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proceeds the plate is gradually invaginated into the substance of the head, and is
FIG. 67.-TRANSVERSE SECTION OF A Rat EMBRYO. Showing the relation of the paraxial mesoderm of the head to the lateral plates, the commencement of the
formation of the otic vesicles and hyomandibular clefts, and the relation of the primitive heart to the pericardium and fore-gut. EC. Ectoderm. SOM. Somatic mesoderm.
SpM. Splanchnic mesoderm. transformed into a pear-shaped vesicle, the otic vesicle, which remains for a time in communication with the exterior by means of a short tubular stalk, the recessus labyrinthi, which is subsequently converted into the ductus endolymphaticus.
After it is separated from the surface the otic vesicle alters its position, until its ventral end lies in close relation to the dorsal wall of the pharynx, and, at the same
า 1 time, it undergoes alteration of shape. The ventral part of the vesicle grows towards the median plane, along the ventral wall of the hind-brain. It forms the cavity and the lining epithelium of the cochlea; but it remains in con
Fig. 68.- TRANSVERSE SECTION THROUGH THE HEAD nexion with the dorsal part
OF AN EMBRYO. by means of a narrow tube, Showing the rudiments of the three parts of the ear and their the canalis reuniens, and as it
relation to the hyomandibular cleft. grows in length it becomes BV. Blood-vessels.
OV. Otic vesicle. converted into a spiral tube.
EM. Ext. acoustic meatus. P. Pharynx. The portion of the dorsal ET. Auditory tube.
RL. Recessus labyrinthi. section of the primitive vesicle,
SC. Semicircular canal.
T. which lies to the lateral side
HM. Hyomandibular cleft.
Tympanum. of the recessus labyrinthi, first expands and then becomes compressed and 1 See note 3, p. 79.
constricted into the form of three flat purse-like diverticula which, by the partial obliteration of their cavities, become converted into the three semicircular canals (see Sense Organs). The more ventral part of the dorsal section of the vesicle is divided, by a constriction of its lateral wall, into a dorsal part, the utricle, which remains in connexion with the semicircular canals, and a ventral part, the saccule, which is united to the cochlea by the canalis reuniens. The apex of the constriction which separates the utricle from the saccule passes into the mouth of the ductus endolymphaticus, which is thus transformed into the Y-shaped canal which connects the utricle with the saccule. At a later period the closed extremity of the ductus endolymphaticus dilates and forms a small saccule, the saccus endolymphaticus. In the adult the saccus endolymphaticus lies in the posterior fossa of the skull,
in relation with the posterior surface of the petrous part of the temporal bone and ternal to the dura mater.
and the auditory tube (O.T. Eustachian)
developed from the first visceral pouch.
The ventral part of the pouch -5 disappears at an
early stage. The dorsal extremity expands and is converted into the cavity of the tympanum,
whilst the stalk FIG. 69.-FIGURES, MODIFIED FROM HIS, ILLUSTRATING THE FORMATION OF
of connexion THE PINNA.
with the pharynx 1. Tuberculum tragicum=Tragus.
Tuberculum antitragicum = Anti-
is gradually conintermedium helicis
7. Tuberculum lobulare = Lobule. stricted off from 4. Cauda helicis
HM. Hyomandibular cleft.
its lateral to5. Tuberculum anthelicis = Antihelix. OV. Otic vesicle.
wards its medial end, and is converted into the auditory tube. The constriction commences when the embryo has attained a length of about 20 mm., that is about the beginning of the eighth week, and is completed about the end of that week when the embryo is about 25 mm. long.
After the auditory tube is defined it grows rapidly in length, and cartilage appears in its walls during the fourth month.
As the tympanic cavity increases in size the auditory ossicles-stapes, incus, and malleus, which are differentiated from the dorsal ends of the cartilages of the first and second branchial arches, are invaginated into it.
The membrana tympani, which separates the tympanum from the external acoustic meatus, is formed from the separating membrane which intervenes between the first branchial pouch and the first cleft. It consists, therefore, of an external covering of ectoderm, an internal lining of entoderm, and an intervening layer, of fibrous tissue, derived from the mesoderm.
The external ear is developed from the cavity and the boundaries of the first branchial cleft. The cavity of the cleft is transformed into the cavity of the external acoustic meatus, and on the mandibular and on the hyoid margins of the
cleft three eminences appear. From the eminences on the two arches, and the skin immediately posterior to the eminences on the hyoid arch, are formed the various parts of the auricle, but the exact part played by the individual eminences in the human subject is as yet a matter of some doubt.
THE PROTECTION AND NUTRITION OF THE EMBRYO DURING
ITS INTRA-UTERINE EXISTENCE.
