BIO 1201 LECTURE 39

Cleavage
Approximately 24 hours after fertilization the impregnated oocyte begins with the first cleavage
division.
The morula, a collection of around 30 cells (blastomere), is created at about 96 hours. Because
these cells arise only through the cleavage of the zygote and all are found inside the pellucid
zone, which cannot expand, no growth is seen. Every new cell is thus only half as large as the
cell from which it derives. The name of this stage comes from its resemblance to a mulberry,
since it really looks like a collection of spherical cells.
The cleavage divisions of the individual cells do not have to happen synchronously and thus
phase shiftings can occur.
On the 4th day after insemination the outermost cells of the morula that are still enclosed
within the pellucid zone begin to join up with each other (so-called compaction). An epithelial
cellular layer forms, thicker towards the outside, and its cells flatten out and become smaller.
The cells contact one another by means of tight junctions and gap junctions. A cavity forms in
the interior of the blastocyst into which fluid flows (the so-called blastocyst cavity). The two to
four innermost cells of the preceding morula develop into the so-called inner cell mass of the
blastocyst. The actual embryo will develop solely from these cells (embryoblast). These cells are
concentrated at one pole, the embryonic pole of the blastocyst. What has thus been formed is
an outer cell mass (the trophoblast), consisting of many flat cells, and the embryoblast, formed
from just a few rounded cells. From the trophoblast the infantile part of the placenta and the
fetal membranes will arise.
Around the end of the fifth day the embryo frees itself from the enveloping pellucid zone.
Through a series of expansion-contraction cycles the embryo bursts the covering. This is
supported by enzymes that dissolve the pellucid zone at the abembryonic pole. The rhythmic
expansions and contractions result in the embryo bulging out of and emerging from the rigid
envelope. This "first birth" is called hatching.
The volume of an embryo begins to grow as soon as the blastocyst cavity within the morula
starts to form.
Organogenesis
- the process by which the ectoderm, endoderm, and mesoderm develop into the internal
organs of the organism.
The facial region develops ventral to the rostral neural tube from parts of the notochord, and
the pharynx, which is surrounded on both sides by a series of aortic arches. Between these
structures and the ectoderm there are a large number of neural crest cells (neurectodermal
origin) and mesenchyma cells, which stem from the mesoderm. They have a tendency for
segmental ordering and form the various sections of the facial anlage and the pharyngeal
arches. The face and the jaw stem from an unpaired forehead prominence (frontonasal
process) and on both sides from the maxillary and mandibular processes, which stem from the
first pharyngeal arch. Through differing growths of the individual processes the frontonasal
process forms the upper half of the face (frontal process) and the nose while the middle part of
the upper jaw and lip are formed by the medial and lateral nasal processes. The maxillary
process fuses with the medial nasal process and forms the lateral part of the upper jaw and lip.
The mandibular process forms the lower jaw, lip and the chin. Between the lateral nasal
process and the maxillary process arises the nasolacrimal duct that connects the orbit with the
nasal cavity.

The palate arises through the fusion of the unpaired median palatine process (primary palate)
with the two lateral palatine processes (secondary palate).

The nose arises from the two nasal placodes in the frontonasal prominence. They subside and
form two nasal sacs that are outwardly delimited on both sides by the lateral and medial nasal
processes (the first forming later the nasal wings and the second the nasal septum). The nasal
sacs (primary nasal cavity) open into the oral cavity via the posterior nasal orifices. Only
somewhat later do these two nasal cavities widen towards the rear. They remain separated
from each other in the middle by the nasal septum. The secondary palates separate them from
the primary oral cavity.

The teeth form through the interaction between dental lamina (ectodermal ridge) in the upper
and lower jaws and neural crest tissue. Initially they are cap-shaped and later bell-shaped. The
ectodermal portion forms the enamel organ (ameloblasts) that forms the hard tooth enamel
layer. In the interior the odontoblasts form from the neural crest tissue, which is responsible for
dentin production. The salivary glands also stem from ectodermic sproutings.

The tongue arises from various anlagen in the pharyngeal floor region. From this the
complicated innervation pattern of the tongue can be explained. The lingual musculature stems
from the occipital somites that migrate into the tongue (lingual cord) with the glossopharyngeal
nerve (cranial nerve XII).

