BS Zoology: 5th Semester

ZOO-504 DEVELOPMENTAL BIOLOGY Cr. Hours: 4(3+1)

Unit: 08 Placentation in Mammals

Hafiz Mehmood Ul Hassan
Lecturer in Zoology, GPGC Nowshera
M.Phil Zoology/ Fisheries & Aquaculture (UoP)
M.Sc Zoology/Fisheries (UoP)
Genotoxicology (Humans)
M.Ed Science Education (AIOU)
mmzoologist@gmail.com
0302-5772747
 Placentation .

Placentation refers to the formation, type and structure, or arrangement of the placenta.
During pregnancy, placentation is the formation and growth of the placenta inside the uterus.
It occurs after the implantation of the embryo into the uterine wall and involves the
remodeling of blood vessels in order to supply the needed amount of blood. In humans,
placentation takes place 7–8 days after fertilization.
The function of placentation is to transfer nutrients, respiratory gases, and water from
maternal tissue to a growing embryo, and in some instances to remove waste from the
embryo. Placentation is best known in live-bearing mammals (theria), but also occurs in
some fish, reptiles, amphibians, a diversity of invertebrates, and flowering plants.
In vertebrates, placentas have evolved more than 100 times independently, with the majority
of these instances occurring in squamate reptiles.
Note: In flowering plants, placentation is the attachment of ovules inside the ovary. The
ovules inside a flower's ovary (which later become the seeds inside a fruit) are attached
via funiculi, the plant part equivalent to an umbilical cord. The part of the ovary where the
funiculus attaches is referred to as the placenta.
 Definition of Placenta
• Placenta: The placenta can be defined as an organ formed by the sustained apposition or
fusion of fetal membranes and parental tissue for physiological exchange.
• The term placenta was delved from Greek word it means flat cake. Placenta is a
special connective tissue, which contains the uterus of mother and foetal membranes of
foetus.
• A placenta is an organ built up of maternal and foetal tissues jointly. It serves for the
transport of nutrients from the mother tissues with those of the embryo as well as the
exchange of gases between the tissues of the two. Thus a placenta may be defined as a
temporary connection between the maternal and foetal tissues for the purpose of shelter,
nutrition, respiration, excretion and defense.

• a fetomaternal organ
• begins developing at implantation of the blastocyst
• shortly after birth the placenta is expelled from the uterus – as the Afterbirth
 Placenta
The placenta is composed of three layers. The innermost placental layer surrounding the
fetus is called the amnion. The allantois is the middle layer of the placenta (derived from
the embryonic hindgut); blood vessels originating from the umbilicus traverse
this membrane.
 Extra Embryonic Membranes / Placental
membranes
• In vertebrate embryonic development, only a part of the egg or the cleavage mass of cells
forms the actual embryo, while other parts lying outside the embryonic territory develop
into extra-embryonic regions, called embryonic or foetal membranes. Embryonic
membranes are auxiliary organs, which have arisen partly for protection of the embryo,
and more specially to provide for its nutrition, respiration and excretion until the
independent existence is attained.

