Book - Embryology of the Pig 13
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Patten BM. Embryology of the Pig. (1951) The Blakiston Company, Toronto.
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Patten BM. Developmental defects at the foramen ovale. (1938) Am J Pathol. 14(2):135-162. PMID 19970381
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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
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., Contrib. to E-mbryoL, Vol. 2, pp. 69-94.
Corner, G. W., 1917. Variations in the amount of phosphatids in the corpus luteum of the sow during pregnancy. Jour. Biol. Chem., Vol. 29, pp. 141-143.
Corner, G. W , 1919. On the origin of the corpus luteum of the sow from both granulosa and theca interna. Am. Jour. Anal., Vol. 26, pp. 117-183.
Cornet, G. W., 1921. Cyclic changes in the ewaries and uterus of swine, and their relations to the mechanism of implantaticm. Carnegie Inst., Contrib. to Embryol., Vol. 13, pp. 117-146.
Corner, G. IT., 1923. Cyclic variation in uterine and tubal contraction waves. Am. Jour. Anat., Vol. 32, pp. 345-351.
Corner, G IT., 1928. Physiolcjgy of the corpus luteum. I. The efTect of \Try early ablation of the corpus lutcnun upon embryos and uterus. Am. Jour. Physiol., Vol. 86, pp. 74-81.
Coiner, G IV., 1942. The Hormones in Human Reproduction. Princeton Univ. Press, Princeton, xix & 265 pp.
Corner, G. IT., and Allen, IT, M., 1929. Physiology of the corpus luteum. II. Production (3f a special uterine reac tion (progestational proliferation) by extracts of the cwpus luteum. Am. Jour. Physiol., Vol. 88, pp. 326-339.
Coinn, G. IT., and Allen, IT. A/., 1929. Physiology of the corpus luteum. III. Normal growth and implantation of embryos after very earlv ablation of the ovaries, under the influence of extracts of the corpus luteum. Am. lour. Physiol., Vol. 88, pp. 340-346.
Cornn, G. IT, and Amsbaugh, A. E., 1917. Oestrus and ovulation in swine. Anat. Rec., Vol. 12, pp. 287-291.
Corner, G. IT., and Snyder, F. F., 1922. Observations on the structure and function of the uterine ciliated epithelium in the pig, with reference to certain clinical hypothesep. Am. Jour. Obs. & Gyn., VcjI. 3, pp. 358-366.
Crew, F. A. E., 1925. Prenatal death in the pig and its effect upon the sex-ratio. Proc. Roy. Soc. Edinburgh, V^ol. 46, pp. 9-14.
Hammond, J., 1934. The fertilisation of rabbit ova in relation to time: A method of controlling the litter size, the duration of pregnancy and the weight of the young at birth. Jour. Exp. Biol., Vol. 11, pp. 140-161.
Hartman, C. G., 1929. The homology of menstruation. J. A. M. A., Vol. 92, pp. 19921995.
Hartman, C. G., and Squier, R. R., 1931. The follicle-stimulating effect of pig anterior lobe on the monkey ovary. Anat. Rec., Vol. 50, pp. 267-273.
Keye, J. D., 1923. Periodic variations in spontaneous contractions of uterine muscle in relation to the oestrous cycle and early pregnancy. Bull. Johns Hopkins Hosp., Vol. 34, pp. 60-63.
Lewis, L. L., 1911. The vitality of reproductive cells. Agric. Exp. Sta. Oklahoma, Bull. No. 96.
322
BIBLIOGRAPHY
Lillie^ F. /?., 1919. Problems of Fertilization. Univ. Chicago Press, xii & 278 pp.
Loeb^ Z.., 1923. The mechanism of the sexual cycle, with special reference to the corpus luteum. Am. Jour. Anat., Vol. 32, pp. 305-343. ^
Markee, J. E., Pasqualetti^ R. A., and Hinsey, J. C., 1936. Growth of intraocular endometrial transplants in spinal rabbits. Anat. R^c., Vol. 64, pp. 247-253.
Marshall^ F, H. A., 1922. The Physiology of Reproduction. Longmans, Green & Co., London, 2nd Ed., xvi & 770 pp.
McKenzie^ F. F, 1926. The normal oestrous cycle in the sow. Univ. Missouri Coll. Agric. Exp. Sta. Res. Bull., Vol. 86, pp. .S-41.
Papanicolaou^ G. A'"., 1923. Oestrus in mammals from a comparative point of view. Am. Jour. Anat., Vol. 32, pp. 285-292.
Parker^ G. IL, 1931. Passage of sperms and of eggs through oviducts in terrestrial vertebrates. Phil. Trans. Roy. Soc. London, Ser. B, Vol. 219, pp. 381-419.
