DEVELOPMENT OF THE CHICK DURING THE THIRD AND FOURTH DAYS OF INCUBATION External Features In 3-day

chicks torsion has progressed to the point where the embryo lies with its left side on the yolk to a level just posterior to
the heart (Fig. A-35). By 4 days the entire body has been turned so that its entire left side lies on the yolk (Fig. A-36).
Meanwhile, the cranial and cervical flexures have increased to somewhat greater than right angles in 3-day chicks.
Because of the broad attachment to the yolk, the midbody of 3-day chicks is slightly concave dorsally (Fig. A-36). By 4
days the caudal end of the body has also undergone flexion, leaving the embryo C shaped (Fig. A-36). The face and oral
region are similar in their development to those of a 5-week human embryo (cf. Fig. 14-IB), with nasal pits, each
surrounded by a U-shaped elevation consisting of a basolateral and a nasomedial process. The two nasomedial processes
merge with each other in the midline and fuse laterally with the maxillary processes, as in mammals, to form the upper jaw
(Fig. 14-1B to 14-IE). The lower jaw arises by merging in the midline of the right and left mandibular processes of the first
branchial arch. Formation of the beak does not begin for several more days. The branchial-arch system continues to
develop, and by 3 days a fourth branchial cleft has appeared (Fig. A-35). Both the anterior and posterior appendage buds
are grossly evident at 3 days. At this stage they consist of homogeneous mesoderm. The outer covering of ectoderm is
marked by the conspicuously thickened apical ectodermal ridge (Figs. 11-5B and A-39E). The allantois in the 3-day chick is
still small and is concealed by the posterior appendage buds. By 4 days it has undergone rapid enlargement and projects
beyond the confines of the body as a stalked vesicle of considerable size (Fig. A-36). In subsequent days (Fig. 7-5) the
allantois becomes very large.

The Nervous System The five major divisions of the brain are well delineated. As early as the third day but more
prominently in the fourth, the telencephalic vesicles have invaginated from the lateral walls of the forebrain (Fig. A-36).
Their cavities are continuous with the lumen of the median portion of the brain through the foramina of Monro (Fig.
12-13E). The telencephalic vesicles eventually be- come the cerebral hemispheres. The lateral walls of the diencephalon at
this stage show little differentiation except ventrally, where the optic stalks merge into the walls of the brain. The epiphysis
continues as a median evagination from the roof of the diencephalon. The infundibular depression in the floor of the
diencephalon has become appreciably deepened and lies close to Rathke's pocket (Fig. A-37) with which it is destined to
fuse in the formation of the hypophysis. Changes in the walls leading to the formation of the thalami and the
hypothalamus have not yet occurred. The mesencephalon does not show any specializations beyond a thickening of its
walls. Later its dorsal walls will become thicker to form the corpora quadrigemina, four symmetrically placed elevations in
the adult brain. The superior pair (superior colliculí) constitute the brain center for visual reflexes and are very prominent
in birds. The inferior colliculi are the center for auditory reflexes. The metencephalon remains well delineated from the
mesencephalon by a constriction, but it remains poorly delimited from the myelencephalon. The metencephalon shows
practically no differentiation in 4-day chicks. Later, its roof undergoes extensive enlargement to form the cerebellum. Its
base, like that of the mesencephalon, contains the main nerve fiber tracts that connect the brain with the spinal cord. The
myelencephalon is recognizable by its thin dorsal wall, which is characteristic of its adult derivative, the medulla. Its
ventral and lateral walls will serve as a conduction path between the brain and spinal cord and as a reflex center for
involuntary activities such as breathing. Prominent in 4-day embryos are ganglia of the sensory components of the cranial
nerves. The most prominent is the semilunar (gasserian) ganglion of the hfth (trigeminal) nerve (Fig. A-38), which lies
opposite the most anterior neuromere of the myelencephalon. Just cephalic to the auditory vesicle is a mass of neural
crest cells which constitute the primordium of the ganglia of the seventh and eighth nerves. Caudal to the auditory vesicle
one can make out the superior and inferior sensory ganglia of the ninth and tenth cranial nerves (Fig. A-38). As
development progresses, the lateral walls of the spinal cord become arcady thickened in contrast to the dorsal and ventral
walls, which remain thin. In this process the lumen (central canal) becomes compressed laterally until it appears in cross
section as little more than a vertical slit (Fig. A-41). Prominent along the spinal cord are regular spinal (dorsal root) ganglia
which contain the cell bodies of the neural crest-derived sensory neurons. When first formed from the neural crest cells,
the spinal ganglion has no connection with the cord. The dorsal root is established by the growth of nerve fibers from
cells of the spinal ganglion into the cord. At the same time fibers grow distad from these cells to form the peripheral part
of the nerve. The fibers arising from cells of the dorsal root ganglion conduct sensory impulses toward the cord.