• Definition: The thin protective membranes formed on the periphery of the embryo in
amniotes (reptiles, birds and mammals) are known as extraembryonic membranes.
• a structure (such as the amnion) that derives from the fertilized ovum but does not form a
part of the embryo
There are four main types namely amnion, yolk sac, allantois, and chorion
 Extra Embryonic Membranes: Amnion
Amnion: thin ectodermally-derived membrane
The amnion is the innermost layer, enclosing the embryo in the amnion fluid, which protects it
from mechanical stress.
Characteristics:
• Non-vascular, fluid filled, Fluid produced by fetus
• Protective cushion
• Fluid allows embryo to move, change position, and shape
• Ruptures at birth (not breaking of water)
Functions:
The primary function of amnion is to protect the embryo from dessication and provides
“private salty” to an embryo to float. Amniotic fluid serves as an efficient shock
absorber.
 Extra Embryonic Membranes: Yolk Sac
Yolk Sac: endodermal,
Tissue layer growing over the yolk – digests and absorb yolk material
• In birds to nourish embryo
• In mammal atrophies but source of blood cells and primordial germ cells
Functions:
• In vertebrate embryonic development, only a part of the egg or the cleavage mass of cells
forms the actual embryo, while other parts lying outside the embryonic territory develop into
extra-embryonic regions, called embryonic or foetal membranes. Embryonic membranes
are auxiliary organs, which have arisen partly for protection of the embryo, and more
specially to provide for its nutrition, respiration and excretion until the independent existence
is attained.
 Extra Embryonic Membranes: Allontois
Allantois: endodermal, originate from ventral surface of early hindgut.
Act as reservoir for storing or removing urinary wastes and mediate gas exchange
between embryo and surrounding.
Characteristics:
• Blood vessels
• Fuses with chorion
• Allantochorion or chorioallantois
• Brings blood vessels to chorion
Functions:
In birds, cavity of allantois serves as an urinary bladder. In mammals, the original
function of allantois as urinary bladder becomes all together lost. The carbon dioxide
produced by the embryo diffuses into the maternal blood and is excreted by the kidney of
the mother.
 Extra Embryonic Membranes: Chorion
Chorion: outermost membrane; Attachment to mother
The chorion and the amnion together form the amniotic sac. It is formed by
extraembryonic mesoderm and the two layers of trophoblast that surround the embryo
and other membranes. The chorionic villi emerge from the chorion, invade the
endometrium, and allow the transfer of nutrients from maternal blood to fetal blood.
The function of the chorion is to contribute to the development of the placenta in
placental mammals.
Functions:
A part of chorion forms finger like out growths known as chorionic villi that
penetrate into the wall of uterus for exchange of substances between embryo and uterus in
the body of the mother.
Amniote embryos develop in a fluid-
filled sac within a shell or uterus
• The amniote embryo is the solution to reproduction
in a dry environment.
reptile & bird mammal

chorion
amnion
embryo

allantois
yolk sac

Fetal portion of placenta

Maternal portion of placenta

Extraembryonic membrane
 Amniote embryos develop in a fluid-
filled sac within a shell or uterus
• The amniote embryo is the solution to reproduction
in a dry environment.

chorion
amnion
embryo

allantois
yolk sac

Fetal portion of placenta

Extraembryonic membrane Maternal portion of placenta

 Development of Placenta
• When the mammalian embryo enters the uterus, it remains bathed in the uterine fluid
containing organic substances produced by the tubular glands of the uterine wall. The
early embryo may absorb some of these substances through the epithelial covering till
the placenta formation. For its further development, the embryo completely depends
on substances supplied to it from the maternal tissue.
• A mammalian placenta is a composite structure based on double origin. It is produced
by the development and apposition of extra-embryonic membranes (chorion, allantois
or yolk sac) with the endometrium of the uterine wall (decidua basalis) which
includes both the compact and spongy layers for the purpose of physiological
exchange. In between these two parallel plates (the chorionic membrane and the
endometrium) a huge blood sinus, the intervillous space, contains an enormous
number of chorionic villi.
• In mammalian placenta there are two possible sources of chorionic vascularization-
the vitelline circulation provided by the allantois.
•
 Placenta
Uterine stromal cells around blastocyst - enlarge, form
glycogen and lipid droplets -- decidual reaction, decidual
cells
The endometrium (lining of the uterus) of the mother
consisting of three regions named by location.

