’s cartilage
 
 
tooth germ
 
mandible
 
 
Fig. 178. Drawing (X 10)tDf a transverse .section of the snout
of a 71 mm. pig embryo. The area included in the rectangle is
shown in detail in the following figure.
 
 
for the milk teeth are budded off from the dental ledge first, but the
cell clusters which later give rise to the enamel of the permanent teeth
are formed at a surprisingly early time (Fig. 180). They remain
dormant, however, during the growth period of the milk teeth and
begin to develop actively only after the jaws have enlarged sufficiently
to accommodate the permanent dentition.
 
The histogenetic processes involved in the formation of milk teeth
and permanent teeth are essentially the same. It is, therefore, sufficient
to trace them in the case of the milk teeth only, keeping in mind that
the same process is repeated later in life in the formation of the permanent teeth.
 
In a section of the developing mandible which cuts the dental ledge
 
 
THE DEVELOPMENT OF THE TEETH
 
 
305
 
 
at a point where an enamel organ is being formed, the shape of the
enamel organ suggests that of an irregularly shaped, inverted goblet,
the section of the dental ledge appearing somewhat like a distorted
stem (Fig. 178). The epithelial cells lining the inside of the goblet early
take on a columnar shape. Because they constitute the layer which
secretes the enamel cap of the tooth, they are called ameloblasts (enamel
 
 
 
formers) (Fig. 179). The outer layer of the enamel organ is made up
of closely packed cells which are at first polyhedral in shape but which
soon, with the rapid growth of the enamel organ, become flattened.
They constitute the so-called outer epithelium of the enamel organ (Fig.
179). Between the outer epithelium and the ameloblast layer is a
loosely aggregated mass of ceils called collectively, because of their
characteristic appearance, the enamel pulp or the stellate reticulum
(Fig. 179).
 
 
306
 
 
DEVELOPMENT OF FACE, JAWS AND TEETH
 
 
 
Fig. 180. Developing tooth from lower jaw of a 120 mm. pig embryo
(X 14). The small sketch including half of the tongue (left) and part of the
lip (right) gives the relations of the region drawn. The area in the rectangle
is shown in detail in the following figure.
 
 
 
Fig, 181. Projection drawing (X 350) of segment of enamel organ and
adjacent pulp from a 120 mm. pig embryo to show ameloblast and odontoblast layers. For location of area represented see preceding figure.
 
 
 
THE DEVELOPMENT OF THE TEETH
 
 
307
 
 
The Dental Papilla. Inside the goblet-shaped enamel organ
there is caught a mass of mesenchymal cells which are said to constitute the dental papilla (Fig. 179). The cells of the dental papilla
proliferate rapidly and soon form a very dense aggregation. The outer
cells of this mass are destined to secrete the dentine of the tooth and
the inner cells to give rise to the pulp of the tooth.
 
A little later in development the enamel organ begins to assume
the shape characteristic of the crown of the tooth it is to lay down
(Fig. 180). At the same time the outer cells of the dental papilla
take on a columnar form similar to that of the ameloblasts (Fig. 181).
They are now called odontoblasts (dentine formers) because they are
about to become active in secreting the dentine of the tooth.
 
In the central portion of the dental papilla vessels and nerves arc
beginning to make their appearance so that the picture is already
suggestive of the condition seen in the pulp of an adult tooth. Meanwhile the growth of the dental papilla toward the gum has crowded
the stellate reticulum of the enamel organ in the crown region so it is
nearly obliterated (Fig. 180). This brings the ameloblasts of this region
much closer to the many small blood vessels which lie in the surrounding mesenchyme. The approach of the ameloblasts to the neighboring
vascular supply would appear to be significant, since it is precisely
here at the tip of the crown where the ameloblasts first begin to secrete
enamel (Fig. 182).
 
By the time the enamel organ has been well established the dental
ledge has lost its connection with the oral epithelium, although traces
of it can still be identified in the mesenchyme at the lingual side of the
tooth germ (Fig. 180). The cluster of cells which is destined to give
rise to the enamel organ of the permanent tooth of this level can be
seen budding off from the ledge close to the point from which the
enamel organ of ffie milk tooth arose (Figi >^).
 
Formation of Dentine. With these preparatory developments
complete, the tooth-forming structures are, so to speak, ready to go
about the fabrication of dentine and enamel. As is the case with bone,
enamel and dentine are both composed of an organic basis in which
inorganic compounds are deposited. We may use the same comparison
that was used in describing bone : that of the familiar use in construction operations of a steel meshwork into which concrete is poured, the
steel giving the finished structure some degree of elasticity and increasing the tensile strength while the concrete gives body and solidity.
In the case of such hard structures in the body as bone, dentine, and
 
 
 
308
 
 
DEVELOPMENT OF FACE, JAWS AND lEETH
 
 
pr^ordiutn enamel organ
' permanent tooth
 
 
odontoblast layer
pulp of tooth
 
 
 
enamel pulp (stellate reticulum)
periosteum of alveolar socket
outer epithelium of enamel organ
 
 
Fig. 182. Developing tooth from lower jaw of a 130 mm. pig embryo
(X 30). The small sketch gives the relations of the regions drawn. The area
in the rectangle is shown in detail in the following figure.
 
 
blood vessel dentine enamel blood vessel in
 
1 i, 1 mesenchyme
 
 
 
fiber lirocess of enamel organ
 
 
Fio, 183. Projection drawing (X 350) of small segment of developing
 
 
THE DEVELOPMENT OF THE TEETH
 
 
309
 
 
enamel, these interlacing organic strands in the matrix give the tissue
its resilience and tensile strength, and the calcareous compounds
deposited in the organic framework give form and hardness.
 
Although bone, dentine, and enamel are similar in having both
organic and inorganic constituents in their matrix they are quite
different in detail, both as to composition and microscopical structure.
Bone has approximately 45 per cent of organic material while dentine
has but 30 per cent and adult enamel 5 per cent or less. There is also
considerable difference in the kind and proportion of inorganic compounds present in each. Structurally they are totally unlike. Bone
matrix is formed in lamellae and has cells scattered through it.
Dentine is formed without lam^llation and has its cellular elements
lying against one face and sending long processes into tubules in the
matrix. Enamel is prismatic in structure and the cells which form
it lie against its outer surface while it is being deposited, but are
destroyed in the eruption of the tooth.
 