In their interior all pharyngeal arches are delimited by the pharyngeal pouches and on the
outside by the pharyngeal clefts. They form many of the various structures in the neck region.
From the 1st pharyngeal arch arise the upper and lower jaws. With mesenchyma of the 1st and
2nd pharyngeal arches that surrounds it, the 1st pharyngeal pouch and cleft forms the
numerous small parts of the middle ear and the external acoustic meatus, respectively.
The 2nd, 3rd and 4th pharyngeal clefts obliterate and form the surface of the neck.
Components of the 2nd to 4th pharyngeal arches form the skeleton, muscle and connective
tissue portions of the neck.
Finally, from the 3rd and 4th pharyngeal pouches arise on both sides portions of the thymus,
the parathyroid and the ultimopharyngeal body (only from the 4th pharyngeal pouch). The
thyroid arises as an unpaired ventral sprouting from the upper pharynx (later base of the
tongue) region (foramen cecum).

The intestines arise from a tube that forms from the endoderm. Cranially this tube ends at the
oropharyngeal membrane and caudally at the cloacal membrane. It is subdivided into a foregut,
midgut and hindgut. The midgut is connected with the umbilical vesicle. The differing
development of the various intestinal sections is based on the local interactions with the
surrounding mesenchyma (epithelio-mesenchymal interactions). Thus in the uppermost
section, beside the thyroid, the primordia of the respiratory system, the pancreas, liver and gall
bladder form as ventral sproutings.

The esophagus remains as a tube, obliterates, though, during the course of the embryonic
development and only in the fetal period does it undergo a recanalization. It lengthens
considerably, just like almost all intestinal sections, with the growth of the embryo. The
stomach appears quite early as a spindle-shaped widening. It is hung on a dorsal and ventral
mesogastrium and through rotation reaches its adult position. Due to its large increase in
length the midgut forms transiently a loop into the umbilical coelom (physiologic umbilical
hernia). With the straightening up of the embryo in the late embryonic period the intestinal
loops can move back again into the abdominal cavity, but experience a rotation (270 degrees)
about their own axis. This leads to the characteristic arrangement of the colon around the loops
of the small intestine. Parts of the mesenterium also coalesce later with the posterior
abdominal wall so that the ascending and descending colon lies secondarily retroperitoneal.
Just as with the esophagus the lumen of the small intestine also obliterates for a certain time in
the early fetal period in order to be recanalized again later.

The pancreas, liver and gall bladder form as sproutings from the intestine. The primordium of
the liver arises at the level of the transverse septum. Through the enormous increase in size it
extends, though, into the abdominal cavity but remains still connected with the transverse
septum (diaphragm) by the area nuda. The liver primordium, together with the
omphalomesenteric vessel, form the complicated sinusoid system of the liver. The pancreas
arises from two components, the ventral and dorsal pancreases. Through a shifting of the
ventral part around the duodenum the two join and form dorsally the definitive pancreas in the
mesogastrium, which somewhat later, adheres to the posterior abdominal wall. Through the
coalescence of the mesentery in this region the pancreas finally also comes to be positioned
secondary retroperitoneal.

In the entire intestinal tract many abnormalities can arise. They range from stenoses, atresias,
duplications, fistulas, diverticula to abnormal rotations. Genetic disorders can also be
responsible for malformations in this region.
The urinary tract develops from the 3rd week of the embryonic period from the intermediate
mesoderm as well as from the urogenital sinus. The kidneys develop from the 4th week in three
steps: As a first one, a cranial anlage, the pronephros, forms that then later atrophies in the 8th
week and is never active functionally. It is followed by a further anlage from the intermediate
mesoderm, the mesonephros, that is formed between the 6th and 10th weeks, but is only
transitory, and the anlage of the definitive kidneys, the metanephros. They develop from a
metanephrogenic anlage (mesodermal origin) and the ureter anlage (that has its origin in the
caudal part of the wolffian duct)

The urine-excreting part of the kidneys, the nephron, mainly arises from the metanephrogenic
anlage (glomerulus, proximal, intermediate and distal tubules), while the rest of the upper
urinary tract (collecting ducts, calices, renal pelvis and ureter) develop from the ureter anlage.

The lower urinary tract differentiates from the cloaca between the 5th and 8th weeks in that it
becomes subdivided by the urorectal septum. The ventral part of the cloaca forms the primary
urogenital sinus, out of which the urethra forms in the lower part and the bladder in the upper
part. The ureter anlage discharges into the upper posterior wall of the urogenital sinus. In
males, the wolffian duct remains present and forms a connection to the genital tract in the
lower part of the urogenital sinus.

The numerous induction mechanisms between ureter anlage and metanephric mesenchyma
during the development of the renal system, as well as the ascent of the kidneys, originating at
the level of the sacrum and moving up to the diaphragm at the end of the development, make
it possible for a large number of abnormalities to arise. Many remain asymptomatic whereas
others are not compatible with survival.