SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

A self-study of early embryology:

fertilization, implantation, gastrulation and the embryonic period

Embryology, in simple words, is the study of life before birth. It explains how structures in the body come to be
what they are and explains their interrelationships. It holds the keys to where development may go wrong, resulting
in anatomical defects, malfunction and biochemical abnormalities; all under the umbrella term congenital errors.
This is an area of medical specialty, but even to non-medical persons, it can explain many common phenomena
that we encounter in this field.
Fertilization is the meeting of male and female germ cells. The single cell thus formed divides to form a mass
which is embedded in the wall of the mothers uterus, a process called implantation. The second week of
development leads to the formation of a bilaminar (two-layered embryo). The third week of development
is characterized by the formation of the three-layered embryo (gastrulation). During weeks three to eight,
the embryonic period, the flat three-layer disc folds in a complex manner and forms a tube. This is followed by the
development of the organs and systems of the body. Within these few weeks the embryo takes on the human form.
At the end of this stage the organs systems are not necessarily in a fully functional state.
The events of life before birth take place in the mothers reproductive system. We therefore begin with a review of
the female reproductive system and the formation of germ cells. Fetal development is a complex occurrence with
multiple events occurring concurrently. For simplicity, in this synopsis we will study each of those events
independently. Realize however that these are snapshots of isolated events among many simultaneously activities.

Readings:
Langmans Medical Embryology, 12th edition, by TW Sadler
Chapters 3, 4, 5 & 6
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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

Female Reproductive Anatomy

The female reproductive organs consist of the vagina,
the uterus, the fallopian tube and the ovary.
The ovary (1) is the egg-producing organ. The
uterine tube (also called the oviduct or fallopian tube)
carries the egg to the uterus. The end of the tube facing
the ovary has a number of fimbriae (singular:
fimbria). The fimbriae sweep the surface of the
ovary and pick up the ocyte. The next part is called
infundibulum (2), followed by a dilated portion, the
ampulla (3). Next to it is a narrow portion, the
isthmus (4). The last part is within the thick wall of
the uterus, and is called the intramural part (5).
In the uterus, the great thickness of the wall is largely
due to muscle. The lining is formed by epithelium and
supporting connective tissue. This lining is the
endometrium (metros = uterus). The embryo, when it
reaches the uterus, is embedded in the endometrium.

Anatomically, the top of the uterus is the fundus, followed by the body and the narrow cervix (= neck). The
vagina is the birth passage with a muscular wall.

Germ cells (Gametes).

The male and female germ cells are quite distinctive. In biological terminology we use the common term
gamete for both male and female germ cells. The male gamete is a spermatozoon, sperm for short.
The female gamete is an ovum. It is often called the egg. It is one of the largest cells in the body with a diameter
of over 100 m. Henceforth we avoid the term germ cell and use the term gamete.
Gametes have 23 single chromosomes, as opposed to 23 pairs in other cells. The reduction in the number is due to
meiotic cell division (meiosis).

SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

Meiosis
Meiosis is a process involving two divisions. The first division (Meiosis I) has two features of importance.
1. Replicated homologous chromosomes (maternal and paternal chromosomes of the same pair) are arranged next
to each other as if in an embrace. Their corresponding arms cross at some points. At the points of crossing, they
exchange equivalent parts of their lengths that is, they exchange genetic material. This process is called
crossing over. When they separate, each of the homologous chromosomes has a mixture of paternal and maternal
genetic material. The exact lengths and therefore the proportion of material exchanged differs from pair to pair, and
even for the same pair in different divisions. This means that each chromosome, after crossing over, is a unique mix
of maternal and paternal genes.
2. The chromatids do not separate during meiosis I. Instead, replicated chromosomes move to opposite ends of the

cell, one member of each pair moving to one end, the other member to the other end.

At the end of meiosis I, there are two daughter cells, each with a haploid number (1N) of replicated
chromosomes
In meiosis II, the two cells divide. The chromatids of the single replicated chromosomes separate. This may sound
like mitosis, but remember that it is a single replicated chromosome that splits, and that the single chromosome
has a mixture of genetic material from the two homologous chromosomes.
.