Whilst it is passing down the uterine tube, and for a brief period after it enters the uterus, the zygote, or impregnated ovum, depends for its nutrition upon the yolk granules (deutoplasm) embedded in its cytoplasm, and upon the fluid medium surrounding it which is secreted by the walls of the uterine tube and the uterus.
As the human ovum is very small, and as it contains but little deutoplasm, its nutrition is practically dependent, almost from the first, upon external sources of supply. The urgent necessity for the formation of adequate arrangements whereby the external sources may be utilised leads to the early establishment of an intimate connexion between the zygote and the mother, which is one of the characteristic features of the development of the human embryo.
During the third week after fertilisation, as the embryo is beginning to be moulded from the embryonic region, and before the paraxial mesoderm commences to separate into mesodermal somites, a primitive heart and the rudiments of some well-defined blood - vessels are distinguishable in the embryo; but the details of the development of the vascular system and the establishment of the embryonic circulation cannot be well understood until the formation and structure of a group of closely associated extra-embryonic organs or appendages, derived from the zygote, has been considered.
This group includes the chorion, the placenta, the amnion, the umbilical cord, and the yolk-sac.
THE MEMBRANES AND APPENDAGES.
The Chorion.— It has already been noted that when the zygote becomes a blastula it consists of three vesicles, a large vesicle enclosing two smaller vesicles and a mass of primary mesoderm (Fig. 29).
The wall of the large vesicle is composed of trophoblast (trophoblastic ectoderm), and its inner surface is in direct contact with the primary mesoderm.
A little later a cavity, the extra-embryonic cælom, appears in the primary mesoderm, separating it into two layers, one lining the inner surface of the trophoblast and the other covering the outer surfaces of the two inner vesicles (Figs. 70, 71).
As soon as the extra-embryonic coelom is established the chorion is formed ; it consists of the trophoblast and its inner covering of mesoderm.
In the meantime the trophoblast has differentiated into two layers, an inner cellular layer, and an outer plasmodial layer. In the plasmodial layer cell territories are not defined, and it consists, therefore, of nucleated protoplasm.
The differentiation of the trophoblast into two layers occurs after the zygote is embedded in the mucous membrane of the uterus which is modified for its reception and which, after the modification has occurred, is called the decidua.
As development proceeds the trophoblast increases in thickness and it invades the decidua. As this invasion occurs the plasmodial layer of the trophoblast becomes permeated with spaces which are continuous with the lumina of the maternal blood-vessels in the decidua, and are filled with maternal blood.
By means of the spaces the plasmodial trophoblast is separated into branching processes which intervene between the blood-filled spaces. The processes are the primary chorionic villi, and they soon develop cellular interiors (Fig. 72).
After a time the primary villi are invaded by the chorionic mesoderm, and are thus converted into the secondary chorionic villi, which become vascularised by the growth of fætal vessels into the fatal mesodermal cores. The secondary villi, therefore, consist of a mesodermal core covered by a layer of cellular trophoblast and a layer of plasmodium, the latter lying outside the former. Still later the
secondary villi send out numer-
ous branches into the blood
in complexity (Figs. 75, 76, 77). Mesoderm of amnion
As development progresses
is converted into the fætal Body stalk mesoderm portion of an organ called the Extra-embryonic cælom placenta, and thus the chorion
is divided into placental and Entoderin Mesoderm covering of
non-placental regions. Upon 'entoderm vesicle the placental part the villi conNeurenteric canal
tinue to increase, but they disappear entirely from the non
placental part, which is then Cavity of entodermal vesicle
called the chorion læve (Fig. Fig. 70.–Schema of SAGITTAL SECTION OF ZYGOTE ALONG LINE A. 77). Plasmodial trophoblast Neural groove
The Amnion, the BodyChorion
Cellular trophoblast | Mesoderm liningoftrophoblast
Stalk (Allantoic Stalk), and Amnion cavity
the Umbilical Cord.—The Extra-embryonic colom
amnion is formed from that Mesoderm of amnion
portion of the wall of the larger Ectoderm of amnion
of the two inner vesicles of the Mesoderm covering
zygote, the ecto-mesodermal entoderm
vesicle (p. 22), which does Entoderm
not take part in the formation Cavity of entodermal
of the embryo. It consists of ectoderm cells covered. externally by a layer of extra-em
bryonic mesoderm, and it is Notochord
continuous with the margin of the embryonic area (Figs.
70, 71). Fig. 71.—SCHEMA OF TRANSVERSE SECTION OF ZYGOTE ALONG LINE B (Fig. 31).
