The three villus types

Quiz

Quiz 11

While the embryo is nourished in the first weeks through simple diffusion, later, due
to its rapid growth, it needs a more powerful gas and nutrient exchange system.
This is made possible by the development of the utero-placental circulation
system in which the circulation systems of the mother and of the embryo get
closer together, thus allowing an exchange of gases and metabolites via diffusion.
It must be always kept in mind, though, that maternal and fetal blood never come
into direct contact with each other.
This system decays after the ninth day in the lacunar stage

Quiz

Quiz 03

5b .

Through the lytic activity of the syncytiotrophoblast (Fig. 18 and 19) the maternal
capillaries are eroded and anastomose with the trophoblast lacunae, forming the
sinusoids. At the end of the pregnancy the lacunae communicate with each other
and form a single, connected system that is delimited by the syncytiotrophoblast
and is termed the intervillous space.

Fig. 18: 9th .10th day - Lacunar stage

Fig. 19: 9th -10th day Primary villus

Legend
Lacunar stage (Fig.
18) and primary
villus (Fig. 19)
Spaces form in the
trophoblast (Fig. 18).
Subsequently, due to
the erosion of the
maternal capillaries,
blood gets into the
vacuoles,
engendering the
maternal sinusoids.
(Fig. 19)

1
2
3
4

Cytotrophoblast
Syncytiotrophoblast
Spaces between
syncytiotrophoblast (Lacunae)
Maternal vessel

Maternal vessel, eroded by the ST,

which form the maternal sinusoids
through communication with the
lacunae
See enlarged version in figure 20

Between the 11th and 13th day cytotrophoblast cells penetrate into the cords of
the syncytiotrophoblast creating the primary trophoblast villi 5b .

Fig. 20: 11th -13th day

11th -13th day

Legend

Primary villus with

the cytotrophoblast,
which penetrates into
the processes of the
syncytiotrophoblast,
forming the primary
trophoblast villi.

1
2

Cytotrophoblast
Syncytiotrophoblast

After the 16th day the extra-embryonic mesoblast also grows into this primary
trophoblast villus, which is now called a secondary villus 5c and expands into
the lacunae that are filled with maternal blood. As was already mentioned, the ST
forms the outermost layer of every villus.

Fig. 21: 16th day

16th day

Legend
Secondary villi with
extra-embryonic
mesoblast in the
center, surrounded by
cytotrophoblast and
syncytiotrophoblast.

1 Extra-embryonic mesoblast
2 Cytotrophoblast
3 Syncytiotrophoblast

At the end of the 3rd week the villus mesoblast differentiates into connective
tissue and blood vessels. They connect up with the embryonic blood vessels. Villi
that contain differentiated blood vessels are called tertiary villi 6 .

Fig. 22: 21rst day

21rst day

Legend

Tertiary villi with

extra-embryonic
mesoblast (EEM) in
the center and
additional embryonic
blood vessels. The
EEM remains in this
stage, still surrounded
by cytotrophoblast.
The outer envelope of
the villus is still
formed by the ST.

1
2
3
4

Extra-embryonic mesoblast
Cytotrophoblast
Syncytiotrophoblast
Fetal capillaries

From this time on gases, nutrients, and waste products that diffuse through the
maternal and fetal blood must pass through a total of four layers:

Capillary endothelium of the villus

Loose connective tissue that surrounds the endothelium

Cytotrophoblast

Syncytiotrophoblast

These four elements together form the placental barrier.

Note! The endothelium that surrounds the maternal blood vessels never
penetrates into the trophoblast lacunae, but comes just to their boundaries.
Numerous "daughter" villi arise out of the tertiary villi. These remain either free and
project into the intervillous space (free villi), or they anchor themselves to the basal
plate (anchoring villi). (Interactive diagram).

After the 4th month the cytotrophoblast in the tertiary villi disappear slowly, the villi
divide further and become very thin, whereby the distance between the intervillous
space with maternal blood and the fetal vessels gets smaller. The villi that arise in
this way are called free villi.