At the end of meiosis I I, there are tour daughter cells, each with a single haploid (1N) chromosome
To summarize:
Meiosis comprises two divisions.
The end products are four haploid cells.
The unpaired chromosomes differ from each other in the mixture of parental genetic material.
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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino
Note : This discussion is limited to understanding the ocyte, its haploid nature and fertilization.

In the male, some cells are set aside in the testis for this purpose of division. These are called spermatogonia.
After puberty, these cells divide by mitosis, some of the daughter cells undergo meiosis while others stay as
spermatogonia to maintain the pool. This process continues throughout life. Thus, a male can produce sperms
indefinitely.
In contrast, the cells set aside in the female form a fixed pool. They begin meiotic division even before female
is born. But this division is not completed a female is born with a fixed number of such cells, in a state of
suspended meiosis. After puberty, approximately every month a few of these cells (ocytes) mature and proceed
with meiosis. Normally only one of these is released for fertilization by a sperm. This cell completes the second
stage of meiosis after the entry of the sperm. Therefore we say that it is an oocyte that is released for
fertilization.

An ocyte has a non-cellular covering in addition

to the cell membrane. This covering is the
zona pellucida (the clear zone, so called
because of its translucent pink appearance under
the microscope. Outside the zona pellucida is a
layer of supporting cells. One layer of these
supporting cells surrounds the ocyte even as it is
released. This layer appears like a radiating
crown in a section and is called the corona
radiata. During each stage of meiosis two cells are
produced. However, one of these receives almost
all cytoplasm, is large and capable of being
fertilized. The other small cell remains within the
zona pellucida and is called the polar body. There
can be two or three polar bodies.
Understand the relationship of the ocyte
with the zona pellucida and the corona radiate.

SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino
Fertilization the beginning

Sperms are deposited deep into the vagina just below the
opening of the uterus. Though sperms are motile, their
speed is far from adequate to carry them to the ocyte in time.
The ocyte has a long journey along the length of the
fallopian tube. Other factors operate in transporting sperms
to the ocyte, probably movements in the smooth muscle wall
of the uterus.

The nucleus and the scanty cytoplasm of the sperm are all in the
head of the sperm. At the very tip is a tiny structure called
acrosome. (akron = highest point, soma = body). The acrosome
(shown as a green line in the adjacent image) has enzymes
which allow it to penetrate the zona pellucida. In the female
reproductive tract, the acrosome undergoes changes which allow
the acrosomal enzymes to be exposed. This is called capacitation.
Fertilization takes place most commonly in the dilated part of the
uterine tube, the ampulla. When a sperm comes in contact with
the zona pellucida, the acrosomal enzymes act on it and facilitate
entry of the sperm.

The cell membranes of the sperm and the ocyte fuse and the
sperm enters the ocyte. The zona pellucida undergoes
molecular changes which prevent the entry of any other sperm.
The fertilized ocyte immediately completes meiosis II and the
second polar body is released.

SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

The fertilized cell (also called the zygote) now has two
haploid nuclei. These are called the male and female
pronuclei. The pronuclei enlarge as the DNA within them is
replicated. The nuclear envelopes disappear and the 23 pairs
of replicated chromosomes (diploid or 2N) are lined up for
separation. Depending upon the sex chromosome present in
the sperm, the sex of the individual is determined at
fertilization. The ocyte can only have an X chromosome, the
sperm can have either X or Y.

The embryo becomes multicellular.

The zygote now undergoes the first series of mitotic divisions increasing the number of cells. These divisions, called
cleavage, are preceded by DNA replication, but the cytoplasm does not increase. Remember: the ocyte has an
enormous amount if cytoplasm. Thus, gradually the amount of cytoplasm in the zygote is distributed among the
cells. The nucleo-cytoplasmic ratio comes down to normal.
The cells, which become smaller with each cleavage division, are known as blastomeres. After a number of
cleavages the blastomeres maximize their contact forming a compact clump of loose cells.

All this while the embryo is being propelled through the uterine tube
towards the uterus. After the 16th cell stage, at approximately 3 days,
the cell mass resembles a mulberry and is called morula. The zona
pellucida is still intact when it enters the uterus. This is important,
because it prevents the attachment (implantation) of the embryo to the
wall of the uterine tube.

SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

Blastocyst formation (Days 8-12)

The cells of the morula are initially a
loose clump. They gradually get
compacted, and after a while show two
distinct regions, an inner cell mass and
an outer cell mass, a process known as
compaction. The inner cell mass
eventually develops into the embryo
proper. The outer mass is called the
trophoblast (nutrition forming). It is
somewhat like a shell around the inner
cell mass. It gives rise to the placenta
which is the link between the
embryo/fetus and the mother.

About the time the embryo enters the

uterine cavity, fluid seeps in through the
zona pellucida into the intercellular spaces
of the inner cell mass forming a cavity
called the blastocele within the embryo.
At this stage t h e e m b r y o is called the
blastocyst, consisting of the compact
inner cell mass, the embryoblast , the
outer cell mass, the trophoblast, and the
blastocyst cavity.

SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

Implantation and Formation of the Placenta

(for the purposes of this narrative the

changes in the embryoblast will not be discussed until later in the discussion)
The wall of the uterus has an outer layer
of muscle and an inner layer, the
endometrium which is comprised of
connective tissue and epithelium.
The endometrium has a rich vascular
supply.
Implantation begins at the end of the first
week and is complete in the second week.

The cells of the trophoblast flatten and

form the epithelial wall of the blastocyst.
The zona pellucida degenerates,
allowing implantation to begin. The
trophoblast has an invasive property.
When it comes in contact with the
endometrium, it burrows into the
endometrium by loosening the
epithelium and destroying the connective
tissue. Gradually the embryo will sink
deeper into the endometrium until finally
it is covered by the uterine epithelium,
completing this process called
implantation.

As the blastocyst embeds in the

endometrial stroma, the trophoblast
differentiates into two layers: an inner
layer of mononucleated cells, the
cytotrophoblast and an outer
multinucleated zone without distinct
boundaries, the syncytiotrophoblast.
These cells are collectively termed
the syncytium (a multinucleated mass
of cytoplasm that is not separated into
cells).
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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

As the blastocyst penetrates the

endometrium, maternal capillaries
within the endometrium enlarge
and are become filled with
maternal blood forming sinusoids.
The blastocyst is more deeply
embedded in the endometrium and
the penetration defect in the
surface epithelium is closed by a
fibrin coagulum (plug). At the
embryonic pole vacuoles appear in
the synctium. When these vacuoles
fuse they form large lacunae,
called trophoblastic lacunae.

The cells of the

syncytiotrophoblast penetrate
deeper into the endometrial
stroma and erode the
endothelial lining of the
maternal sinusoids. The
trophoblastic lacunae become
continuous with the material
sinusoids and maternal blood
enters the synctiums lacunae
system. The trophoblast is
now characterized by villous
sinusoids.

Further differentiation of the placental villi will be covered later in this synopsis.

SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

Changes in the inner cell mass (embryoblast).

Recall, the cells of the embryonic mass are initially a
loose clump, called a blastocyst. During the second
week of development, they gradually get compacted,
and after a while show two distinct regions, an inner
cell mass and an outer cell mass.

The inner cell mass eventually

develops into the embryo proper,
thus is termed the embryoblast.

Formation of the Bilaminar Embryoblast

By the end of the second week the
embryonic development, cells of the
inner cell mass or embryoblast
differentiate into two layers a layer
of tall columnar cells, the epiblast
and one of smaller cuboidal cells, the
hypoblast.
The embryo is now a bi-laminar disc
(bi = two, lamina = plate or layer)
composed of the epiblast above and
the hypoblast below.
Concurrently, a small cavity appears
in the embryoblast. This is the
amniotic cavity. The cells lining
this cavity, facing the cytotrophoblast
are called amnioblasts. This
complex is termed the amnion.
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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino
At the time the bilaminar disc is
forming, cells from the hypoblast
migrate and form a thin layer of
cells that expands towards the
cytotrophoblast. This is the
exocoelomic membrane, known
as Heusers membrane. This
layer forms the lining of the
exocoelomic cavity which will
later become the primitive yolk
sac.

At the end of the second week the

embryo is a bi-laminar disc,
surrounded by above and below
by cavities. The junction between
the cavities is compressed. This
is the embryo proper (epiblast and
hypoblast). Think of this
arrangement as two soap bubbles
touching each other. The junction
between the bubbles is flat, the
bilaminar embryoblast. Above
(top bubble) is the anmion and
amniotic cavity; below (lower
bubble) is the primitive yolk sac.