The first dentine is deposited against the inner face of the enamel
organ, the odontoblasts drawing their raw materials from the small
vessels in the pulp and secreting their finished product toward the
enamel organ. It is significant in this connection that in an active
odontoblast the nucleus, which is the metabolic center of the cell, has
gravitated toward the source of supplies and come to lie in the extreme
pulpal end of the cell (Fig, 183). Also, the end of the odontoblast
toward the enamel organ, where the elaborated product of the cell
is being accumulated preparatory to its extrusion, can be seen to
take the stain especially intensely. Although our knowledge of intracellular chemistry is as yet exceedingly fragmentary and we do not
know the exact chemical nature of the product in this stage, the staining reaction is clearly indicative of the presence of calcium compounds
of some sort.
 
If attention is turned now to the recently formed dentine, two
zones distinctly different in staining reaction can be seen. The zone
nearer the cells is pale, taking but little stain (Fig. 183). This zone
consists of the recently deposited organic part of the matrix not as yet
impregnated with calcareous material. The zone nearer the enamel
organ will be found, by contrast, very intensely stained. This is the
older part of the dentine matrix which has had the organic framework
impregnated with calcareous material.
 
As the odontoblasts continue to secrete additional dentine matrix
the accumulation of their own product inevitably forces the cell
 
 
 
310
 
 
DEVELOPMENT OF FACE, JAWS AND TEETH
 
 
layer back, away from the material previously deposited. Apparently
strands of their cytoplasm become embedded in the material first laid
down and are then pulled out to form the characteristic processes of
the odontoblasts known as the dentinal fibers (Fig. 183). As the layer
of secreted material becomes thicker and the cells are forced farther
from the material first deposited, these dentinal fibers become progressively longer. Even in adult teeth where the dentine may be as much
as 2 mm. in thickness they extend from the odontoblasts which line
the pulp chamber to the very outer part of the dentine. These dentinal
fibers are believed to be concerned with maintaining the organic
portion of the dentine matrix in a healthy condition. When the pulp
is removed from a tooth, taking with it the odontoblasts, we know
that the dentine undergoes degenerative changes which involve,
among other things, increase in brittleness. This would seem to be
attributable to the degeneration of the organic framework of a matrix
no longer nourished by the odontoblasts.
 
Formation of Enamel. While the dentine is being laid down by
the cells of the odontoblast layer, the enamel cap of the tooth is being
formed by the ameloblast layer of the enamel organ. As was the case
with the odontoblasts, the active cells of the ameloblast layer are
columnar in shape and their nuclei, too, lie in the ends of the cells
toward the source of supplies, in this case the small vessels in the
adjacent mesenchyme (Fig. 183). The amount of organic material
laid down as the framework of enamel is much less than is the case
with either bone or dentine, and it is therefore more difficult to make
out its precise character and arrangement. It is, nevertheless, possible
to see in decalcified sections, delicate fibrous strands projecting from
the tips of the ameloblasts into the areas of newly formed enamel
(Fig. 183). It seems probable that these strands {Tomes'' processes) are
in some way involved in the formation of the organic matrix of
enamel. The problem of tracing the relations of Tomes’ processes to
the organic framework of enamel is greatly complicated by the fact
that where the ameloblasts have deposited calcium compounds the
calcium has rendered the organic part of the matrix so avid in its
affinity for stains that it is not possible to discern fine structural details
because of the very density of the resulting coloration (Fig. 183). This
reaction of the tissue to stains persists even after the inorganic calcium
compounds have been removed by decalcification, indicating that
the organic framework itself has been chemically altered by the
calcium deposited in it.
 
 
 
THE DEVELOPMENT OF THE TEETH
 
 
311
 
 
In spite -of these difficulties in getting at the exact nature and
arrangement of the organic matrix of enamel, it is quite possible to
see the genesis of its fundamental prismatic structure. Each amcloblasl
builds up beneath itself a minute rod or prism of calcareous material.
These prisms are placed with their long axes approximately at right
angles to the dento-enamel junction. Collectively these enamel prisms
form an exceedingly hard cap over the crown of the tooth which in
its structural arrangement suggests a paving of polygonal bricks laid
on end. There is sufficient difference in the rate at which the different
 
 
growth lines
 
in enamel
 
 
pulp chamber
 
growth lines
in dentine
 
 
root canal
 
 
cementum
 
 
oral epithelium
 
 
osteolilasts
 
of periosteum
of alveolus
 
 
connective
tissue ftbers
 
 
cementoblasts
( from
dental sac)
 
 
blood vessels
and nerves to pulp
 
 
Fig. 184. Schematic diagram showing the topography of a tooth and its
relations to the bone of the jaw. The numbered zones indicate empirically
the sequence of deposition of the dentine and enamel. The so-called growth
lines in the dentine and enamel follow the general contours indicated by
the dotted lines in the figure but arc much more numerous.
 
 
 
312
 
 
DEVELOPMENT OF FACE, JAWS AND TEETH
 
 
ameloblasts work so that in actively growing enamel the surface is
jagged and irregular due to the varying extent to ^<^hich the different
prismatic elements have been calcified (Fig. 183).
 
Both enamel formation and dentine formation begin at the tip
of the crown and progress toward the root of the tooth (Figs. 182 and
184). But the entire crown is well formed before the root is much more
than begun. The progressive increase in the length of the root is an
important factor in the eruption of the tooth, for as the root increases
in length the previously formed crown must move closer to the surface
of the gum. Even when the crown of the tooth begins to erupt the
root is still incomplete, and it does not acquire its full length until the
crown has entirely emerged.
 
The Formation of Cementum. The so-called cementum of the
tooth is virtually a bone encrustation of its root. No cementum is
formed until the tooth has acquired nearly its full growth and its
definitive position in the jaw. But the first indications of specialization
in the tissue destined to give rise to it can be seen long before the
cementum itself appears.
 
Outside the entire tooth germ, between it and the developing bone
of the jaw, there occurs a definite concentration of mesenchyme.
The concentration becomes evident first at the base of the dental
papilla and extends thence crownwards about the developing tooth,
which it eventually completely surrounds.
 
This mesenchymal investment is known as the dental sac (Fig. 182).
In the eruption of the tooth the portion of the dental sac over the
crown is destroyed, but the deeper portion of the sac persists and
becomes closely applied to the growing root. At about the time the
tooth has acquired its final position in the jaw, the cells of the dental
sac begin to form the cementum. Histologically and chemically
cementum is practically identical with subperiosteal bone. When we
consider the manner of origin of the dental sac and of the periosteum
of the bone socket (alveolar socket) in which the root of the tooth lies,
and see how they arise side by side from the same sort of tissue, this
seems biit natural. The dental sac is essentially a layer of periosteal
tissue facing the root of the tooth and back to back with the periosteal
tissue of the alveolar socket (Fig. 182).
 