Region Description
Decidua basalis Region between the blastocyst
and the myometrium
Decidua capsularis Endometrium that covers the
implanted blastocyst
Decidua parietalis All the remaining endometrium
 Development of Placenta: Humans
• In humans, the placenta develops in the following manner.
• Chorionic villi (from the embryo) on the embryonic pole grow, forming chorion
frondosum. Villi on the opposite side (abembryonic pole) degenerate and form the
chorion laeve (or chorionic laevae), a smooth surface. The endometrium (from the
mother) over the chorion frondosum (this part of the endometrium is called the decidua
basalis) forms the decidual plate. The decidual plate is tightly attached to the chorion
frondosum and goes on to form the actual placenta. Endometrium on the opposite side
to the decidua basalis is the decidua parietalis. This fuses with the chorion laevae, thus
filling up the uterine cavity.
• In the case of twins, dichorionic placentation refers to the presence of two placentas
(in all dizygotic and some monozygotic twins). Monochorionic placentation occurs
when monozygotic twins develop with only one placenta and bears a higher risk of
complications during pregnancy. Abnormal placentation can lead to an early termination
of pregnancy, for example in pre-eclampsia.
Umbilical cord
• A composite structure formed by contributions from:
• Fetal connecting (body) stalk
• Yolk sac
• Amnion
• contains the right and left umbilical arteries, the left umbilical vein,
and mucous connective tissue. Presence of only one umbilical artery
may suggest the presence of cardiovascular anomalies.
 Types of placenta: in prototherian mammals
• Prototherian mammals are egg laying mammals. Hence placenta is not formed in uterus.
In marsupials the embryo develops incompletely in the uterus. They show two types of
placenta.
• Yolk sac placenta and
• Primitive allantoic placenta
 Chorio-vitelline / yolk-sac placenta
• Some marsupials (Didelphis, Macropus), allantois remains relatively small and
never makes contact with the chorion, where as the yolk-sac with its network of
vitelline blood vessels becomes very large and fuses broadly with the chorion to
complete a transitory yolk-sac placenta or chorio-vitelline placenta.
• Further, the chorion never advances beyond a smooth membrane in close
apposition with the vascular uterine endometrium. The uterine wall secretes a
viscous fluid, the uterine milk. It is absorbed by the villi of yolk-sac placenta and
through vitelline circulation is carried to the embryo.
OR
• In these animals the developing embryo shows small allantois. It will never come
in contact with chorion. Their yolk sac was large. It comes in contact with chorin.
This part will gain blood vessels. This part will come in contact with endometrium
of uterus. This is only a contact, but not fusion. Through this contact the embryo
will absorb nourishment from mother. This is called chorio vitelline placenta or
Yolk sac placenta.
 Chorio-allantoic placenta
• In some other marsupials (Parameles, Dasyuris) and all the eutherian mammals, the yolk
sac remains rudimentary but allantoic blood vessels are well developed. The allantois fuses
with the chorion to form a composite membrane. This membrane serves as a component
part of the so-called chorio-allantoic placenta.
• The chorion grows out into root like vascular processes, the chorionic villi that engage the
uterine mucosa in a more or less intimate functional relation persisting throughout
pregnancy. Remnants of the chorio-vitelline placenta may be found either temporarily or
even permanently in higher mammals playing a subsidiary role in their Placentation. When
the allantois is insignificant or lacking, the placenta is known as chorionic placenta.
• OR
• In paramoles simple allantoic placenta is developed. Allantoic will enlarge. It comes In
contact with chorion This structure will be closely applied to mother’s uterus. It is called
chorio-allantoic placenta. In these animals yolk sac placenta is not seen.
 Allonto-chorionic villi
The chorionic villi are finger-like outgrowths that penetrate into depressions in the wall of the uterus
through which all kinds of interchanges take place between the mother and the foetus. The early villi
are compact, bush-like tufts with a few small branches. Their main stems arise from the chorionic
membranes and almost all the eroded surfaces of compact endometrium (decidua basalis).
Side branches begin to develop in the early period of pregnancy and produce many villi as well.
During the middle and late months of pregnancy, the villi become much more branched, tree like
with permanent trunks and long and slender branches bearing innumerable small twigs.
Many terminal branches fuse with the decidua basalis of the endometrium at some distance from
their ends, and then recurve into the inter-villous space in a shaped manner. All parts of the villous
tree have the same structural plan. At the center is a connective tissue core, in which are embedded
the allantoic blood vessels (in case of chorioallantoic placenta) or the vitelline blood vessels (in case
of chorio-vitelline placenta).
These appear as arterioles and venules but taper to prominent capillaries, which continue to the
villous tips where they complete a continuous system of closed vessels. The connective tissue core is
covered with a double layer of trophoblast.
Inside, next to the connective tissue, is a single layered cyto-trophoblast or cellular trophoblast with
its separate cuboidal cells, it is also known as the layer of Langhans. The cellular layer gives rise to
syncytium, the syntrophoblast, which covers the villi externally.
 Placenta In Eutheria
In Eutherian mammals the embryo develops in the uterus of mother. The developing
embryo will get nourishment from mother through the placenta. Placenta is not
common to all mammals. It is developed well in Eutheria.
In Eutherian mammals true allantoic placenta is seen. Allantoic becomes big and
comes in contact with chorion. This part will show close association with uterine wall.
This connection is called placental connection. The structure of placenta will vary in
different orders of Eutheria.
Placenta is classified in three ways.
• The placenta classification on nature of contact.
• Placenta is classified basis on the distribution of villi.
• Classification of placenta basing on histology.
 NATURE OF CONTACT
It is two types, Indeciduate and deciduate type.