Phillips, R. W., and Andrews, F, j\., 1937. The speed oi travel of ram spermatozoa. Anat. Rec., Vol. 68, pp. 127-132.
Pincus, G., 1936. The Eggs of Mammals. Macmillan, New York, ix & 160 pp.
Schott, R. G., and Phillips, R. W., 1941. Rate of sperm travel and time of ovulation in sheep. Anal. Rec,, Vol. 79, pp. 531-540.
Secktnger, D, L., 1923. Spontaneous contractions of the Fallopian tube of the domestic pig with reference to the oestrous cycle. Bull. Johns Hopkins Hosp., Vol. 34, pp. 236-239.
Snyder, F. F., 1923. Changes in the Fallopian tube during the ovulation cycle and early pregnancy. Bull. Johns Hopkins Hosp., Vol. 34, pp. 121-125.
Soderwall, A, L., and Blandau, R. J., 1941. The duration of the fertilizing capacity of spermatozoa in the female genital tract of the rat. Jour. Exp. Zool , Vol. 88, pp. 55-64.
Stockard, C. /?., 1923. The general morphological and physiological importance of the oestrous problem. Am. Jour. Anat., Vol. 32, pp. 277-283.
Surface, F. M., 1908, Fecundity of swine. Biometrika, Vol. 6, pp. 433-436.
Swingle, W, W., 1926. The determination of sex in animals. Physiol. Rev., Vol. 6,
pp. 28-61.
Toothill, M. C., and Toung, W. C.,\93\. The time consumed by spermatozoa in passing through the ductus epididymidis of the guinea-pig as determined by means of India-ink injections. Anat. Rec., Vol. 50, pp. 95-107.
Wilson, K. M., 1926. Histological changes in the vaginal mucosa of the sow in relation to the oestrous cycle. Am. Jour. Anat., Vol. 37, pp. 417-432.
Witschi, E., 1932. Physiology of embryonic sex differentiation. Am. Nat., Vol. 66, pp. 108-117.
Cleavage, Germ Layer Formation^ and the Structure of Young
Embryos
Assheton, R., 1899. The development of the pig during the first ten days. Quart. Jour. Micr. Sci., Vol. 41, pp. 329-359.
Clark, R, T,, 1934. Studies on the physiology of reproduction in the sheep. II. The cleavage stages of the ovum. Anat. Rec., Vol. 60, pp. 135-159.
Conklin, E, G., 1909. The application of expieriment to the study of the organization and early differentiation of the egg. Anat. Rec., Vol. 3, pp. 149-154.
CI.EAVAGE, GERM LAYERS, YOUNG EMBRYOS
323
Defrise^ A.^ 1933. Some observations on living eggs and blastulae of the albino rat.
Anat. Rec., Vol. 57, pp. 239-250.
Green^ W, W., and Winters, L. Af., 1946. Cleavage and attachment stages of the pig. Jour. Morph., Vol. 78, pp. 305-316.
Heuser, C\ H., and Streeter, G. L., 1 929. Early stages in the development of pig embryos, from the period of initial cleavage to the time of the appearance of limb-buds. Carnegie Inst., Contrib. to Embryol., Vol. 20, pp. 1-29.
Keibel, F., 1891. Uebcr die Entwicklungsgeschichte des Schweines. Anat. Anz., Bd. 6, S. 193-198.
Keibel, F., 1894. Studien zur Entwicklungsgeschichte des Schweines (Sus scrofa domesticus). I. Morph. Arbeiten, Bd. 3, S. 1-139.
Keibel, F., 1895. Ueber einige Plattenmodelle junger Schwein-embryonen. Verhandlungen d. Anat. Ges., Erganzungsheft Bd. 10, Anat. Anz., S. 199-201. Keibel, F., 1896. Studien zur Entwicklungsgeschichte des Schweines (Sus scrofa domesticus). II. Morph. Arbeiten, Bd. 5, S. 17-168.
Kingsbury, B, F., 1920. I'he developmental origin of the notochord. Science, N. S., Vol. 51, pp. 190-193.
Kingsbury, B. F., 1924a. The developmental significance of the notochord (Chorda dorsalis). Zeitschr. f. Morphologic u. Anthropologic, Vol. 24, pp. 59-74. Kingsbury, B. F,, 1924b. The significance of the so-called law of cephalocaudal differential growth. Anat. Rec., Vol. 27, pp. 305-321.
Kingsbury, B, F., 1926. On the so-called law of antero-posterior development. Anat. Rec., Vol. 33, pp. 73-87.