The Attachment of Tooth in the Jaw. The attachment of the
tooth in its socket is brought about by the development, between the
dental sac and the periosteum of the tooth socket, of an exceedingly
tough fibrous connective tissue. As the periosteum of the alveolus
 
 
 
THE DEVELOPMENT OF THE TEETH
 
 
313
 
 
adds new lamellae of bone to the jaw on the one side, and the dental
sac adds lamellae of cementum to the root of the tooth on the other,
the fibers of this connective tissue are caught in the new lamellae.
Thus the tooth comes to be supported by fibers which are literally
calcified into the cementum of the tooth at one end and into the bone
of the jaw at the other (Fig. 184). The mechanism involved is precisely
the same as that which occurs in the burying of tendon fibers in a
growing bone, where the buried ends of the fibers are known as the
penetrating fibers of Sharpey.
 
Replacement of Deciduous Teeth by Permanent Teeth. The
 
replacement of the temporary or “milk” (deciduous) dentition by the
permanent teeth is a process which varies in detail for each tooth.
The general course of events is, however, essentially similar in all
cases. The enamel organ of the permanent tooth arises from the
dental ledge near the point of origin of the corresponding deciduous
tooth (Fig. 180). With the disappearance of the dental ledge, the
permanent tooth germ comes to lie in a depression of the alveolar
socket on the lingual side of the developing deciduous tooth (Fig. 185).
 
When the jaws approach their adult size the hitherto latent primordia of the permanent teeth b<-gin to go through the same histo
 
 
Fio. 185. Photomicrograph (X 5) of upper jaw of 160 mm. pig embryo
showing the milk cuspids just breaking through the gum.
 
 
314
 
 
DEVELOPMENT OF FACE, JAWS AND TEETH
 
 
 
Fig. 186. Photomicrograph (X 6) of section through the jaw of a puppy
showing a deciduous tooth nearly ready to drop out and the developing
permanent tooth deeply embedded in the jaw below it. The space about the
crown of the permanent tooth was occupied in the living condition by
enamel. Fully formed enamel, being approximately 97 per cent inorganic
in composition, is almost completely destroyed by the decalcification with
acids which must be carried out before such material can be sectioned.
(From a preparation loaned by Dr* S. W. Chase.)
 
 
THE DEVELOPMENT OF THE TEETH
 
 
315
 
 
genetic changes we have already traced in the case of the temporary
teeth. As the permanent tooth increases in size, the root of the deciduous tooth is resorbed and the permanent tooth comes to lie underneath
its remaining portion (Fig. 186). Eventually nearly the entire root
of the deciduous tooth- is destroyed and its loosened crown drops out,
making way for the eruption of the corresponding permanent tooth.
 
 
 
 
BiLlio grapky
 
 
Although this docs not purport to be a complete bibliography
on the development of the pig, I have tried to make it comprehensive.
At least one reference has been included on every phase of the subject
concerning which I could find published information. Under each
of the main subject headings some of the articles referred to have
extensive bibliographies of the literature in their special field. But,
with the exception of a few outstanding contributions, no papers have
been included which are merely of historical interest. For such articles
reference should be made to the exhaustive bibliographies compiled
by Minot (1893) and by Keibel (1897). Furthermore, the list is largely
restricted to contributions based directly on pig embryos. Exception
to this rule has been made in favor of a few general articles which
are of especial assistance in acquiring a perspective on some phase of
the subject. Also, a number of articles based on other forms have been
included when no work appeared to have been done on corresponding
phases of development in the pig. It is hoped that such a list of
selected references will furnish a starting point for following up any
desired line of inquiry without involving one in a discouraging
multiplicity of titles.
 
Texts and Manuals
 
Arey^ L. i?., 19^6. Developmental Anatomy. Saunders, Philadelphia, 5th Ed., ix &
 
616 pp.
 
Baumgartner, W, J,, 1924, Laboratory Manual of the Foetal Pig, Macmillan, New
York, vii & 57 pp.
 
Boyden, E. A., 1936. A Laboratory Atlas of the 13-mm. Pig Embryo. (Prefaced by
younger stages of the chick embryo.) The Wistar Institute Press, Philadelphia,
iv & 104 pp.
 
Hamilton, W. J., Boyd, J. D., and Mossman, H. W., 1945. Human Embryology.
 
Williams and Wilkins, Baltimore, viii & 366 pp.
 
Hertwig, O., 1901-07. Handbuch der Vergleichenden und experimentellen Entwicklungslehre der Wirbeltiere. (Edited by Dr. Oskar Hertwig and written by
numerous collaborators.) Fischer, Jena.
 
Huettner, A, F., 1941. Fundamentals of Comparative Embryology of the Vertebrates.
Macmillan, New York, xiv & 416 pp,
 
Jordan, H. E,, and Kindred, J. E., 1948. A Textbook of Embryology. Appleton, New
York, 5th Ed., xiv & 613 pp.
 
 
317
 
 
 
 
318
 
 
BIBLIOGRAPHY
 
 
Keihel^ F., 1897. Normentafeln zur Entwicklungsgeschichte der Wirbelthiere. I. Des
Schweines. Fischer, Jena, 114 S.
 
Martin^ P., 1912. Lehrbuch der Anatomic der Haustiere. ^chickhard & Ebner,
Stuttgart. (Bd. 1, Allgemeine und vergleichende Anatomic mit Entwicklungsgeschichte, xii & 811 S.)
 
Minot ^ C, S., 1893. A Bibliography of Vertebrate Embryology. Memoirs, Boston
Soc. Nat. History, Vol. 4, pp. 487-614.
 
Mtnot, C, S., 1911. A Laboratory Textbook of Embryology. The Blakiston Company,
Philadelphia, 2nd Ed., xii & 402 pp.
 
Needham^ J'., 1931. Chemical Embryology. Macmillan, New York, xxii & 2021 pp.
 
Patten^ B. A/., 1929. Early Embryology of the Chick. The Blakiston Company, Philadelphia, 3rd Ed., xiii & 228 pp.
 
Patten, B. M., 1946. Human Embryology. The Blakiston Company, Philadelphia,
XV & 776 pp.
 
Stssori, S., 1921. The Anatomy of the Domestic Animals. Saunders, Philadelphia,
2nd Ed., 930 pp.
 
Wetss, P.y 1939. Principles of Development. A Text in Experimental Embryology.
Holt, New York, xix & 601 pp.
 
