MODUL ASISTEN

SELAPUT EKSTRAEMBRIONIK DAN PLASENTA

The embryo not only grows and differentiates, but undergoes those physiological processes that are
necessary for life, some of which are possible only with the aid of the extra-embryonic membranes. These
structures are continuous with the embryonic tissues, but are not part of the embryonic body proper and are
discarded at hatching. They are the following:

• the yolk sac, which encloses the yolk and is so constructed that the partially digested yolk is passed
into the embryonic blood stream;

• the chorioallantois, which has three functions, respiratory exchange, calcium transport and
storage of nitrogenous wastes produced by the embryo;

• the amnion, which is a sac surrounding the embryo and which secretes fluid that cushions the embryo
and prevents it from becoming dehydrated.

To summarize, each of the extraembryonic membranes are composed of the following:

- Chorion: ectoderm and underlying avascular mesoderm
- Amnion: avascular mesoderm and underlying ectoderm
 - Allantois: vascular mesoderm and underlying endoderm
- Yolk sac: vascular mesoderm and underlying endoderm
Yolk sac membrane
The first of the extra-embryonic membranes to develop is the yolk sac membrane. Initially recognized
as the area vitellina, this is the outer area of the blastoderm, consisting of three cell layers, which are
a continuation of the cell layers of the embryonic disc. There is an ectodermal layer adjacent to the
vitelline membranes, an endodermal layer adjacent to the yolk and in between a layer of mesoderm
cells. The mesoderm is split into two by a cavity, the extra-embryonic coelom, and only the mesoderm
next to the endoderm develops blood vessels (called the vascular mesoderm). It is these two cell
layers that form the definitive wall of the yolk sac. The vitelline membranes enclose the yolk sac until
day 4 of incubation. Contact of embryonic tissue with the vitelline membranes alters their structure
(Jensen, 1969), probably facilitating their rupture when yolk sac volume increases at this time. This
causes the yolk to lose its spherical shape and the embryonic-yolk structure adopts the shape of the
egg. The top of this structure is bounded by the yolk sac membrane and the lower by the vitelline
membrane, which slips down to the pole of the yolk sac opposite to the embryo. In this way the
partition of the yolk sac from the albumen (the so called yolk sac umbilicus) is maintained and at this
‘‘vegetal’’ pole, the yolk sac membrane remains incomplete until day 17.
The yolk sac membrane passes the equator of the yolk sac at 5–7 days and achieves its maximum
area around 10–11 days finally surrounding yolk at 14–15 days. From day 12 the yolk sac membrane
changes into a flabby three-lobed mass. After day 10–11 the area of the yolk sac membrane actually
decreases, although the membrane is still increasing in mass and reaches its peak weight at around
15 days (Table 1). This is due to a decrease in the size of the yolk sac as yolk is absorbed.
The primary blood system of the yolk sac membrane, the area vasculosa, is evident by day 2– 3 of
incubation when the vitelline arteries that carry blood from the heart to the periphery of the
blastoderm become clearly visible (Figure 2). At the same time a vein develops at the margin of the
blastoderm, the sinus terminalis. The blood in the peripheral capillary vessels empties into this vein,
which carries blood anteriorly to the anterior vitelline vein, which then conveys blood back to the
heart. A secondary blood system of new veins starts to appear whilst the primary blood system is
still developing. By day 5, the lateral vitelline veins have grown parallel to the vitelline arteries, and
in the peripheral capillaries, in which blood was formerly conveyed in a peripheral direction into the
sinus terminalis, the flow is reversed. Blood now flows from periphery of the blastoderm to heart via
the lateral vitelline veins, a more direct route. By day 6 the sinus terminalis starts to regress (Figure
2).
Amnion
The amnion is formed from a layer of ectoderm and underlying avascular (does not develop blood
vessels) mesoderm immediately adjacent to the embryo. The flat tissue rises up to form folds over
both the head and tail of the developing embryo and by day 4 of incubation the folds fuse together
over the embryo to enclose the amniotic sac. Consequently, the inner layer of the amnion is ectoderm,
and the outer one is avascular mesoderm. Muscle cells appear within this mesoderm so that the
amnion becomes contractile by day 5, but the blood vessels of the amnion arise from the subsequent
fusion of this avascular mesoderm with vascular mesoderm from the allantois. On day 12 a duct
develops between the albumen sac and the amniotic sac; this sero-amniotic connection (Figure 1) is
the point where the head and tail folds of the amnion met on day 4 of incubation. This duct
allows the movement of albumen proteins into amniotic fluid where they are swallowed ending up
in the yolk sac.
Chorion
The chorion develops from tissue continuous with the amnion and away from the embryo. It is
comprised of an outside layer of ectoderm adjacent to the inner shell membrane and an inner
avascular mesoderm, which lines the extra-embryonic coelom. Eventually this mesoderm fuses with
vascular mesoderm ofallantois, so supplying the blood vessels and forming the chorio-allantois. By
day 11 the allantois has completely lined the chorion and the chorio-allantois covers 98% of the area
of the eggshell membranes, thereby acting as the primary respiratory surface for the embryo during
the second half of incubation.
Allantois
At day 2 the allantois is a small bud of endodermal cells and by day 4 it is visible as a free sac growing
out from the primitive hindgut of the embryo. Its inner side is endoderm and the outer surface is
vascular mesoderm. It expands into, and eventually fills, the extraembryonic coelom, so that by day
6 its vascular mesoderm is fused both with the chorion, to form chorioallantois, and with the
avascular mesoderm of the amnion. Hence, the allantois supplies the blood system for both the
chorio-allantois and the amnion. The allantois is a fluid-filled and is a repository for kidney excretions
that first appear by day 5 and enter via the allantoic duct from the cloacal region of the hindgut.
PLACENTA TYPES