Lewis, F. T., 1902. The gross anatomy of a 12-mm. pig. Am. Jour. Anat., Vol. 2,
pp. 211-226.
Lewis, W, H,, and Gregory, P. W,, 1929. Cinematographs of living developing rabbiteggs. Science, Vol. 69, pp. 226-229.
Lewis, W. H., and Hartman, C, G., 1933. Early cleavage stages of the egg of the monkey (Macacus rhesus). Carnegie Inst., Contrib. to Embryol., Vol. 24, pp. 187-201. Lewis, W. PL, and Wright, E, S,, 1935. On the early development of the mouse egg.
Carnegie Inst., Contrib. to Embryol., Vol. 25, pp. 113-144.
Macdonald, E., and Long, J, A., 1934. Some features of cleavage in the living egg of the rat.^Am. Jour. Anat., Vol. 55, pp. 343-361.
Nicholas, J. S., and Hall, B. V., 1942. Experiments on developing rats. II. The development of isolated blastomeres and fused eggs. Jour. Exp. Zooi., Vol. 90, pp. 441-459.
Patten, B. M,, and Philpott, R., 1921 . The shrinkage of embryos in the processes preparatory to sectioning. Anat. Rec., Vol. 20, pp. 393-413.
Pincus, G,, 1939. The comparative behavior of mammalian eggs in vivo and in vitro. IV. The development of fertilized and artificially activated rabbit eggs. Jour. Exp. Zodl., Vol. 82, pp. 85-131.
Robinson, A,, 1892. Observations upon the development of the segmentation cavity, the archenteron, the germinal layers, and the amnion in mammals. Quart. Jour. Micr. Sci., Vol. 33, pp. 369-455.
Streeter, G. L,, 1926. Development of the raesoblast and notochord in pig embryos.
Carnegie Inst., Contrib. to Embryol., Vol. 19, pp. 73-92.
'J'hyng, F, W,, 1911. The anatomy of a" 7.8-mm. pig embryo. Anat. Rec., Vol. 5, pp. 17-45.
324
BIBLIOGRAPHY
Waldo, C, M., and Wimsatt, W. A., 1945. The effect of colchicine on early cleavage of
mouse ova. Anat. Rec., Vol. 93, pp. 363-375.
Wallin, E,, 1917. A teaching model of a 10-mm. pig embryo. Anal. Rec., Vol. 13, pp. 295-297.
Weysse, A, W,, 1894. On the blastodermic vesicle of Sus scrofa domesticus. Proc. Am. Acad. Arts & Sciences, Vol. 30, pp. 283-321,
Williams, L. W., 1908. The later development of the notochord in mammals. Am. Jour. Anat., Vol. 8, pp. 251-284.
Fetal Membranes and the Relations of the Embryos to the Uterus
Abromavich, C. E., 1926. The morphology and distribution of the rosettes on the foetal placenta of the pig. Anat. Rec., Vol. 33, pp. 69-72.
Assheton, R., 1906. The morphology of the ungulate placenta. Phil. Trans. Roy. Soc. London, Ser. B, Vol. 198, pp. 143-220.
Brambel, C. E., Jr., 1933. Allantochorionic differentiations of the pig studied morphologically and histochemically. Am. Jour. Anat., Vol. 52, pp 397-459.
Corner, G. W., 1921. Internal migration of the ovum. Bull. Johns Hopkins Hosp., Vol. 32, pp. 78-83.
Fawcett, D. W., Wislocki, G. B., and Waldo, C. M., 1947. The development of mouse ova in the anterior chamber of the eye and in the abdominal cavity. Am. Jour. Anat., Vol. 81, pp. 413-443.
Gellhorn, A., Flexner, L. B., and Pohl, H. A., 1941. I’he transfer of radioactive sodium across the placenta of the sow. Jour. Cell. & Comp. Physiol., Vol. 18, pp. 393400.
Goldstein, S. R., 1926. A note on the vascular relations and areolae in the placenta of the pig. Anat. Rec., Vol. 34, pp. 25-35.
Grosser, 0., 1909. Vergleichende Anatomic und Entwicklungsgeschichte der Eihaute und der Placenta, Braumiiller, Leipzig, xi & 314 S.
Grosser, 0., 1927. Friihentwicklung, Eihautbildung und Placentation des Mcnschen und der Saugetiere. Verlag von J. F. Bergmann, Munchen, viii & 454 pp.
Heuser, C. H., 1927. A study of the implantation of the ovum of the pig from the stage of the bilaminar blastocyst to the completion of the fetal membranes. Carnegie Inst., Contrib. to Embr^ol., Vol. 19, pp. 229-243.