Indeciduate type placenta: Ex: Ungulate, Cetaceans, Sirenians. Lemurs:

The chorionic villi are simple projections, they lie in contact with uterus. They have
a loose contact. There is no fusion. At the time of birth of embryo uterus is not
damaged.

Deciduate type Placenta: Ex: Primates, Rodentia, Insectivora, chiroptera.

The allanto-chorionic villi penetrate into uterine villi. They are intimately fused.
Hence at the time of birth, the uterus is damaged. Bleeding occurs, the uterine wall
enters into formation of placenta is called deciduas.
 Basing on distribution of villi / Placental Shape/
Contact Point
According to the distribution of villi five kinds of placenta are seen.
Diffused type placenta: Ex: Horse, pig. The villi are uniformly
(Almost the entire surface of the allantochorion is involved in
formation of the placenta) distributed on the surface of blastocyst,
except at the extreme ends.
Cotyledonary placenta: Ex: Sheep, Cow, Deer (ruminants) Cotyledonary

The villi are arranged in groups. Each group is called cotyledon.

Each cotyledon fits into caruncle of uterus.
Multiple, discrete areas of Diffuse Placenta Diffuse Placenta: horse
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 are called
caruncles, and the cotyledon-
caruncle complex is called a
placentome
 Basing on distribution of villi / Placental Shape/
Contact Point
Intermediate type Placenta: Ex: Camel, Giraffe. It is
a rare type, it shows free villi on cotyledons. Hence it Zonary Placenta
is called intermediate type placenta.
In these three types of placenta during parturition the
foetus will not damage uterus.

• Zonary placenta: the placenta takes the form of a

complete or incomplete band of tissue surrounding
the fetus. The villi are in the form of transverse
zones.
• In dog a single girdle of villi will be present. In fox
two girdles of villi are present. The villi penetrate
Into uterine wall. Hence during parturition uterine
wall is damaged.
Ex: in carnivores like dogs and cats, seals, bears,
and elephants.
 Basing on distribution of villi / Placental Shape/
Contact Point
Discoidal Placenta: Ex: Rat, Bat, Rabbit.
• A single placenta is formed and is discoid in shape.
• On the entire surface of blastocyst the villi are in the form of
discs. When the embryo is growing, it moves away from
uterus hence the with look like a disc. These villi are
Intimately connected with uterus. Hence during parturition
much uterine tissue is damaged.
• At birth the placenta and decidua expelled as the afterbirth
Types of Placenta Basing on distribution of villi