Type # 1. Classification Based on the Degree of Intimacy:

Based on the degree of intimacy of foetal and maternal tissues, the following types of placenta
are seen:

i. Non-Deciduous Placenta or Semi Placenta:

In non-deciduous placenta the implantation is superficial. This occurs in most mammals where the
blastocysts lie in the uterine cavity. At the point of contact with the wall of the uterus, the blastocyst
surface gives out finger like projection called chorionic villi that penetrate into the depressions of
the uterine wall and are loosely united.
At the time of birth, when parturition takes place, the chorionic villi are simply withdrawn from
the cavities of the uterine wall without causing any damage or bleeding. This type of placenta
formation is seen in pig, cattle, horse etc., where the less hazardous birth process allows the females
to resume full running speed soon after birth.

As the chorionic villi do not fuse with the endometrium, such a placenta is also called semi
placenta.

ii. Deciduous Placenta or Placenta Vera:

In cat, dog, primates, rodents etc., the degree of intimacy between the chorionic villi and the
endometrium is greatly increased. The uterine wall gets eroded. The chorionic villi fuse with the
eroded uterine mucosa. Such a placenta is termed as placenta vera (true placenta).

At the time when parturition takes place the uterine wall does not remain intact. It tears away and
extensive haemorrhage takes place at birth. Such a type of placenta is termed as deciduous
placenta.

This phenomena of shedding (tearing off) and replacement of maternal tissue is termed as decidua
(meaning, to shed). Here the placenta is physiologically more efficient, where the mothers are
protected enough to recover fully after child birth.

iii. Contra-Deciduate Placenta:

A somewhat modified type of deciduate placenta is seen in Parameles and Talpa (mole), where
there is loss of both maternal tissue as well as foetal portion of placenta. Such a placenta is called
contra-deciduate placenta.

Type # 2. Classification Based on the Types of Implantation:

Implantation is the process by which the embryo becomes attached to a nutritional substance. The
term in case of placental mammals is referred to the process by which the embryo remains
associated intimately with the uterine wall.

Generally three types of implantation are seen which are as follows:

i. Central or Superficial Implantation:

The chorionic sac of the embryo grows and makes superficial attachment with the uterine mucosa.
This type of implantation is called central or superficial implantation and the embryo remains
within the lumen of the uterus (Fig. 5.53A).

It is seen in all cases of implantation in lower vertebrates. It is also present in Preambles and
Dasyurus among the marsupials, while among eutherians it is seen in pig, cow, rabbit, sheep, dog,
cat etc.
ii. Eccentric Implantation:

In mouse, rat, beaver, squirrel etc., the blastocyst in its early stage comes to lie between the uterine
epithelial folds or pocket (Fig. 5.53B), and this type of implantation is called eccentric
implantation. The epithelial folds at a later stage, encloses the blastocyst almost completely.

iii. Interstitial Implantation:

In interstitial implantation the embryo burrows into the uterine mucosa below the epithelium and
becomes surrounded completely by the endometrial tissue of the uterus (Fig. 5.53C). This type of
implantation is seen in hedgehog, guinea-pig, some bats, chimpanzee, man etc.