Heusers membrane continues to

expand. It approaches the inner surface
of the cytotrophoblast, enlarging the
exocoelomic cavity, which now called
the primitive (primary) yolk sac.
The amniotic cavity is situated above
around the cells of the epiblast and is
lined by the surrounding amnioblasts.

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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

Concurrent with the formation of the primitive

yolk sac, some of the migrating hypoblast
cells trans-differentiate into mesenchymal
cells that fill the space between Heuser's
membrane and the cytotrophoblast, forming
the extraembryonic mesoderm.

As development progresses, small lacunae

begin to form within the extraembryonic
mesoderm. These lacunae expand and
separate Heusers membrane from the
cytotrophoblast. This cavity enlargers and
forms the extraembryonic cavity or coelom.
It divides the extraembryonic mesoderm into
two layers: extraembryonic splanchnic
mesoderm, which lies adjacent to Heuser's
membrane around the outside of the primitive
yolk sac, and extraembryonic somatic
mesoderm, which lies adjacent to the
cytotrophoblast layer of the embryo.

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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

With time the yolk sac shrinks away from the

cytotrophoblast and giving rise to what is
termed the secondary (definitive) yolk sac,
lined by hypoblast cells and extraembryonic
splanchnic mesoderm.
As the amnion increases in size, it begins to
expand and form a cleft between the amniotic
cavity and the yolk sac. This space is the
intra-embryonic cavity, as it will eventually
be the inside of the embryo.
The extra-embryonic cavity and the intraembryonic cavity are continuous at this stage
and freely communicate.

The extra-embryonic cavity will later become

the chorionic cavity.
The extraembryonic somatic mesoderm lining
the inside of the cytotrophoblast becomes the
chorionic plate.
Condensations of the extraembryonic
mesoderm attach the developing embryo to the
chorionic plate and ultimately the placenta.
With the development of blood vessels this
becomes the umbilical stalk.

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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

Development of the placenta villi:

The following is an overview of aspects of the basic fetal subunit of the placenta - the placental villi. In early
placentation, the villi proceeds through a similar initial program of development. In later placentation, villi
morphologically differentiate into a range of villi in accordance with functional changes reflecting their
specialization. The major initial contribution is from the trophoblast shell that surrounds the fetus and later by the
development of extraembryonic mesoderm and blood vessel differentiation.

The cells of the cytotrophoblast

proliferate locally and penetrate into the
syncytiotrophoblast, forming cellular
columns surrounded by the syncytium.
These cellular columns with the syncytical
(syncytiotrophoblast) covering are known
as the primary villi of the placenta.

During further placenta development

extraembryonic mesodermal from the chorionic
plate penetrate the core of the primary villi and
grow towards the decidua of the endometrium
(secondary villi). Some mesodermal cells in the
core of the villus differentiate into blood vessels and
cells, forming the villous capillary system known as
tertiary or definitive placenta villi.

Finally, at about the end of the third week, cytotrophoblastic cells

penetrate progressively into overlying syncytium until they reach the
maternal endometrium. There they establish a thin outer
cytotrophoblastic shell which attaches to the maternal endometrial
tissue. Spiral arteries from the material vessels form connections
between the maternal capillaries in the endometrium and the
intervillous spaces of the syncytium. The intervillous space is lined
by a thin layer of syncytiotrophoblast cells and the cytotrophoblastic
cells of the villi. The outer shell of cytotrophoblast becomes the
placenta barrier.