The cavity of the ecto
mesodermal vesicle, enclosed between the amnion and the embryonic area, is the cavity of the amnion; it is filled with fluid, which raises the amnion in the form of a cupola over the embryonic region (Fig. 70).
The Body-Stalk (Allantoic Stalk).—It has been noted already that the mesoderm of the median part of the posterior or caudal portion of the amnion becomes
Plasmodial Plasmodial. trophoblast
Afferent vessel of villus Cellular
of villus trophoblastMesoderm
Fused mesoderm Ectoderm of chorion
Fused mesoderm of and amnion
Famnion and chorion Ectoderm
Ectoderm of amnion of amnion Fig. 72.-SCHEMA OF THREE STAGES IN THE FORMATION OF A CHORIONIC Villus. thickened. In the thickened strand lies the allantoic diverticulum of the entodermal vesicle (Fig. 70), whilst through it, on either side of the allantoic diverticulum, pass the umbilical arteries and veins, by means of which blood is conveyed between the embryo and the chorion.
This segment of the wall of the amnion vesicle was termed by His the body-stalk. It takes no direct part in the formation of the embryo, and as it
contains the rudimentary allantoic diverticulum and represents the much more highly developed allantois of other forms, it would, perhaps, be better to term it the allantoic stalk. For the present purpose it is important to note that the bloodvessels which pass through the body-stalk enter or leave the body through the umbilical orifice, which is, at first, a relatively large aperture (Fig. 50).
As the embryonic area is folded into the form of the embryo the amnion increases in extent, filling more and more of the extra-embryonic cælom, and the embryo rises into the interior of its cavity. In other words, the walls of the amnion bulge ventrally round the cranial and caudal extremities and the lateral borders of the embryo (Figs. 75, 76, 77). As the distension of the amnion still continues, the ventral bulging, round the margin of the umbilical orifice, becomes more pronounced, the yolk-sac is forced farther
Plasmodial trophoblast and farther away from the embryo, the vitello-intestinal duct is elongated, and
Cellular of villus
trophoblast it is surrounded by a hollow tube. The cavity of the tube is an elongated part of
of villus the extra-embryonic ccelom, and its walls are formed by the amnion (Figs. 57,62,63).
of villus The caudal wall of the tube necessarily consists of the elongated body-stalk
Efferent vessel (allantoic stalk).
As the distension of the amnion still continues, the walls of the tube are forced against the vitello-intestinal duct, and Fig. 73.—SCHEMA OF Transverse Section of a
SECONDARY CHORIONIC VILLUS. the amniotic mesoderm fuses with the
A loop of the
afferent vessel has been cut at two points. mesoderm of the vitello-intestinal duct. When the fusion is completed, a solid cord, the umbilical cord, is formed (Figs. 77, 78, 80). It consists of an external covering of amniotic ectoderm, and a core of mesoderm in which lie the two umbilical arteries of the body-stalk, a single umbilical vein formed by the fusion of the two primitive veins, and the remains of the vitello-intestinal duct and the vitelline vessels. The proximal end of the umbilical cord is connected with the embryo; the distal end is attached to the chorion, and in its neighbourhood lies the now relatively small vesicular yolk-sac (Fig. 62)
As the amnion grows still larger, all that part of its outer surface which does not take part in the formation of the umbilical cord is ultimately pressed into contact with the inner surface of the chorion, with which it fuses, and the cavity of the extra-embryonic part of the coelom is obliterated (Fig. 78).
The outer wall of the zygote now consists of the fused chorion and amnion, and it contains in its interior the amniotic cavity and the embryo, which is attached to the chorion by the umbilical cord.
When it is first formed the umbilical cord is comparatively short, but, as the amniotic cavity increases, the cord elongates, until it attains a length of from 18 to 20 inches, a condition which allows the embryo to float freely in the fluid in the amniotic cavity, whilst its nutrition is provided for by the flow and return of blood, through the umbilical cord, to and from the placenta, where interchanges take place between the maternal and the fatal blood.
The Yolk-Sac or Umbilical Vesicle.— When the embryonic area is folded into the form of the embryo, the entodermal vesicle is differentiated into three parts: (1) a part enclosed in the embryo, where it forms the primitive entodermal alimentary canal; (2) a part which lies external to the embryo in the extraembryonic ccelom—this is the yolk-sac or umbilical vesicle ; (3) the third portion is the vitello-intestinal duct, which connects the primitive alimentary canal and the yolk-sac together (Figs. 40, 62).
The walls and the cavity of the yolk sac are, therefore, continuous with the walls of the primitive alimentary canal, and the structural features of the two are identical, each consisting of an internal layer of entodermal cells and an external layer of splanchnic mesoderm.
Free communication between the yolk-sac and the primitive alimentary canal