Coincident with the establishment of the dorsal root, the ventral root is formed by fibers which grow out from cells located
in the ventral part of the lateral plate of the cord. Most of the fibers which thus arise from cells in the erd and pass out
through the ventral root conduct motor impulses from the brain and cord to the muscles with which they are associated,

The Digestive and Respiratory Systems During the third day the oral plate, the thin membrane of ectoderm and endoderm
that separates the stomodeal depression from the foregut, breaks down, allowing the foregut to communicate with the
outside. Following the rupture of the oral plate, the originally shallow stomodeal depression deepens as a result of growth
of the surrounding structures. The original site of the oral plate becomes the region of transition from the oral cavity to the
pharynx. Caudal to the oral opening the foregut has become flattened dorsoventrally and widened laterally to form the
pharynx. On either side the pharyngeal lumen shows a series of extensions or bays known as the pharyngeal pouches
(Fig. A-38). Each pharyngeal pouch lies opposite an external branchial groove (Fig. A-41B). This leaves in these areas only
a thin layer of tissue (the branchial plate) separating the pharyngeal lumen from the outside. This layer is com- posed
internaily of endoderm and externally of the ectoderm of the bottom of the branchial groove (Fig. A-41B). Between
adjacent branchial grooves or clefts the lateral walls are greatly thickened and filled with closely packed mesenchymal
cells (Fig. A-39B). These thickened areas are known as the branchial arches. One of the most important relationships to
grasp in the study of young embryos is the manner in which the aortic arches lie embedded in the tissue of the branchial
arch of corresponding number. The stereogram appear- ing in Fig. A-40 emphasizes these relationships. Several important
glandular structures arise from portions of the pharyngeal endoderm. The thyroid gland starts out as a median
diverticulum from the floor of the pharynx between the first and second pairs of pharyngeal pouches (Figs. A-37 and A-38).
Sometime after the fourth day, two pairs of parathyroid Primordia, as well as primordia of the thymus gland, arise from the
endodermal lining of the third and fourth pharyngeal pouches. The respiratory tract appears in 3-day chicks as a
midventral groove in the Pharynx. Beginning just posterior to the level of the fourth pharyngeal pouches and extending
caudad, the laryngotracheal groove (Fig. A-39B) deepens rapidly closes off to form the tracheal tube, which continues to
grow caudad and bifurcates to form a pair of lung buds. 4 days the esophagus remains a thin tube, but the segment of the
gut that tdecome stomach is already slightly dilated (Fig. A-38). Just caudal to the daye diverticulum from the ventral wall
of the gut at the end of the second endod original hepatic evagination has grown as a mass of branching cords of stomach
one finds the primordia of the liver and pancreas. After arising as a the invagination closest to the gut remains open and
will eventually endodermal cells, and by 4 days it has become quite massive (Fig. A-38). The part of the invagination
closest to the gut remains open and will eventually serve as the drainage duct of the liver and gallbladder. Regional
differentiation of the drainage system occurs later in development. In birds, the pancreas arises as a single median dorsal
and a pair of ventral pancreatic buds. The dorsal evagination appears at about 72 hours, and the ventral evaginations
appear toward the end of the fourth day. Later in development, the masses of cellular cords derived from the three
pancreatic primordia grow together and become fused into a single glandular mass. In chicks of 4 days, the midgut has
been practically replaced by the extension of the foregut and hindgut. It consists only of the restricted region where the
yolk stalk leads from the gut tract to the yolk sac (Fig. A-37). The beginning of the formation of the cloaca is indicated in
chicks of 4 days' incubation by a dilation of the posterior portion of the hindgut (Fig. A-38). Although extensive
differentiation in the cloacal region does not appear until later in development, certain of its fundamental relationships are
established at this stage. The cloaca of an adult bird is the common chamber into which the intestinal contents, the urine,
and the products of the reproductive organs are received for discharge. Arising from the dorsal wall of the cloaca in birds
is an endodermal outgrowth known as the bursa of Fabricius. Later in development it becomes heavily infiltrated with
lymphoid cells. Its function long remained obscure. It is now recognized that the bursa of Fabricius is one of the major
components of the immune defense system of birds and that if it is extirpated, the ability of the bird to produce humoral
antibodies is sharply reduced (Glick and Chang, 1957). Although the urinary system is not yet developed to conditions
which resemble those in the adult, the parts of it which have been established are already definitely associated with the
cloaca. The proximal portion of the allantoic stalk, which is the homologue of the urinary bladder of mammals, opens
directly into the cloaca (Fig. A-38). The ducts which drain the developing excretory organs also open into the cloacal
region on either side of the allantoic Stalk. There is at this stage little indication of the formation of the gonads. The
Relation of the sexual ducts to the cloaca can be discerned only by the study of older embryos.