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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

Gastrulation: Formation of Ectoderm, Mesoderm and Endoderm

The primitive streak

The most characteristic event during the third week is Gastrulation. Gastrulation begins with the formation of the
primitive streak on the surface of the epiblast. Looking down at the epiblast, the primitive streak is seen as a faint
midline ridge on the epiblast side of the embryo. At one end the streak has a knot-like swelling, the primitive
node. This establishes the axes of the embryo. The part of the embryo beyond the node will be the cephalic (head
end) of the embryo. The other end, near the narrow tip of the streak is the caudal (tail end) of the embryo.
The primitive streak and node are temporary structures that play vital role in establishing the laterality (left and
right sidedness) of the embryo. Fibroblast growth factor 8 (FGF8), secreted by the primitive node establishes the
expression of Nodal, a secretory protein that belongs to the Transforming Growth Factor (TGF-beta) superfamily.
Nodal initiates and maintains the primitive streak. Once the streak is formed the nodal protein accumulates on the
left side of the embryo and up regulates a number of genes (Lefty and PITX1) which are responsible for the
establishment of left sidedness of the embryo. Importantly, the neurotransmitter serotonin (5HT) plays a role in
the signaling cascade that establishes laterality. 5HT is concentrated on the left side and is upstream from FGF8.
5HT is broken down by its metabolizing enzyme monoamine oxidase MAO) on the right side.

Formation of the germ layers

The head end of the embryo begins to differentiate first. The sequence of development is often described a
cephalocaudal. (From head to tail. kephale = head, cauda = tail).

The cells of the epiblast proliferate and migrate towards the primitive
streak. Upon arrival at the region of the primitive streak they enlarge,
detach from the epiblast and slip beneath the primitive streak. This inward
movement of the modified epiblast cells is known as invagination.
Migration and specification of the cells of the epiblast is controlled by
fibroblast growth factor 8 (FGF8).

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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

Once the cells have invaginated into the

primitive streak, they sink deeper inside
where they displace the hypoblast laterally,
giving rise to what will become the
endoderm. The endoderm gives rise to the
epithelium of the digestive system and
respiratory system.

With the formation of the endoderm, more

cells proliferate and pour between the
epiblast and the newly formed endoderm
giving rise to the intra-embryonic
mesoderm. The intra-embryonic mesoderm
gives rise to mesenchyme (connective tissue).
Cells remaining in the epiblast then form the
ectoderm. Ectoderm will give rise to
epidermis and related structures as well as
portions of the brain and nervous system.

Expansion of the Mesoderm

Mesodermal cells pouring into

the primitive streak spread out
laterally and towards the head end
in a definite pattern. Migrating
laterally and cranially the
mesoderm fills the gap between
the ectoderm (epiblast) and
endoderm (hypoblast).

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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

Formation of the Notochordal Process

Prechordal plate
Prenotochordal cells are a group of specialized cells
that migrate through the primitive node located at the
cranial end of the primitive streak, and give rise to the
prechordal plate and notochordal process. The cells
migrating most anteriorly form the prechordal plate,
whereas the ones migrating most posteriorly form the
notochordal process. Prechordal plate cells participate
in the formation of the endodermal layer of the mouth.
In this area the ectoderm is attached directly to the
endoderm without intervening mesoderm. This area
is known as the oropharyngeal membrane, and it
will become the mouth.

Notochordal process

Additional prenotochordal
cells invaginating into the
primitive node move
forward cranially in the
midline behind the
prechordal. This layer forms
a tubular mass between the
epiblast and hypoblast
known as the notochordal
process. The prenotochordal
cells of the notochordal
process become intercalated
in the developing hypoblast
(future endoderm): (see
below).This will become the
future notochord.

As the intra-embryonic mesoderm spreads out from the primitive streak, the whole embryo increases in size and the
primitive streak becomes relatively smaller. When the process of gastrulation is complete the primitive streak
disappears.
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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

Formation of the Notochord

The notochord is vital link in the chain of events. In vertebrates, it exists as a continuous column only during
embryonic life. Parts of it persist as normal structures in the vertebral column.
Embryologically the notochord is of great importance. After the primitive streak disappears, it is the definitive
axis of the body. Moreover, it directs a strip of ectoderm to develop into the neural plate, the precursor of the
nervous system.

The notochordal process becomes intercalated within the developing endoderm. As more of the hypoblast is
replaced by endodermal cells, the tubular notochordal process fuses with the endodermal layer beneath it and is
morphologically modified from a tube shape to a flattened plate shape, the notochordal plate.

The notochordal plate then detaches from the endoderm and rolls into a solid mass of cells, which migrates into
the intraembryonic mesoderm. This mass of cells is now the definitive notochord.

