Embryonic Folding

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The placenta | Main Anatomy Index | Heart
development
Last updated 30 March 2006
This page was contributed by David Boshell
Embryonic Folding
At the end of the third week, the germ disc begins to overgrow the
yolk sac, ballooning into a convex shape, with the peripheral areas of the germ disc
becoming the ventral surface of the embryo.
Cephalocaudal folding
Cranial folding begins on about day 21, with cephalic folding
taking place as the neural plate overgrows the yolk sac, whilst caudal folding begins on
about day 23.
The forebrain of cephalic neural plate folds
ventrally, forming the mesencephalic (cranial) flexure in the midbrain region, whilst other cephalic
flexures will appear later.
The cranial region of the embryonic disc, from caudal to cranial contains:
- The buccopharyngeal membrane,
just cranial to the neural plate
- The cardiogenic area, a
horseshoe shaped area cranial and lateral to the buccopharyngeal membrane
- Thickened mesoderm called the septum transversum
that will give rise to portions of the liver and diaphragm.
- The cranial rim of the disc folds under the
cephalic neural plate, wedging the septum transversum
between the heart region and the yolk sac, and placing the cardiogenic area and the
buccopharyngeal membrane in the future thoracic and mouth regions, respectively.
- The caudal rim of the embryonic disc contains:
- The cloacal membrane
- The connecting stalk, which
itself contains the slender allantois.
- Caudal folding brings the cloacal membrane onto the ventral surface of the embryo, with
the embryo rotating ventrally until the
connective stalk lies against the neck of the yolk sac.
Lateral Folding
The lateral edges of
the germ disc bend ventrally, meet and fuse
along the ventral midline from the cranial and
caudal ends toward the neck of the constricting yolk sac, converting the embryo into a
tubular structure with three concentric layers:
- An outer layer of ectoderm
- An intermediate layer of mesoderm
- An inner layer of endoderm
- Thus, ectoderm covers the whole of the embryo except for the closing neuropores and the
ubilicus, from which the connecting stalk and yolk sac neck emerge.
- Fusion of the lateral edges of the endoderm creates the gut
tube, with blind ends in the cranial and caudal regions forming the foregut and hindgut
respectively.
- The midgut is temporarily open to the
constricting yolk sac neck, which will narrow into the slender vitelline
duct by the end of the sixth week.
- The buccopharyngeal membrane, capping the
foregut, ruptures at the end of the fourth week, connecting the oral cavity to the pharynx;
whilst the cloacal membrane, capping the
hidgut, breaks down in the seventh week,
forming the anal and urogenital
orifices.
Formation of the body cavities
The intraembryonic coelom
- When the 2 layers of lateral plate mesoderm meet and fuse, the
space between them becomes a cavity called the
intraembryonic coelom.
- Until there is complete fusion in the ventral midline, the intraembryonic coelom
communicates with the extraembryonic coelom.
- The serosal (mesothelial) membranes
develop to line the body cavity:
- Parietal serosa develops from the
somatopleuric lateral plate mesoderm
- Visceral serosa develops from the
splanchnopleuric lateral plate mesoderm
- The ventrolateral body wall of the embryo is called the embryonic
somatopleure whilst the gut wall of the embryo is called the embryonic splanchnopleure.
- The mesenchyme connecting the gut tube dorsally to the body wall in the abdominal region
disperses, and the overlying serosal layer meets dorsal to the gut tube to become the
double layed dorsal mesentery, suspending the
gut tube in the abdominal cavity.
- Visceral organs are classified in relation to the body wall as follows:
- Organs suspended in the coelom by a mesentery are intraperitoneal
- Organs that develop in the body wall are retroperitoneal
- Organs initially suspended in mesentery that become attached to the body wall are secondarily peritoneal.
Formation of the body cavities
The septum transversum,
placed just caudal to the heart by cephalic folding, initially divides the intraembryonic
coelom into 2 cavities:
- A cranial primitive pericardial cavity,
containing the developing heart
- A caudal peritoneal cavity, containing
developing abdominal and pelvic viscera
- These 2 cavities communicate through a pair of dorsolateral canals called the pericardioperitoneal canals, dorsal to the septum
transversum on either side of the foregut mesenchyme.
- The folding and differential growth of the embryo causes the septum tranversum to
descend from the cervical region to the level of the future diaphragm, carrying with it
its cervical innervation (C3, C4, C5) to form the phrenic nerve.
- During the fifth week, two coronal pleuropericardial folds
develop in the lateral walls of the primitive pericardial cavity, growing medially to
partition the primitive pericardial cavity into:
- The definitive pericardial cavity ventrally,
containing the heart
- A pair of dorsolateral pleural cavities that
will house the lungs, and are still continuous
with the peritoneal cavity through the
pericardioperitoneal canals.
- The bases of the pleuropericardial folds migrate ventrally around the body wall so that
the heart becomes enveloped by the pleuropericardial folds, forming the pericardial
sac.
- During the fifth and sixth weeks, 2 pleuroperitoneal
membranes grow from the walls of the pericardioperitoneal canals to fuse
with the septum transversum, separating the pleural cavities from the peritoneal cavity,
completing partitioning of the intraembryonic coelom.
- Thus, the diaphragm has 4 components:
- The muscle sheets of the pleuroperitoneal membranes
- The central tendon, derived from the septum transversum
- The right and left crura which develop from oesophageal
mesenchyme
- A peripheral ring of muscle from the body wall mesenchyme
adjacent to the developing diaphragm, innervated by spinal nerves T7-T12.