Fig. 23 - After the end of the 4th month

Placenta at term

Legend

Free villi with extraembryonic mesoblast

(EEM) and fetal blood
vessels in the center.

1
2
3
4

Extra-embryonic mesoblast
Remains of cytotrophoblast
Syncytiotrophoblast
Fetal capillaries

he cytotrophoblast layer
The cytotrophoblast of the anchoring
villus expands until a further layer
outside the syncytiotrophoblast arises,
forming the cytotrophoblast layer
(interactive diagram). It slips in between
the syncytiotrophoblast and the uterine
endometrium.

Fig. 24 - Development of the

cytotrophoblast layer

1
2
3
4
5

Anchoring villus
Syncytiotrophoblast
Cytotrophoblast
Cytotrophoblast layer
Uterine endometrium

Over the course of the 4th month the cytotrophoblast cells slowly disappear out of
the villus wall and the chorionic plate. They persist, however, in the cytotrophoblast

Legend

To be recognized are
the anchoring
villus with the
cytotrophoblast
shown in dark green
that infiltrates more
and more the basal
plate
and gets in
between the
syncytiotrophoblast
(light green) and the
compact layer of the
decidua in order to
form the
cytotrophoblast layer.
Note that the
syncytiotrophoblast
completely covers the
interior of the
intervillous spaces.

layer. The cytotrophoblast cells penetrate into the decidua and the myometrium
and also colonize the wall of the spiral arteries close to their openings.

Fig. 25 - Growth of the CT into the

walls of the maternal vessels

Fig. 26 - Growth of the CT into the

walls of the maternal vessels

Legend

Fig. 25, Fig. 26

Progressive growth of
the cytotrophoblast
cells into the decidua
and into the wall of
the spiral arteries.

1
2
3
4

Syncytiotrophoblast (ST)
Cytotrophoblast (CT)
Endothelial cells
Smooth muscle cells

5
6

Spiral arteries
Endovascular cytotrophoblast

This invasion of the maternal vessels by the cytotrophoblast leads to the

destruction of the smooth muscle layer and to a partial replacement of the
endothelial cells. It is responsible for the change in elasticity of the spiral arteries,
whereby the blood circulation of this fetoplacental unit is adapted to the rapid
growth of the fetus. This phenomenon of cell exchange is absent
in preeclampsia or an intra-uterine growth retardation.An excessive proliferation of
the cytotrophoblast can lead to tumor formation, especially to a chorion carcinoma.

More info

Supplementary
information
regarding the
physiopathology of
the mechanisms of
preeclampsia
Physiopathology of
preeclampsia
(French)
HTA of the
pregnancy (French)

More info

During a normal pregancy the maternal spiral arteries that nourish the placenta
are continuosly pulled into the lacunar system.
These structural adaptations are accompanied by an edema, the dissolution of
the endothelium and destruction of the tunica media and the membrana elastica
interna, which are replaced by fibrous tissue. Through these alterations the

arteries are removed from neuro-vascular control and the influence of the toneproducing vessel mediators (prostaglandin, nitrous oxide, endothelin). Thus, a
larger blood flow is allowed in the placenta.
The migration of the trophoblast cells stands under a strict temporal-spatial
control, an alteration of which can provoke a disorder in placental function. This
ranges from a preeclampsia (characterized by insufficient penetration by the
trophoblast) to a chorion carcinoma (characterized by an excessive trophoblast
invasion).

Placental tissue structure

Maternal and fetal tissues form two units that are closely bound together at the
placental level.

Quiz

Quiz 12

The fetal part of the placenta is made up of the chorionic plate with its
placental villi, the cytotrophoblast layer and the intervillous spaces. The
chorionic plate (great part of the placenta on the fetal side) consists of the
amnion, the extra-embryonic mesenchyma, the cytotrophoblast and the
syncytiotrophoblast.

The basal plate, the peripheral region of the placenta on the maternal side that
is in contact with the uterine wall, is made up of two tissues: embryonic tissue
(cytotrophoblast, syncytiotrophoblast), on the one side, and of maternal tissue
(decidua basalis) on the other.