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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

Neurulation (Formation of the neural tube)

Neurulation is the process of conversion of the neural plate to form the neural tube.
A band of ectoderm across the midline
becomes thickened to form a plate called
the neural plate. Since it forms nervous
tissue and it is a part of ectoderm, it is
called neuroectoderm. As mentioned
prior, the notochord induces the ectoderm
in the formation and subsequent
modification of the neural plate.

By the third week the lateral edges of the

neural plate become elevated to form
neural folds and the depressed mid-region
becomes the neural groove.

Gradually the neural folds approach each

other turning the neural groove into a
deeper tube-like depression.

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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

Soon, the groove closes as the neural

folds come together to form the
primitive neural tube. The edges of
the neural folds fuse. The fusion
begins in the middle of the embryo and
proceeds towards both ends. As
closure progresses there are cranial
and caudal neuropores.

The neural tube sinks in the

surrounding mesoderm, losing its
contact with the ectoderm on the
surface. The surface ectoderm
becomes continuous across the
midline when the neural tube closes.
The neural tube forms the brain
and the spinal cord

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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

Neural Crest
The main derivatives of the neural crest are the ganglia of peripheral nervous system and sensory nerve fibers.
The neural crest also gives rise to a number of non-neural structures. These will be referred to as we come across
them.

Neural crest cells are specified at the

border of the neural plate. During
neurulation, the borders the neural folds
converge at the dorsal midline to form the
neural tube. The lips of the groove
separate out as a long column on either
side of the tube. These form the neural
crest, which also sinks deeper along with
the tube.

Subsequently, neural crest cells from the

roof plate of the neural tube undergo an
epithelial to mesenchymal transition,
delaminating from the neuroepithelium.

These cell clusters detach from the

neurotube and migrate through the
developing embryo where they
differentiate into varied cell types.

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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

The spread of the mesoderm.

Mesodermal cells spread out laterally and towards the head end in a definite pattern. They follow arc-like courses
between the ectoderm and the endoderm. When the spread is complete the mesoderm is seen mainly as three
columns along the length of the embryo.
Cells from the head end of the streak keep
close to the midline, spreading in narrow
arcs. In the final picture, this column of
mesoderm is close to the midline or the
axis. This column is the paraxial (para +
axial) mesoderm.
Cells from the middle part of the streak
occupy a position just lateral to the
paraxial mesoderm. They form a column
called the intermediate mesoderm.
Cells from the tail end (caudal part) of the
streak spread out in broad arcs and finally
form a plate close to the lateral boundaries
of the embryo. This part of the mesoderm
is called the lateral plate mesoderm.
Some cells from the region near the head
end are special they reside at the cranial
tip of the lateral plate and indeed form a
horseshoe-like band at the head end of the
embryo (heart field). These cells form the
mesodermal precursor of the heart. This
will be further discussed with the
development of the heart.
The spread of the mesoderm to the head end, however, leaves a small area where ectoderm and endoderm are in
contact. This is called the oropharyngeal membrane. At the tail end there is a similar small area where ectoderm
and endoderm are in contact with no mesoderm in between. This is called the cloacal membrane.

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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

Divisions of the mesoderm.

The three divisions are shown as above are completely separate from each other with gaps in between. This is for
the sake of clarity in the picture! However, the fates of these divisions are quite specific.

At the beginning of the third week, the column of

paraxial mesoderm begins to form segmental
masses called somites. Somites appear as two
columns of mesodermal masses adjacent to the
notochord.
The formation of paraxial mesoderm into somites
begins at the head end and progresses caudally.
The sequence of development is cephalocaudal
(head to tail). As more and more somites
appear towards the tail end, the cephalic somites
begin develop into various structures.
These segments (slices or blocks) are the
basis of the structure of the vertebrae and dermis,
connective tissue & musculature of the body wall
and limbs.
The intermediate mesoderm is mainly
involved in the formation of the urinary and
reproductive systems.

The lateral plate mesoderm will split

into two parts. One layer of mesoderm
follows the ectoderm (somatic
mesoderm) and the other (splanchnic
mesoderm) migrates towards the
endoderm. During further development
the somatic mesoderm will give rise to
body wall connective tissue structures,
while the splanchnic mesoderm will
give rise to the mesenteries of the gut
tube.