The Circulatory System The 4-day chick embryo possesses two extraembryonic circulatory arcs-the vitelline and allantoic
arcs-in addition to the intraembryonic system of vessels. The heart is already well developed. The pattern of the vitelline
circulation in chicks of 4 days is shown in Fig. A-43. Blood from the dorsal aorta leaves the body via the paired vitelline
arteries, which branch repeatedly and end up as a large capillary network lying on the yolk sac. After picking up nutritive
materials from the yolk, the blood flows into venous collecting channels and finally makes its way either into the marginal
sinus or directly into one of the larger vitelline veins. The proximal portions of the main vitelline veins have fused to form
an unpaired median vessel within the body of the embryo. Through this vessel, the blood returning from the vitelline
circuit eventually reaches the heart. The allantoic circulatory arc is not yet highly developed in 4-day embryos because of
the small size of the allantois. The allantoic arteries arise by means of the prolongation and enlargement of a pair of
ventral segmental vessels arising from the aorta at the level of the allantoic stalk. Their size increases rapidly as the
allantois increases in extent. From them the blood is distributed in a rich plexus of vessels which spread over the
mesoderm of the allantois (Fig. A-45). Later, when the allantois has expanded to meet the chorion as the chorioallantoic
membrane, the allantoic vascular plexus serves as the main site of gas exchange for the embryo. The blood from the
allantois is collected and returned to the heart by way of the allantoic veins, which enter the body of the embryo through
the allantoic stalk (Fig. A-42), course through the lateral body walls (Fig. A-4IE to A-41G), and ultimately empty into the
sinus venous (Fig. A-44). The intraembryonic circulation of the 4-day chick begins with the ventral aorta, which leads into
the series of aortic arches. Aortic arches I and often II have disappeared as main channels, leaving only the third, fourth,
and sixth pairs of arches. The right and left arches are still symmetrical at this time. The regression of the first two aortic
arches plays a prominent role in the formation of important arteries supplying the head-the external and internal carotid
arteries, which are extensions of the ventral and dorsal aortic roots, respec- tively. The dorsal aortae, originally paired
throughout much of their length, are by 4 days fused as far cephalically as the posterior pharynx (Fig. A-41B and C), As
the dorsal aorta extends caudally, it gives off the paired vitelline arteries. Occasionally, early traces of the unpaired coeliac
artery can be seen in 4-day chicks (Fig. A-44), Farther caudally, the paired allantoic arteries branch out toward the allantois
(Fig. A-44). Throughout the length of the aorta, paired segmental arteries supply the somites and their derivatives. The
intraembryonic venous system is represented principally by the anterior and posterior cardinal veins, which drain the
head and trunk and converge into the common cardinal veins as they turn medially to enter the sinus venosus (Fig. A-44).
The posterior cardinal veins lie just dorsal to the mesonephric kidneys throughout their length (Figs. A-39D and E and
A-41E to A-41H). Situated ventrally in the mesonephroi are the small, irregular subcardinal veins (Fig. A-44), which are a
relic of the renal portal system of more primitive ancestral forms. The role of the veins in forming the inferior vena cava is
discussed in Chap. 17 (Fig. 18-12). The early morphogenesis of the heart is summarized in Fig. A-46. By the third and
fourth days, the heart has already twisted upon itself to form a loop and the major gross divisions of the heart are
recognizable. Expansion of the atria is prominent during this period. During the fourth day the truncus arteriosus becomes
closely applied to the ventral surface of the atrium, which expands around the truncus and becomes subdivided into right
and Left chambers. Separation of the common ventricle into right and left ventricles has just begun.

