Gastrulation and Segmentation of

Mesoderm

Dr. AKINGBADE A.M

The most characteristic event occurring during the third week of gestation is
gastrulation, the process that establishes all three germ layers (ectoderm,
mesoderm, and endoderm) in the embryo.

Gastrulation begins with formation of the primitive streak on the surface of the
epiblast Initially, the streak is vaguely defined , but in a 15- to 16-day embryo, it is
clearly visible as a narrow groove with slightly bulging regions on either side.

The cephalic end of the streak, the primitive node, consists of a slightly elevated
area surrounding the small primitive pit . Cells of the epiblast migrate toward the
primitive streak . Upon arrival in the region of the streak, they become flask-shaped,
detach from the epiblast, and slip beneath it . This inward movement is known as
invagination.

Once the cells have invaginated, some displace the hypoblast, creating the
embryonic endoderm, and others come to lie between the epiblast and newly
created endoderm to form mesoderm.
Cells remaining in the epiblast then form ectoderm. Thus, the epiblast, through the
process of gastrulation, is the source of all of the germ layers , and cells in these layers
will give rise to all of the tissues and organs in the embryo. As more and more cells
move between the epiblast and hypoblastlayers, they begin to spread laterally and
cephalad .

Gradually they migrate beyond the margin of the disc and establish contact with the
extraembryonic mesoderm covering the yolk sac and amnion. In the cephalic
direction, they pass on each side of the prechordal plate.

The prechordal plate itself forms between the tip of the notochord and the
buccopharyngeal membrane and is derived from some of the first cells that migrate
through the node in a cephalic direction.
 A. Dorsal side of the germ disc from a 16-day embryo indicating the movement
of surface epiblast cells (solid black lines) through the primitive streak and node
and the subsequent migration of cells between the hypoblast and epiblast (broken lines).
B. Cross section through the cranial region of the streak at 15 days showing invagination
of epiblast cells. The first cells to move inward displace the hypoblast to create
the definitive endoderm. Once definitive endoderm is established, inwardly moving epiblast
forms mesoderm. C. Scanning electron micrograph through the primitive streak
of a mouse embryo showing migration of epiblast (eb) cells. The node region appears
Later, the prechordal plate will be important for induction of the forebrain. The
buccopharyngeal membrane at the cranial end of the disc consists of a small region of
tightly adherent ectoderm and endoderm cells that represents the future opening of the
oral cavity.
Derivatives of the Mesodermal Germ Layer

Initially, cells of the mesodermal germ layer form a thin sheet of loosely
woven tissue on each side of the midline . By approximately the
17th day, however, cells close to the midline proliferate and form a thickened
plate of tissue known as paraxial mesoderm.

More laterally, the mesoderm layer remains thin and is known as the lateral plate. With
the appearance and coalescence of intercellular cavities in the lateral plate, this tissue
is divided into two layers (a) a layer continuous with mesoderm covering the amnion,
known as the somatic or parietal mesoderm layer; and (b) a layer continuous with
mesoderm covering the yolk sac, known as the splanchnic or visceral mesoderm layer.

Together, these layers line a newly formed cavity, the intraembryonic cavity, which is
continuous with the extraembryonic cavity on each side of the embryo. Intermediate
mesoderm connects paraxial and lateral plate mesoderm
.
In the head region, somitomeres form in association with segmentation of the
neural plate into neuromeres and contribute to mesenchyme in the head.

From the occipital region caudally, somitomeres further organize into somites. The
first pair of somites arises in the occipital region of the embryo at approximately the
20th day of development.

From here, new somites appear in craniocaudal sequence at a rate of

approximately three pairs per day until, at the end of the fifth week, 42 to 44 pairs
are present
There are four occipital, eight cervical, 12 thoracic, five lumbar, five sacral, and eight to 10
coccygeal pairs. The first occipital and the last five to seven coccygeal somites later disappear,
while the remaining somites form the axial skeleton .

During this period of development, the age of the embryo is expressed in number of somites.
Table 5.2 shows the approximate age of the embryo correlated to the number of somites.

By the beginning of the fourth week, cells forming the ventral and medial walls of the somite
lose their compact organization, become polymorphous, and shift their position to surround
the notochord .

These cells, collectively known as the sclerotome, form a loosely woven tissue, the
mesenchyme. They will surround the spinal cord and notochord to form the vertebral column
Cells at the dorsolateral portion of the somite also migrate as precursors of the limb and body
wall musculature
 After migration of these muscle cells and cells of the sclerotome, cells at the
dorsomedial portion of the somite proliferate and migrate down the ventral side of the
remaining dorsal epithelium of the somite to form a new layer, the myotome .

The remaining dorsal epithelium forms the dermatome, and together these layers
constitute the dermomyotome .
Each segmentally arranged myotome contributes to muscles of the back (epaxial
musculature, while dermatomes disperse to form the dermis and subcutaneous tissue of the
skin .

Furthermore, each myotome and dermatome retains its innervation from its segment of
origin, no matter where the cells migrate. Hence each somite forms its own sclerotome (the
cartilage and bone component), its own myotome (providing the segmental muscle
component), and its own dermatome, the segmental skin component.

Each myotome and dermatome also has its own segmental nerve component.
INTERMEDIATE MESODERM
Intermediate mesoderm, which temporarily connects paraxial mesoderm
with the lateral plate, differentiates into urogenital structures.
In cervical and upper thoracic regions, it forms segmental cell cluster
(future nephrotomes), whereas more caudally, it forms an unsegmented
mass of tissue, the nephrogenic cord.
Excretory units of the urinary system and the gonads develop from this
partly segmented, partly unsegmented intermediate mesoderm.
Lateral plate mesoderm splits into parietal and visceral layers, which line the
intraembryonic cavity and surround the organs, respectively. Mesoderm from the
parietal layer, together with overlying ectoderm, will form the lateral and ventral body
wall. The visceral layer and embryonic endoderm will form the wall of the gut.

Mesoderm cells of the parietal layer surrounding the intraembryonic cavity will form
thin membranes, the mesothelial membranes, or serous membranes, which will line
the peritoneal, pleural, and pericardial cavities and secrete serous fluid.

Mesoderm cells of the visceral layer will form a thin serous membrane around each
organ