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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

Folding of the embryo

The flat trilaminar embryonic disk becomes more a cylindrical embryo due to the longitudinal and transverse
folding that occurs as a result of embryonic growth. The folding occur simultaneously and are not separate
sequential events.
The longitudinal fold or cephalic-caudal fold creates a
cranial and caudal region of the embryo. This is caused
by the rapidly developing neural tube. Essentially the
head and tail of the embryo fold towards each other.

Transverse fold
At three weeks the embryo is a slightly curved flattened
disk. The lateral plate mesoderm has separated into
somatic (following the ectoderm) and splanchnic (around
the yolk sac) mesoderm. The space or cavity between the
two layers is called the intraembryonic coelomic cavity. .

The transverse fold (called flexion) produces right and

left lateral folds. Flexion, a process of curving, transforms
the embryo into a sort of "tube". The left and right sides of
the embryo curve and migrate toward the mid-line.
The intraembryonic coelom communicates freely with the
extra-embryonic coelom.

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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

The combination of somatic mesoderm and adjacent

ectoderm, (termed the somatopleure) along with the
overlying amniotic membranes and cavity, folds toward
the midline, rolling the edges of the embryonic disk
ventrally to form a cylindrical embryo.
As the sides come together, they pinch off the yolk sac
drawing the splanchnopleure (the combination of the
splanchnic mesoderm and the adjacent endoderm) into the
intraembryonic coelom.

As lateral and ventral body walls form, part of the yolk sac
is incorporated into the embryo as the midgut;
simultaneously, the connection of the midgut with the yolk
sac is reduced to a yolk stalk or vitelline duct (the
connection between the yolk sac and the primary intestinal
loop of the midgut).

With complete closure the intra-embryonic coelom or

cavity is closed off. The mid-gut is incorporated into the
intra-embryonic coelom surrounded by splanchnic
mesoderm. The outer periphery of the intra-embryonic
cavity is lined by somatic mesoderm.
The amniotic cavity enlarges and obliterates the extraembryonic coelom.

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SELF STUDY: EARLY EMBRYOLOGY

Dr. Leon J. Martino

SUMMARY
Do not try to memorize this list! Understand the explanations from the discussions above.

Concepts and facts fundamental to study of the embryology of the human structure:

Haploidgametesunite:beginningofanewhumanbeing.

Eachgametehasauniquemixtureofgeneticmaterialfromtwoparents.

Fertilizationmostcommonlytakesplaceintheampullaoftheuterinetube.

Zona pellucida prevents the entryof more than one sperm. It also prevents implantation aslongas
itisintact.

The early embryonic cell mass specializes into two groups: the inner cell mass embryoproper,
andtheoutercellmasstrophoblast(significantcontributiontotheinterfacebetweenmotherand
fetus).

Trophoblastictissueisinvasive.Thispropertyisresponsibleforimplantation

Theinnercellmass(embryoproper)dividesintotwolayers(bilaminarembryo),theepiblastaboveand
thehypoblastbelow.

Theepiblastlayerofthebilaminarembryogivesrisetothetrilaminarembryo.The
hypoblast,forthemostpartdisappears.

The primitive streak, a thickening of the epiblast, is the site where epiblast cells migrate toform
thethreelayersofthetrilaminarembryo.

Thetrilaminarembryoiscomprisedofthreebasiclayersoftheembryonicbody:ectoderm,mesoderm
andendoderm.

Thenotochord,whicharisesfromthetipofthestreak,definestheaxisandsymmetryoftheembryo.
Italsoinducestheformationoftheneuraltube.

Theneuraltubeisanectodermalstructure.

Themesodermallayerhasthreesubdivisionsparaxial,intermediateandlateralplate.

Paraxialmesodermformsblocksstretchingfromtheheadendtothetailendoftheembryo.These
blocks,calledsomites,establishthestructuralpatternofthebodywall(dermis,connectivetissueand
muscle).

Lateralplatemesodermsplitsintosomaticandsplanchnicmesoderm

Somaticmesodermformsbodywallconnectivetissuestructures,whilesplanchnicmesodermwillgive
risetothemesenteriesoftheguttube.

Theflattrilaminarembryoundergoescomplexchangesinform,withtheheadandtailends(celphalic
caudalfold)andthesides(transversefold)foldingupmakingitathreedimensionaltubularstructure.
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