The maternal side of the placenta is made up of the decidua basalis, uterine
vessels and glands.

Fig. 27 - Chorionic plate

Fig. 28 - Basal plate

Legend

1
2
3
4

Amnion
Extra-embryonic mesoblast
Cytotrophoblast
Syncytiotrophoblast

5
6
7
8

Zona compacta
Zona spongiosa
Decidua basalis
Myometrium

Development of the placenta

After the 4th month the cytotrophoblast slowly disappears from the walls of the
tertiary villi (interactive diagram), whereby the distance between the maternal and
fetal vessels diminishes. They also disappear from the chorionic plate. In the basal
plate the cytotrophoblast remains mainly at the level of the cytotrophoblast layer.
Together with the decidua and fibrin deposits, they form protrusions (intercotyledon septa) that project into the intervillous space, dividing it to some extent
into so-called cotyledons.The formative mechanism of these inter-cotyledon septa
remains speculative and probably depends on the folding together of the basal
plate which, for its part, has resulted from the proliferation of the stem villi. They
push the basal plate back. In addition, the spread of the placenta into the uterine
cavity also appears to contribute to the creation of the septa (15) .

Fig. 29 - Development of the placenta (> 4th month)

Quiz

Quiz 13

Legend

Fig. 29
The cytotrophoblast
islands move into the
periphery of the
cotyledons and,
together with the
decidual tissue, are
involved with
formation of the
placental intercotyledon septa.

1
2
3
4

Decidual tissue
Syncytiotrophoblast
Cytotrophoblast islands
Septum

These septa delimit the cotyledons but never merge with the chorionic plate.
Maternal blood can accordingly circulate freely from one cotyledon to the other
(interactive diagram).
The villus stems of the placenta lengthen considerably towards the end of the
pregnancy and the fibrinoid deposits (extra-cellular substance made up of fibrin,
placental secretions and dead trophoblast cells), accumulate in the placenta. This
happens especially under the chorionic plate, where they form the
subchorialLanghans' layer, as well as at the level of the basal plate beneath the
stem villi and the cytotrophoblast layer, where the fibrin deposits form Rohr's
layer. Still deeper in the decidua basalis these deposits form Nitabuch's layer.
This is where the placenta detaches itself from the uterus at birth (interactive
diagram).

Fig. 30 - Definitive placenta

Legend

Fig. 30
The fibrinoid deposits
form the subchorionic Langhans'
layer A. Rohr's
layer B is found at
the level of the basal
plate beneath the
stem villi. Lying still
deeper in the decidua
basalis they form
Nitabuch's layerC.
This is located at the
boundary between
the zona spongiosa
and the zona
compacta (where the
release of the
placenta takes place).

A
B
C

Subchorial Langhans' layer

Rohr's layer
Nitabuch's layer

More info

The fibrinoid deposits are structurally and chemically closely related to fibrin
that is formed by the activation of fibrinogen in the blood vessels. Recent
research results emphasize the existence of two types of fibrinoid (16).
The fibrinoid deposits are present in all normal placentas, increase in thickness
during the pregnancy, and can take up a maximum of 30% of the placental
volume without affecting its function. Normally, at the end of the pregnancy, they
do not occupy more than 10 to 20% of the placenta volume.
It seems their generation is connected with micro-lesions of the
syncytiotrophoblast. Through these injuries, the mesenchyma comes into contact
with maternal blood. This provokes the activation of local coagulation
mechanisms. The mechanism resembles those which are brought about through
endothelial lesions in the blood vessels of adults.
Fibrinoid deposits also envelop all necrotic material that ensues from placental
degeneration. When these deposits are massive and block one or more vessels
to the villi, they form white infarcts, which can be seen macroscopically on the
placenta.
The functional importance of the fibrinoid seems to be quite complex. Besides
their sealing effects, they also play a role in the immunologic "barrier" between
feto-maternal tissue as well in the anchoring of the placenta.

he decidua

At the implantation location, the maternal endometrium is changed by the decidual

reaction (epithelial transformation of the fibroblasts of the uterine stroma, in that
lipids and glycogen accumulate) and is called the decidua.