IVieman, H. L., 1930. An Introduction to Vertebrate Embryology. McGraw-Hill,
New York, xi & 411 pp,
 
Windle, W, F., 1940. Physiology of the Fetus. Saunders, Philadelphia, xiii & 249 pp.
 
Zeitzschmann, 0,, 1923-24. Lehrbuch dcr Entwicklungsgeschichte der Haustiere.
R. Schoetz, Berlin, 542 S,
 
General Articles and Articles of Significance for the
Interpretation of Developmental Processes
 
Alexander, J., 1944. The Gene — A Structure of Colloidal Dimensions. Chapt. 37,
pp. 808-819, in “Colloid Chemistry,” Vol. V, edited by Jerome Alexander,
Reinhold Publishing Corp., New York.
 
Allen, W. F., 1918. Advantages of sagittal sections of pig embryos for a medical
embryology course. Anat. Rec., Vol. 14, pp. 183-191.
 
Barth, L. G., 1944. Colloid Chemistry in Embryonic Development. Chapt. 39, pp.
851-859, in “Colloid Chemistry,” Vol. V, edited by Jerome Alexander, Reinhold Publishing Corp., New York.
 
Chambers, R., 1944. Some Physical Properties of Protoplasm. Chapt. 41, pp. 864-875,
in “Colloid Chemistry,” Vol. V, edited by Jerome Alexander, Reinhold
Publishing Corp., New York.
 
Conklin, E, G., 1914. The cellular basis of heredity and development. Pop. Sci.
Monthly, Vol. 85, pp. 105-133. '
 
Hartman, C. G., 1 931 . Development of the egg as seen by the physiologist. Sci. Monthly,
Vol, 33, pp, 17-28.
 
Medley, 0. F., 1926. Quantitative study of growth of certain organs in pig fetus. Bull.
Med. Coll. Va., Vol. 23, pp. 19-36.
 
Holtfreter, J., 1947. Changes of structure and the kinetics of differentiating embryonic
cells. Jour. Morph., Vol. 80, pp. 57-92.
 
Holtfreter, J., 1947. Observations on the migration, aggregation and phagocytosis of
embryonic cells. Jour. Morph., Vol. 80, pp. 93-111.
 
 
 
THE SEX ORGANS AND GAMETOGENESIS
 


319
[[Category:Draft]]
 
 
Kingsbury, B. F,, 1926. On the so-called law of anteroposterior development. Anat.
Rec., Vol. 33, pp. 73-87.
 
Lewis, W, H., 1947. Mechanics of invagination. Anat. Rec., Vol. 97, pp. 139-156.
 
Spemann, H., 1927. Organizers in animal development. Proc. Roy. Soc. London,
Ser. B, Vol. 102, pp. 177-187.
 
Spemann, H., 1938. Embryonic Development and Induction. Yale Univ. Press,
New Haven, xii & 401 pp.
 
The Sex Organs and Gametogenesis
 
Allen, E., 1923. Ovogenesis during sexual maturity. Am. Jour. Anat., Vol. 31, pp.
439-481.
 
Allen, E., Kountz, IV. B., and Francis, B. F., 1925. Selective elimination of ova in the
adult ovary. Am. Jour. Anat., Vol. 34, pp. 445-468.
 
Bascom, K. F., 1925. Quantitative studies of the testis. I. Some observations on the
cryptorchid testes of sheep and swine. Anat. Rec., Vol. 30, pp. 225-241.
 
Bascom, K. F., and Ouerud, H. L., 1925. Quantitative studies of the testicle. II. Pattern
and total tubule length in the testicles of certain common mammals. Anat. Rec.,
Vol. 31, pp. 159-169.
 
Blandau, R. J., 1945. The first maturation division of the rat ovum. Anat. Rec.,
Vol. 92, pp. 449-457.
 
Corner, G. W., 1917. Maturation of the ovum in swine. Anat. Rec., Vol. 13, pp. 109
112 .
 
Comer, G. IV., 1919. On the origin of the corpus luteum of the sow from both granulosa and theca interna. Am. Jour. Anat., Vol. 26, pp. 117-183.
 
Evans, H. M., and Swezv, 0., 1929. Ovogenesis and the normal follicular cycle in adult
mammalia. Mem. LTniv. Cal., Vol. 9, pp. 119-224.
 
Everett, N. B., 1945. The present status of the germ-cell problem in vertebrates. Biol.
Rev., Vol. 20, pp. 45-55.
 
Gould, H. N., 1923. Observations on the genital organs of a sex intergrade hog. Anat.
Rec., Vol. 26, pp, 241-261.
 
Hargitt, G. T., 1925-30, The formation of the sex glands and germ cells of mammals.
Jour. Morph. & Physiol., Vols. 40, 41, 42, 49.
 
Hartman, C. G., 1926. Polynuclear ova and polyovular follicles in the opossum and
other mammals, with special reference to the problem of fecundity. Am. Jour.
Anat., Vol. 37, pp. 1-52.
 
Hill, R. T., Allen, E., and Kramer, T. C., 1935. Cinemicrographic studies of rabbit
ovulation. Anat. Rec., Vol. 63, pp. 239-245.
 
Kellicott, W. E,, 1913. A Textbook of General Embryology. Holt, New York, v &
376 pp.
 
Kiipfer, M., 1920. Beitrage zur Morphologic der weiblichen Geschlechtsorgane bei
den Saugetieren. Ueber das Auftreten gelber Korper am Ovarium des domestizierten Rindes und Schweines. Vierteljahrsschrift d. Naturf. Gesellsch., Zurich,
Bd. 65, S. 377-433.
 
Latta, J, S., and Pederson, E. S,, 1944. The origin of ova and follicle cells from the
germinal epithelium of the ovary of the albino rat as demonstrated by selective
intravital staining with India ink. Anat. Rec., Vol. 90, pp. 23-35,
 
 
 
320
 
 
BIBLIOGRAPHY
 
 
Morgan^ T. H.j 1926. The Theory of the (iene. Yale Univ. Press, New Haven, xvi &
343 pp.
 
Fainter, T. S,, 1922. Studies in mammalian spermatogenesis. J(/Ur. Exp. Zodl., Vols.
35, 37, 39. Jour. Morph. & Physiol., Vol. 43.
 
Parker, G. H., and Bullard, C., 1913. On the size of litters and the number of nipples
in swine. Proc, Am. Acad. Arts & Sciences, Vol. 49, pp. 399-426.
 