The image to the right shows an incised uterus from a pregnant sheep,
roughly 50 days of gestation. The numerous button-shaped structures
are placentomes, and the surfaces in view are actually cotyledons - the
fetal side of the placentome. The slightly milky-looking membrane
covering and between placentomes is the chorioallantois. The fetus is
clearly visible inside the amnion
 Classification of placenta (basing on histology) / on the basis
of tissues
Mammalian placentas can be classified based on the number of tissues separating the maternal from
the fetal blood.
According to number of layers of cells present between foetus and uterus blood supply the placenta Is
classified into five types.
a) Epithelio chorial placenta: Ex: Pig, Horse, (Ungulates Lemmures),ruminants, whales, ower primates.
The foetal chorion is in contact with epithelium of the uterus hence it is called epithellio- chorial
placenta. Chorionic villi, growing into the apertures of uterine glands (epithelium).
In between foetal, maternal parts six layers are present.
1. Endothelium of mother blood vessel.
2. Maternal syndesmose connective tissue.
3. Epitheliurn of mother
4. Chorion of foetus.
5. Foetus connective tissue (syndesmose
6. Endothellum of foetal blood vessel.
If all the six layers are present the placenta is called epithello chorial placenta.
 Classification of placenta (basing on histology)
/ on the basis of tissues
b) Syndesmose chorial or Syndesmo chorial Placenta: Ex: Sheep, Cow.
A type of placentation characterized by an endometrial attachment to the chorion with a limited
amount of destruction of the endometrial epithelium. Formerly thought to be characteristic of the
ewe and goat doe, these species are now known to have epitheliochorial placentae.
The allanto-chorionic villi will pierce into the uterus of the mother, the chorion will come in contact
with syndesmose of mother’s uterus. Hence it is called syndesmose chorial.
Syndesmose : A membranous vascular organ that develops in female eutherian mammals during
pregnancy, lining the uterine wall and partially enveloping the fetus, to which it is attached by
the umbilical cord.

c) Endothelio chorial placenta: Ex: Cats, Dogs, i.e., in most carnivores.

In this type of placentation, the chorionic villi (or chorion) are in contact with the endothelium
of maternal blood vessels (mother’s uterus).
 Classification of placenta (basing on histology) / on the
basis of tissues
d) Hemochorial placenta: (Haem-blood) Ex: Bat, Man, Primates, Insectivores.
“having the fetal epithelium bathed in maternal blood”. The placental connections are more
intimate/deep. The chorion of foetus will float In the blood pools of mother’s uterus. Hence It Is called
haemochorial placenta.
In hemochorial placentation maternal blood comes in direct contact with the fetal chorion, which it
does not in the other two types. It may avail for more efficient transfer of nutrients etc., but is also
more challenging for the systems of gestational immune tolerance to avoid rejection of the fetus. (e.g.
in higher order primates, including humans, and also in rabbits, guinea pigs, mice, and rats).

e) Hemo endothelial placenta: Ex: Rat, Rabbit,

A placenta in which the trophoblast becomes so attenuated that maternal blood is separated from fetal
blood only by the endothelium of the chorionic capillaries.
Hence guinea-pig will float In mother’s blood. Hence it called hemo endothelial placenta.
Classification of placenta (basing on
histology) / on the basis of tissues
 Functions of placenta
• Placenta will form a physiological barrier between mother and foetus. It will possess foetal and
maternal blood mixing.
• Placenta allows the diffusion of monosacharides, amino adds, hormones, vitamins, oxygen,
.carbondioxide, water and other waste materials, because of this it supplies food, oxygen to foetus.
• It works as an excretory organ of foetus. It releases the nitrogenous waste materials Into mother
blood.
• It works as an endocrine gland. It will secretes lactogen ,progesterone,etc. hormones.
• The placenta will manufacture fructose from glucose.