Quiz

Quiz 05

The decidua consists of various parts, depending on its relationship with the
embryo:

Decidua basalis, where the implantation takes place and the basal plate is
formed. This can be subdivided into a zona compacta and a zona
spongiosa (where the detachment of the placenta takes place following
birth).

Decidua capsularis, lies like a capsule around the chorion

Decidua parietalis, on the opposite uterus wall

At around the 4th month, the fetus is so large that the decidua capsularis comes
into contact with the decidua parietalis. The merging of these two deciduae causes
the uterine cavity to obliterate.

Fig. 31 - 8th week

Fig. 32 - 12th week

Legend

ig. 31, Fig. 32

The three deciduae in the
second month of pregnancy.
After the 4th month the
growing fetus brings the
decidua capsularis into
contact with the decidua
parietalis. The fusion of the
two deciduae leads to the
disappearance of the
uterine cavity.
1
2
3
4

Decidua parietalis
Decidua capsularis
Decidua basalis
Uterine cavity

Development of the villi

5
6
7
8

Smooth chorion (laeve)

Chorionic villi
Amniotic cavity
Decidua capsularis and parietalis,
grown together

In the early stage of development the trophoblastic villi form all around the embryo
and give it the appearance of a "hairy ball".
During the 3rd month most placental villi disappear but remain present at the basal
plate:

The chorion gets here to be a villus-rich chorion (chorion frondosum), a

major component of the placenta.

At other locations, where the villi degenerate, the chorion becomes

smooth (chorion laeve). At these locations, no exchange between the
maternal and fetal blood circulation systems takes place.

Quiz

Quiz 23

The chorion laeve is formed from a chorion layer (consisting of extra-embryonic

mesenchyma and cytotrophoblast).
In this stage all placental villi are tertiary villi. After the 9th week the tertiary villi
lengthen through the proliferation of the terminal villus mesenchyma (see more
info).

Definitive placental villi > 4th month

Legend
Scanning electromicroscopic image of
the placental villus
tree.

More info

After the end of the 4th week all of the placental villi are tertiary villi.
Histologically and morphologically they develop themselves further in the

direction of a new type of villus that is better adapted to the growing embryonic
needs. The tertiary villi, with a diameter between 60 -200 mm, are homogenous
and rich in mesenchyma with few capillaries. They grow first in length and later,
through proliferation of the trophoblast and extra-embryonic mesoblast (EEM) as
well as of endothelial cells, they also increase in density and width. The
trophoblast covering forms numerous trophoblast buds, which at the beginning
are formed by the syncytiotrophoblast (similar to the primary villi), but
nevertheless cytotrophoblast and the EEM grow very rapidly into these villi.
After the ninth and up to the 16th week the tertiary villi mesh and change
through proliferation of the terminal villus mesenchyma, the trophoblast and the
fetal endothelial cells in order to finally form intermediate immature villi. The latter
generate trophoblast buds, out of which new, thinner villi arise. These villi are rich
in large diameter vessels, guaranteeing increased perfusion.
After the 26th week, the (mesenchymatous) tertiary villi develop longer and
thinner villi (60 to 200 mm in diameter) in which the fetal capillary network gets
richer. These are termed mature intermediate villi that, for their part, form small
meshes, terminal or free villi (diameter 40 - 80 mm), as well as new trophoblast
buds.
At the end of the pregnancy six types of villi are to be found in the placenta:
1.

Stem villi

2.

Tertiary mesenchymatous villi

3.

Immature intermediate villi

4.

Mature intermediate villi

5.

Terminal or free villi

6.

Trophoblast buds

The stem villi guarantee the mechanical stability of the villus tree, the
immature intermediate villi are the place of the proliferation of the trophoblast
and the trophoblast buds. Tertiary villi (mesenchymatous) are responsible for the
lengthening of the villus tree.
From the mature intermediate villi the free villi or terminal villi arise which
makes up 40% of the volume and 50% of the exchange surface of the placenta
before birth. (17)