Patten, W., 1925. Life, evolution and heredity. Sci. Monthly, Vol. 21, pp. 122-134.
 
Pincus, G., and Enzrnann, K. V,, 1937. The growth, maturation and atresia of ovarian
eggs in the rabbit. Jour. Morph., Vol. 61, pp. 351-383.
 
Pltske, E. C., 1940. Studies on the influence of the zona pellucida in atresia. Jour.
Morph., Vol. 67, pp 321-349.
 
Schmaltz-, B., 1911. Die Slrucktur der Ge.schlechtsorgane der ITaussaugetiere. P. Parey,
Berlin, xii & 388 S.
 
Smith, J, T., and Kettcrmgharn, R. C., 1937-38. Rupture of the graafian follicles.
Part I. Am. Jour. Obs. & Gyn., Vol. 33, pp. 820-827. Part IT Am. Join. Obs.
& Gyn., Vol. 36, pp. 45.3-460.
 
Stein, K. F., and Allen, E., 1942. Attempts to stimulate proliferaticm of the germinal
epithelium of the ovary. Anat. Rec., Vol. 82, pp. 1 -9.
 
Thanhoffer, L. de, 1934. The structure of the graafian follicle as revealed by microdissection. Zeilschr. f. Anat. u. Entwg., Bd. 102, S. 402-408.
 
Warwick, B. L., 1925. The eflect of vasectomy on swine. Anat. Rec., Vol. 31, pp.
19-21.
 
Wihon, E. B., 1925. The Cell in Development and Heredity. Macmillan, New York,
3rd Ed., ix & 1232 pp.
 
Young, W. C , 1929. A study of the function of the epididymis. I. Is the attainment
of full spermatcjzoon maturity attributable to some specific action of the epididymal secretion? Jour. Morph. & Physiol., Vol. 47, pp. 479-495.
 
Young, W. C., 1929. A study of the function of the epididymis. II. The importance of
an aging process in sperm for the length of the period during which fertilizing
capacity is retained by sperm isolated in the epididymis of the guinea-pig. Jour.
Morph. & Physiol., Vol. 48, pp. 475-491.
 
Young, W. C., 1931. A study of the function cT the epididymis. III. Functional changes
undergone by spermatozoa during their passage through the epididvmis and
vas deferens in the guinea-pig. Jour. Exp. Biol., Vol. 8, pp. 151-162.
 
The Sexual Cycle, Fertilization, Sex Determination
 
Allen, E., 1926. The ovarian follicular hormone: a study of variation in pig, cow, and
human ovaries. Proc. Soc. Exp. Biol. & Med., Vol. 23, pp. 383-387.
 
Allen, E,, Danforth, C. H., and Daisy, ErA,, 1939. Sex and Internal Secretions. Williams
and Wilkins, Baltimore, 2nd Ed., xxxvi & 1346 pp.
 
Allen, E., and Daisy, E. A., 1927. Ovarian and placental hormones. Physiol. Reviews,
Vol. 7, pp. 600-650.
 
Ampolsky, Z)., 1928. Cyclic changes in size of muscle fibers of the fallopian tube of
the sow. Am. Jour. Anat., Vol. 40, pp. 459-469.
 
Blandau, R., and Money, W. L., 1944. Observations on the rate of transport of spermatozoa in the female genital tract of the rat. Anat. Rec., Vol. 90, pp. 25.5-260.
 
 
 
THE SEXUAI, CYCLE, FERTILIZATION, SEX DETERMINATION 321
 
 
Burns, R. K., Jr., 1938. Hormonal control of sex differentiation. Am. Nat., Vol. 72,
pp. 207-227.
 
Comstock, R. E., 1939. A study of the mammalian sperm cell. I. Variations in the
glycolytic power of spermatozoa and their relation to motility and its duration.
Jour. Exp. Zool., Vol. 81, pp. 147—164.
 
Corner, G. W., 1915. The corpus luteum of pregnancy as it is in swine. Carnegie Inst.,
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Corner, G. W., 1917. Variations in the amount of phosphatids in the corpus luteum of
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Cleavage, Germ Layer Formation^ and the Structure of Young
 
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THE NERVOUS SYSTEM AND SENSE ORGANS
 
 
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The Nervous System and Sense Organs
 
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THE CIRCULATORY SYSTEM
 
 
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^ The Circulatory System
 
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KastschenkOyN.y 1887. Das Schicksal der embryonalen Schlundspalten bci Saugelhieren
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Minervini, R., 1904. Des ^Capsules surrenalcs. Developpement, structure, fonctions.
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Mitsukuriy K., 1882. On the development of the suprarenal in mammalia. Quart.
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Moody, R, 0., 1910. Some features of the histogenesis of the thyreoid gland in the
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CONNECTIVE TISSUES, SKELETAL, AND MUSCULAR SYSTEMS 337
 
 
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Connective Tissues, Skeletal, and Muscular Systems
 
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338
 
 
BIBLIOGRAPHY
 
 
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TWINS, DOUBLE MONSTERS, ANOMALIES
 
 
339
 
 
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Twins, Double Monsters, Anomalies
 
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340
 
 
BIBLIOGRAPHY
 
 
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42, pp. 267-280.
 
Poklman, A, G., 1919. Double ureters in human and pig embryos. Anat. Rec., Vol.
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183-194.
 
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Index
 
 
To facilitate the use of this book in connection with otheis in which the terminology
may differ somewhat, many synonyms which were not used in the text have been put into
the index and cross-referenced to the alternative terms used in this book; for example,
Wolffian body, a teim not used in this text, is frequently applied to the mesonephros. It
appears m the index thus: Wolffian body (= mesonephros, q.v.).
 
Both figure and page references are given in the index. The figure references arc preceded by the letter /.
 