Hubrecht, A. A. W., 1895. Die Phylogenese des Amnions und die Bedeutung des Trophoblastes. Verh. Koniklijke Akad. van Wetenschappen te Amsterdam, fiir 1895, S. 3-66.
Lell, W. A., Liber, K. E., and Snyder, F. F., 1931. Quantitative study of placental transmission and permeability of fetal membranes at various stages of pregnancy. Am. Jour. Physiol., Vol. 100, pp. 21-31.
Mossman, H. W., and Noer, H. R., 1947. A study of the amnion with the electron microscope. Anat. Rec., Vol. 97, pp. 253-257.
Noer, H. R., 1946. A study of the effect of flow direction on placental transmission, using artificial placentas. Anat. Rec., Vol. 96, pp. 383-389.
Robinson, A., 1904. Lectures on the early stages in the development of mammalian ova and on the formation of the placenta in different groups of mammals. Jour. Anat. & Physiol., Vol. 38, pp. 186-204.
THE NERVOUS SYSTEM AND SENSE ORGANS
325
Runner, M, JV,, 1947. Development of mouse eggs in the anterior chamber of the eye.
Anat. Rec., Vol. 98, pp. 1-1 7.
Warwick, B. L., 1926. Intra-uterine migration of ova in the sow. Anat. Rec., Vol. 33, pp. 29-33.
Wislocki, G. B.y 1929. On the placentation of primates, with a consideration of the phylogeny of the placenta. Carnegie Inst., Contrib. to Embryol., Vol. 20, pp. 51-80.
Wisloch, G. B., 1935. On the volume of the fetal fluids in sow and cat. Anat. Rec., Vol. 63, pp. 183-192.
Widocki, G. B., and Dempsey, E. W,, 1946. Histochemical reactions of the placenta of the pig. Am. Jour. Anat., Vol. 78, pp. 181-225.
The Nervous System and Sense Organs
Assheton, R., 1892. On the development of the optic nerve of vertebrates and the choroidal fissure of embryonic life. Quart. Jour. Micr. Sci., Vol. 34, pp. 85-103.
Bardeen, C. R., 1903. The growth and histogenesis of the cerebrospinal nerves in mammals. Am. Jour. Anat., Vol. 2, pp. 231-257.
Barnes, W,, 1883-84. On the development of the posterior fissure of the spinal cord and the reduction of the central canal in the pig. Proc. Am. Acad. Arts. & Sciences, Vol. 19, pp. 97-110.
Bedford, E. A., 1904. The early history of the olfactory nerve in swine. Jour. Comp. Ncur., Vol. 14, pp. 390-410.
Bradley, 0. C., 1904. Neuromeres of the rhombencephalon of the pig. Review of Neurology & Psychiatry, Vol. 2, pp. 625-635.
Coghill, G. E., 1926. The growth of functional neurones and its relation to the development of behavior. Proc. Am. Philosoph. Soc., Vol. 65, pp. 51-55.
Detwiler, S. R., 1936. Neuroembryology. An Experimental Study. Macmillan, New York, X & 218 pp.
Dowd, L. W., 1929. The development of the dentate nucleus in the pig. Jour. Comp. Neur., Vol. 48, pp. 471-498.
GrademgOy G., 1887. Die embryonale Anlage des Mittelohrs; die morphologische BedeutiAig der Gehorknochelchen. Mitth. aus dem Embryeflog. Inst. d. Universitat. Wien, 1887, S. 85-232.
Hardesty, /., 1904. On the development and nature of the neuroglia. Am. Jour. Anat., Vol. 3, pp. 229-268.
Hardesty, /., 1905, On the occurrence of the sheath cells and the nature of the axone sheaths in the central nervous system. Am. Jour. Anat., Vol. 4, pp. 329-354.
Held, H., 1909. Die Entwicklung des Nervengewebes bei den Wirbeltieren. J. A. Barth, Leipzig, ix & 378 S.
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Lymphatic Vessels and Organs
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The Digestive and Respiratory Systems, and the Body Cavities
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Swaen, A., 1896 and 1897. Recherches sur le developpement du foie, du tube digestive,
de rarriere cavit6, du p6ritoine et du mesentere. Jour, de TAnat. et de la Physiol.,
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The Urogenital System
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THE DUCTLESS GI.ANDS
335
Henneberg, 1925. Anatomic und Entwicklung der Ausseren Genital organe des
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The Ductless Glands
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336
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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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Teeth, Hair, and Hoofs
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TWINS, DOUBLE MONSTERS, ANOMALIES
339
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Twins, Double Monsters, Anomalies
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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.).
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