Type # 3. Classification Based on the Distribution of Villi:

Examination of placentae from different species reveals striking differences in their shape and the
area of contact between fetal and maternal tissue:

• Diffuse: Almost the entire surface of the allantochorion is involved in formation of the
placenta. Seen in horses and pigs.
• Cotyledonary: Multiple, discrete areas of attachment called cotyledons are formed by
interaction of patches of allantochorion with endometrium. The fetal portions of this type
of placenta are called cotyledons, the maternal contact sites (caruncles), and the cotyledon-
caruncle complex a placentome. This type of placentation is observed in ruminants.
• Zonary: The placenta takes the form of a complete or incomplete band of tissue
surrounding the fetus. Seen in carnivores like dogs and cats, seals, bears, and elephants.
• Discoid: A single placenta is formed and is discoid in shape. Seen in primates and rodents.
Type # 4. Classification Based on the Degree of Involvement of Foetal and Maternal Tissues:

Just prior to formation of the placenta, there are a total of six layers of tissue separating maternal
and fetal blood. There are three layers of fetal extraembryonic membranes in the chorioallantoic
placenta of all mammals, all of which are components of the mature placenta:

1. Endothelium lining allantoic capillaries

2. Connective tissue in the form of chorioallantoic mesoderm
3. Chorionic epithelium, the outermost layer of fetal membranes derived from trophoblast

There are also three layers on the maternal side, but the number of these layers which are retained
- that is, not destroyed in the process of placentation - varies greatly among species. The three
potential maternal layers in a placenta are:

1. Endothelium lining endometrial blood vessels

2. Connective tissue of the endometrium
3. Endometrial epithelial cells
One classification scheme for placentas is based on which maternal layers are retained in the placenta,
which of course is the same as stating which maternal tissue is in contact with chorionic epithelium of the
fetus. Each of the possibilities is observed in some group of mammals.

In humans, fetal chorionic epithelium is bathed in maternal blood because chorionic villi have eroded
through maternal endothelium. In contrast, the chorionic epithelium of horse and pig fetuses remains
separated from maternal blood by 3 layers of tissue. One might thus be tempted to consider that exchange
across the equine placenta is much less efficient that across the human placenta.

i. Epitheliochorial Placenta:

This type of placenta involves the contact of the chorionic epithelium with that of uterine
epithelium and thus the term epitheliochorial. The epitheliochorial placenta involves six tissue
barriers between the foetal and maternal circulation. It is the primitive type from which others have
been derived and is seen in pig, sow, mare, horse, cattle etc.

ii. Syndesmochorial Placenta:

In the ruminant ungulates (cattle, sheep, deer, giraffe etc.), varying amounts of the uterine
epithelium may be absent. As a result the chorion is brought into direct contact with the connective
tissue of the uterus. Only five barriers therefore, lie between the two blood streams. This type of
placenta is termed as syndesmochorial.

iii. Endotheliochorial Placenta:

In endotheliochorial placenta the uterine mucosa is reduced and the chorionic epithelium comes in
contact with the endothelial wall of maternal blood vessels. This type of placenta is characteristic
of carnivores like dog, cat, bear etc. Here, only four barriers remain between the two blood streams.

iv. Haemochorial Placenta:

A reduction of the barriers into three is observed in haemochorial placenta, which is found in
primates, insectivores (mole, shrew) and chiropterans (bat). Here, the endothelial wall of the
maternal blood vessel disappears and the chorionic epithelium is bathed directly in the maternal
blood.

v. Haemoendothelial Placenta:

In this type of placenta the chorionic villi looses their epithelium and mesenchymal layers to such
a degree that the endothelial wall of the foetal blood vessels remain in contact with the maternal
blood. The haemoendothelial placenta is observed in mouse, rat, rabbit, guinea-pig etc. and only a
single barrier separates the two blood streams.
Preparat yang akan diamati
- Preparat seri ayam 72 jam amati amnion, korion, yolk sac.
- Preparat ayam utuh 96 jam amati allantois, amnion, yolk sac.