 
Abdominal pregnancy, 217
Abducens nerve, 169
Accessory nerve, 172
Acoustic ganglion, /92, 117, 169
Acoustico-facialis ganglion ( = early undifferentiated condition of ganglia of
7th and 8th cranial nerves), 169
Action system, 141
 
Adrenal, /1 00, /106, /1 27, /1 28, /1 38, 223
After-birth, 105
Alae of nose, 296
Alar plate of neural tube, 1 57
Allantois, circulation of, /45,/51, 99
formation of,/30,/37, 99
function of, 106
relations of,/49,/55, 103
Alveolar periosteum, 312
Ameloblasts,/179-183, 305, 310
Amnion, formation of,/25,/37,/50, 97
function ofi 96
relations of,/49, 97
Amniotes, 96
Amniotic folds, 97
Ampulla of ductus deferens, /3, 8
Anal plate ( = cloacal plate, q.v.)
Anamniotes, 96
Angioblast, 87 footnote
Animad pole, 37
 
Anterior intestinal portal, /37, 74
Anterior neuropore, /29, 71
Anus,/118,/132, 210, 225
Aorta, see arteries
Aortic arches, sec arteries
Aortic bulb, 256
Aortic chromaffin body, 223
Appendage buds, /3 1-34, 65, 109
 
 
Appendix, of epididymis, /1 24, 213
of testis, /1 24, 217
Appositional growth, 278
Aqueduct of Sylvius, 158
Archenteron, 43
Areola, 105
 
Arterial circle (of Willis), 131
Arteries, allantoic, /45,/51, 89, 132, 240
aorta, dorsal, /67, /1 33-1 37, 88, 131,
235, 240
 
aoita, ventral, /45, /1 33, 88, 235
aortic arches, /45, /1 33-1 37, 88, 129,
 
233
 
basilar, /66, /67, /133, /137, 131, 238
brarhio-cephalic,/133, 234
carotid, common, /1 33, /1 37, 234
carotid, external, /67, /1 33, /1 37, 130,
 
234
 
carotid, internal, /66, /133, /137, 130,
234, 238
caudal, 240
 
cervical, intersegmental,/66,/133,/l37,
130, 235
 
coeIiac,/67,/lll, 132, 240
ductus arteriosus /1 33, /ISO, 235, 262,
268
 
hypogastric, 240, 270
iliacs,/66, /150, 240
innominate, 234
internal mammary, 238
interscgmcntal,/67,/133, 130, 235
mesenteric, ant., /67, /1 11, 132, 238
mesenteric, post»,/lll, 238
omphalomesenteric, /45, 89, 131, 238
pulmonary, /67, /1 33-1 37, 130, 234,
267
 
 
341
 
 
 
342
 
 
INDEX
 
 
Arteries — {Continued)
 
renal, /1 27, /1 28, 240
speimatic, /1 27-1 30
 
subclavian, /67, /133-135, 131, 234,
235
 
umbilical (allantoic), /66, /1 50, 89, 132,
240
 
vertebral,/66,/67,/137, 131, 237
vitelline, /45, 89, 132
Arytenoid process of larynx, /1 38
Astrocytes, 150
Atresia of ovarian follicles, 20
Atrio-ventricular canal, 126, 256
Atrium, 126, 254
 
Auditory, ganglion, see acoustic nerve, 1 69
vesicle, /36,/61, 117, 169
 
Basal plate of neural tube,/91, 157
Belly-stalk, /49, 97
Bicornate uterus, 216
Bile duct, common, 185
Bladder (urinary), /118, /124, /125, /130,
209
 
Blastocoele, j\S, 41
Blastocyst, /1 7, 41
 
elongation of, /1 8, 45
riastodermic vesicle ( == blastocyst, q.v.)
Blastodisc ( — embryonic disc, q.v.)
Blastorneie, 38
Blastula, 41
 
Blood cells, formation of,/48,/152, 90, 251
Blood islands, /48, 89, 251
Blood vessels, formation of, /48, 87
see also arteries and veins
Body axis, 60
Body cavity, see coelom
Body folds, /38, 72
Body-stalk, sec belly-stalk
Bone, cancellous, 272
cells, /1 51, /1 52, 275
compact, 272
endochondral, 272
histogenesis of, 271
intramembranous, 272
lacuna, 275
lamellae, /1 51, 275
marrow, /1 52, 251
matrix, /1 51, 273
trabeculae, /1 51, /1 53, 274
Bone, formation of,
 
compact from cancellous, 281
 
 
Bone — '{Continued)
 
endochondral, /1 55, 276
flat, /1 57, 282 '
 
intramem branous, /1 5 1 , /1 53, 272
long, /1 58, 283
 
primary cancellous, /1 51, /153, 272,
281
 
Bowman’s capsule, /1 16, 203
Brachial plexus, 116
Brain, formation of, /36, 69
neuromeric structure of, 69
regional differentiation of,/87, 1 S4
ventricles of, /88, 156, 160
3-vesicle stage, /36, 70
5-vesicle stage, /60, 110
Branchial arches (= gill arches, q.v.)
 
Bridge of nose, 296
 
Broad ligaments of uterus, 222
 
Bronchi, 189
 
Bulbo-urethral gland, /3, /1 24, 9, 213
 
Clalcification
 
of bone, 274
of teeth, 309
 
Calyces of renal pelvis, /1 19
Canal, atrio-ventncular, 126
Haversian, /1 56, 282
inguinal, /1 29, 222
of Gartner, 217
pleural,/! 10
pleuro- peritoneal, 195
semicircular, 169
Canalicuii, /152
Cancellous bone, /1 53, 276
Capsule, glomerular, /1 21, 203
of Bowman, /1 21 , 203
of cartilage cells, 278
Cardiac loop, /1 42, 125, 254
Cardinal vessels, see veins
“Cartilage bone,” 272
Cartilage, erosion, /1 55, 280
formation, /1 54, 277
matrix, 278
Caval plica, /1 40, 245
Cecum, /102, 122, 182
Cementoblasts, /1 84
Cemcntum,/184, 312
Central canal of spinal cord,/86, 151
Centrum, see vertebrae
Cephalic region, differentiation of, 61
mesoderm of, 1 91
 
 
 
INDEX
 
 
343
 
 
Cephalic region — (Continued)
precocity of, 54
Cerebellar peduncles, 157
Cerebellum, /80,/87, 157
Cerebral aqueduct, 158
Cerebral cortex, 145, 162
Cerebral ganglia, see ganglia, cranial
Cerebral hemispheres, /87, 162
Cerebral peduncles, /1 00, 158
Cerebro-spinal paths, 142
Cervical flexure, 66
Cervical sinus, /32, /1 69, 109
Cervix of uterus, 216
Choanae, of nose, 296
Chondrin, 278
Chondrogenesis, 276
Chondrogenctic layer, 278
Chorda dorsalis ( = notochord, q.v.)
 
Chorion, /55-57, 103
Chorionic vesicle, /54, 103
Chorionic villi, /57
Choroid fissure, of eye,/39, 117
Choroid plexus, anterior (of 3rd ventricle),
/106, 158, 162
 
lateral (of 1st and 2nd ventricles), /1 00.
 