The Urinary System It is possible to recognize pronephric tubules in chick embryos of 38 to 40 hours (Fig. 17-2A). In
chicks of 50 to 55 hours the mesonephric ducts and primordial tubules are clearly identifiable just lateral to the somites
(Fig. A-30D and E). By the fourth day mesonephric tubules are well under way in their development. The mesonephric
tubules, which constitute the excretory units of the perma- nent kidneys, do not appear until considerably later in
development, and the genital organs which become intimately interrelated with the urinary organs are also relatively late
in making their appearance. The later stages of the development of the urinary system as well as the entire sequence of
stages in the formation of the genital organs are covered in Chap. 17.

The Coelom and Mesenteries In adult birds and mammals the body cavity consists of three regions: pericardial, pleural,
and peritoneal. The pleural cavities are paired, each of the pleural chambers being a laterally situated sac containing one
of the lungs. The pericardial chamber containing the heart and the peritoneal chamber contain- ing the viscera other than
the lungs and heart are unpaired. These regions of the adult body cavity are formed by the reshaping and partitioning of
the primary body cavity, or coelom, of the embryo. In the chick the coelom arises by a splitting of the lateral mesoderm of
either side of the body (Fig. A-47A and B). It is therefore at first a paired cavity. Unlike the coelom of some of the more
primitive vertebrates, the coelom of the chick never shows any indications of segmental pouches corresponding in
arrangement with the somites. Instead, the right and left coelomic chambers extend cephalocaudally without interruption
through the entire lateral plates of mesoderm. The coclomic chambers are not limited to the region in which the body of
the embryo is developing. They extend on either side into the mesoderm, which, in common with the other germ layers,
spreads out over the yolk surface. Large parts of the primitive coelomic chambers thus come to be extraembryonic in their
associations (Chap. 7; Figs. 7-1 and 7-4). The portion of the coelom that gives rise to the embryonic body cavities is first
marked off by the series of folds which separate the body of the embryo from the yo!k (Fig. A-47C and D). As the closure
of the ventral body wall progresses (Fig. A-47E and F), the embryonic coelom becomes completely separated from the
extraembryonic. The delayed closure of the ventral body wall in the yolk-stalk region results in the embryonic coelom and
extraembryonic coelom retaining an open commu- nication at this point for a considerable time after they have been
completely separated elsewhere. The same folding process that establishes the ventral body wall completes the gut
ventrally (Fig. A-47C to A-47F). Meanwhile the right and left coelomic chambers are expanded mesiad. As a result the
newly closed gut comes to lie suspended between the two layers of splanchnic mesoderm which constitute the mesial
walls of the right and left coelomic chambers. The double layer of splanchnic mesoderm which thus becomes apposed to
the gut and supports it in the body cavity is known as the primary mesentery. The part of the mesentery dorsal to the gut,
suspending it from the dorsal body wall, is the dorsal mesentery, and the part ventral to the gut, attaching it to the ventral
body wall, is the ventral mesentery. When the dorsal and ventral mesenteries are intact, they constitute a complete
membranous partition dividing the body cavity into right and left halves. The primary dorsal mesentery persists in large
part, but the ventral mesentery soon is extensively resorbed (Fig. A-47H), bringing the right and left coelomic chambers
into confluence ventral to the gut and establishing the unpaired condition of the body cavity characteristic of the adult. In
their relation to the other mesenteries of the body, the mesocardia may be regarded as special regions of the ventral
mesentery. In the most cephalic part of the body cavity, the gut lies embedded in the dorsal body wall instead of being
suspended by a dorsal mesentery as it is farther caudally (cf. Figs. A-21E and A-47F). A ventral mesentery is, however,
developed in the same manner anteriorly as it is posteriorly, and when the heart is formed it is suspended in the most
anterior part of this ventral mesentery. The dorsal and ventral mesocardia may therefore be thought of as the parts of the
primary ventral mesentery lying dorsal to the heart and ventral to the heart, respectively (Fig. A-21D). When the ventral
mesocardium, and a little later the dorsal mesocar- dium, break through, the primary right and left coelomic chambers
become confluent to form the pericardial region of the body cavity (Fig. A-25).