162
 
posterior (of 4th ventricle), /65, /99,
/1 38, 156
 
Chromaffin tissue, 223
Chromosomes, sex, 23
species number of, 21
Circle of Willis, /1 33, 131
Circulation, changes in at birth, 268
early embryonic, /45,/51, 92
hepatic portal, 241
interpreta^tion of embryonic, 227
placental,/45,/51,/55, 93, 263
pulmonary, 262, 267
vitelline, /45, /1 41, 93, 249
Circulatory arcs, 92
Cleavage, /1 2, /1 3, /1 4, 37
Cleavage cavity, see blastococle
aitoris,/132, 225
Cloaca, /65, /1 18, 120, 209
Cloacal plate (membrane), /37, 209
Cochlea, 169
Coelom, /1 09
 
abdominal, /1 11, 194
diflferentiation of, /1 08, 193
formation of,y20, f22,/108, 51, 120, 189
intra- and extra-embryonic, 52, 190
 
 
Coelom — (Continued)
 
partitioning of, /1 11-1 13, 194
pericardial, /26, /44, /1 09, /1 10, /111,
53, 87, 195
peritoneal, /1 10, 189
pleural,/! 11, /1 13, 194
thoracic, 194
 
Colliculi, inferior, suptTior (lobes of corpora
quadngemma), /80, /87, 157, 167
Colon (large intestine), /1 02, 182
Columns of spinal cord, /8 6, 154
Commissural ganglion, /92, 172
Components of spinal nerve, 151
Concrescence, /20, 46
Coordinating centers, 145
Copula, /41
Cord, spinal, 71, 115
 
and reflexes, /80, 142
histogenesis of, /8 1-8 3, 147
white and grey matter of,/86, 151
Cord, umbilical, 250
Corona radiata, 18
Coronary sinus, sec veins
Corpora quadrigemma, /87, 157
Corpus albicans, 27
Corpus hacmorrhagicurn, 26
Corpus luteum, formation of,/6, 24
in pregnancy, /1 0, 26
significance of, 33
Corpus striatum, 1 63
 
Cowper’s gland (= bulbo-urethral gland,
q.v.)
 
Cranial flexure, 66
 
Cranial ganglia, see ganglia
 
Cranial nerves, see nerves
 
Crura cerebri (= cerebral peduncles, q.v.)
 
Cumulus oophorus, 19
 
Cutis plate, see dermatome
 
Cystic duct,/104,/105, 184
 
Decalcification, 274
Deciduous placenta, 103
Deitcr’s nucleus, /80, 144
Dental ledge, /1 75-1 80, 302
Dental papilla, 307
Dental sac, /1 82, 312
Dentinal fiber /1 83, 310
Dentine, /1 83, 307
Dermatome, /42, 81
 
Deutoplasm, effect of on cleavage, /1 2, 37
in pig ovum, 39
 
 
 
344
 
 
INDEX
 
 
Diaphragm, /1 00, /1 12, 194
Diaphragmatic ligament of mesonephros,
/1 23, 218
 
Diaphysis, of long bone, 283
Dicnccphalon, /60, 110, 158
Diestrum, 29
 
Diocoele ( =« lumen of diencephalon)
Dio-mesencephalic boundary, 110
Dio-telcncephalic boundary, 110
Diploe, of bone, 282
Diploid number of chromosomes, 22
Dorsal, aorta, see arteries
flexure, 66
 
mesentery, /1 08, /1 11, 192
mesocardium,/43,/144, 87, 254
nerve roots, see nerves, spinal
root ganglia, see ganglia, spinal
Duct of, Cuvier ( = common cardinal
vein, q.v.)
 
Santorini, 186
Wirsung, 186
 
Ductus, arteriosus, /1 33, /1 38, /1 50, 235,
262, 268
 
choledochus ( = common bile duct, q.v.)
deferens, /3, /1 18, /1 24, 9, 213
endolymphaticus, 117
venosus, see veins
Duodenum, /1 00, /1 02, 184
 
Ear, external, /33,/34, 62
internal, 62, 117, 169
middle, 118, 170
Ectoderm, 59, 98
 
derivatives of,/27
Efferent ductules, 213
Egg nests, 1 5
 
Ejaculatory duct, /3, /1 24, 9, 213
Embryonic disk, /1 6, /1 9, 45, 60
Enamel, /183, 310
Enamel organ, /1 79-1 82, 303
Enamel prisms, 311
 
Endocardial cushion of A-V. canal, /39,
/147,/148, 126, 256
Endocardial cushion tissue, 256
Endocardial primordia,/43,/44, 85
Endocardium, 85
Endolymphatic duct, 117
Endothelium, origin of vascular, /48, 87
Entoderm, 59
 
derivatives of, /27
formation of, /1 6, 43
 
 
Eparterial bronchus, /1 07
Ep>endymal cells, 150
Ependymal layer of c^d,/81, 147
Epicardium, 85
 
Epididymis, /3, /1 24, /1 29, 213
 
Epi-myocardium,/43,/44, 85
 
Epiphyseal cartilage plates, 284
 
Epiphyseal ossification centers, 284
 
Epiphyses of long bones, /1 58, 284
 
Epiphysis, of diencephalon, /1 06, 110, 158
 
Epiploic foramen, 181
 
Epo6phoron,/125, 217
 
Equation division, 22
 
Erythroblasts, /1 52
 
Esophagus, /64,/72, 80, 119, 179
 
Estrus, 29
 
Estrus cycle, /9, 29
 
Eustachian tube, 118, 175
 
Evolution, 198, 229
 
Exocoelom, 52
 
Extra-embryonic coelom, see coelom
Extra-embryonic membranes, /49, 94
Eye, 63, 116, 167
 
Face, development of, /1 68-1 72, 293
Facial nerve, /92, 169
Falciform ligament, /1 1 1 , 193
Fallopian tube (= uterine tube, q.v.)
Fasciculi, see columns of spinal cord
Fertilization, /1 1, 34
Flexion, 65, 107
Floor plate of neural tube, 156
Fetal membranes, see extra-embryonic
membranes
 
Follicle, ovarian, /6,/7, 17
Foramen, ovale, 260, 267, 268 see also
interatrial
 
Foramen of Monro, /88, /1 38, 160
Foramen of Winslow, 181
Fore-brain, see prosencephalon
Fore-gut, /26,/37, 74
Fossa ovalis, 269
 
Fovea cardiaca (= ant. intestinal portal,
q.v.)
 
Frontal lobe, 163
 
Frontal process, /1 68-1 72, 293
 
Froriep’s ganglion, /92, 172
 
Gall-bladder, /46, /103-105, /1 40, 80, 119,
184
 
Gametes, /8, 9, 12, 15
 
 
 
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Patten BM. Embryology of the Pig. (1951) The Blakiston Company, Toronto.

Patten 1951: 1 Foreword to the Student | 2 Reproductive Organs - Gametogenesis | 3 Sexual Cycle | 4 Cleavage and Germ Layers | 5 Body Form and Organs | 6 Extra-Embryonic Membranes | 7 Embryos 9-12 mm | 8 Nervous System | 9 Digestive - Respiratory and Body Cavities | 10 Urogenital | 11 Circulatory System | 12 Bone and Skeletal System | 13 Face and Jaws | Bibliography
Online Editor 
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This historic 1951 embryology of the pig textbook by Patten was designed as an introduction to the topic. Currently only the text has been made available online, figures will be added at a later date. My thanks to the Internet Archive for making the original scanned book available.


By the same author: Patten BM. The Early Embryology of the Chick. (1920) Philadelphia: P. Blakiston's Son and Co.

Patten BM. Developmental defects at the foramen ovale. (1938) Am J Pathol. 14(2):135-162. PMID 19970381


Modern Notes

pig

Historic Disclaimer - information about historic embryology pages 
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Pages where the terms "Historic" (textbooks, papers, people, recommendations) appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms, interpretations and recommendations may not reflect our current scientific understanding.     (More? Embryology History | Historic Embryology Papers)

Embryology of the Pig

Bradley M. Patten
Bradley Patten ( -1971)

Frontispiece

Reconstruction (X 17.5) showing the organ systems of a 9.4 mm. pig embryo. For explanation see figures 60 and 66.

By Bradley M. Patten

Professor of Anatomy in the University of Michigan Medical School


Third Edition


With Colored Frontispiece

And 186 Illustrations In The Text (Containing 412 Figures) Of Which 6 Are In Color


Philadelphia : THE BLAKISTON COMPANY : Toronto


Third Edition


Copyright, October 1948, by The Blakiston Company


By P. Blakiston's Son & Co.


Copyright, 1951, by P Blakiston's Son & Co , Inc.


Patten 1951: 1 Foreword to the Student | 2 Reproductive Organs - Gametogenesis | 3 Sexual Cycle | 4 Cleavage and Germ Layers | 5 Body Form and Organs | 6 Extra-Embryonic Membranes | 7 Embryos 9-12 mm | 8 Nervous System | 9 Digestive - Respiratory and Body Cavities | 10 Urogenital | 11 Circulatory System | 12 Bone and Skeletal System | 13 Face and Jaws | Bibliography

Preface to Third Edition

In making the revision for a new edition of this book it did not seem desirable essentially to change its form or scope. Effort has been concentrated on improving the presentation of the original subject matter and bringing it up to date, rather than on its expansion. The entire book has been reset to a greater page width which has permitted enlarging certain of the illustrations that, in earlier editions, had proved to be too greatly reduced. With the generous cooperation of the publishers several of the important plates on the cardiovascular system have been remade to a larger scale and with color. A number of new illustrations have been drawn for sections where experience has shown that students needed additional graphic assistance in interpreting their laboratory material. It is hoped that these changes will all contribute toward making the book as a whole more serviceable.


Bradley M. Patten

August 1048


Preface to First Edition

This book represents an endeavor to set forth in brief and simple form the fundamental facts of mammalian development. The thread of the story and the illustrations have been based on pig embryos because of their value and availability as laboratory material. But special stress has been laid on the embryological phenomena involved instead of on the details of specific conditions existing in the pig. Throughout the book, every efTort has been made to present developmental processes as dynamic events with emphasis on their sequence and significance, rather than as a series of still pictures of selected stages.


Obviously no book can deal fully with all phases of development, even in a single form, and still remain serviceable as a text. As this book is for the student, it has seemed expedient, for the sake of clearness and simplicity, to omit many things which I should like to have included. My primary aim has been to write an account in which the essentials stand out adequately interpreted and unobscured by a multiplicity of details — to lay a foundation which can be further built upon in accordance with special needs or individual desires.


Bradley M. Patten

January 1927

Acknowledgment

The pleasantest thing about working on this book has been the generous aid I have received from many sources. Throughout the preparation of the initial edition the encouragement, criticism, and suggestions of my colleagues, Dr. F. C. Waite and Dr. S. W. Chase, were of the greatest help. In the preparation of material and in making the illustrations for the first edition the beautifully accurate work of Miss Kathryn Toulmin was of inestimable value. In making the new drawings added in the third edition I was fortunate in securing the unusually able assistance of Mrs. Dorothy Van Eck.


Dr. G. L. Streeter and Dr. C. H. Heuser of the Carnegie Institute allowed me free use of their extensive collection of young embryos and generously gave me many photographs made from their material. To Mrs. Charles S, Minot I am indebted for permission to use several figures from the late Professor Minot's works. The accrediting in the figure legends of these and other borrowed illustrations by no means covers my obligation to other writers. Practically the entire bibliography is a statement of indebtedness for information and ideas.


I wish I might acknowledge individually the helpful services rendered by many of my students, but they are too numerous. Several reconstructions which I have used directly or indirectly have been largely their work. Of even greater assistance have been their suggestions during the shaping of the work — suggestions of especial value because they were made from a point of view difficult for an instructor to appreciate without such aid.


To The Blakiston Company, I am indebted for much helpful cooperation and especially for their liberality with regard to the number and quality of the illustrations.


No person other than my wife could have deciphered and put into usable form manuscript of the character I frequently turned over to her for revision and typing. Without her generous help the preparation of the text would have been much more arduous and long delayed.


Bradley M. Patten


Cite this page: Hill, M.A. (2024, May 7) Embryology Book - Embryology of the Pig. Retrieved from https://embryology.med.unsw.edu.au/embryology/index.php/Book_-_Embryology_of_the_Pig

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© Dr Mark Hill 2024, UNSW Embryology ISBN: 978 0 7334 2609 4 - UNSW CRICOS Provider Code No. 00098G


Patten 1951: 1 Foreword to the Student | 2 Reproductive Organs - Gametogenesis | 3 Sexual Cycle | 4 Cleavage and Germ Layers | 5 Body Form and Organs | 6 Extra-Embryonic Membranes | 7 Embryos 9-12 mm | 8 Nervous System | 9 Digestive - Respiratory and Body Cavities | 10 Urogenital | 11 Circulatory System | 12 Bone and Skeletal System | 13 Face and Jaws | Bibliography
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