MEMBRANOGENESIS AND PLACENTAL FUNCTION

LEARNING OUTCOMES

1. describe the formation of the tubular embryo by creation of body folds

2. note the juxtaposition of ectoderm and endoderm at the oral plate and cloacal membrane

3. describe the formation and fusion of the amnion to create a protective bubble around the embryo

4. be aware of the vestigial nature of the yolk sac in mammals with regard to nutrition but its importance
in terms of haematopoiesis and its transient significance in contributing to the choriovitelline placenta

5. show the development of the allantois as a bud of the gut tube and its importance in the
chorio-allantoic placenta

6. understand the different histological forms that the foetal/maternal placenta interface can take

7. understand the different anatomical forms that the foetal/maternal placenta interface can take

8. Note the emphasis on glucose and amino acids as energy and growth resources in the foetus and
explain how the placenta has an endocrine function in ensuring these resources are directed to the foetus

9. Explain the special foetal adaptations that ensure adequate perfusion of foetal tissues with oxygen
The flat embryo begins to fold downwards at the sides and at the
front and back to enclose a primitive gut

Ectoderm
Neural tube

Mesoderm

Endoderm

Ectoderm
Neural tube Hindgut
Mesoderm
Endoderm
Cloacal plate
Oral plate

FUSION POINTS
OF ECTODERM
AND ENDODERM
Regions of brain FUSION POINTS
WITH EXTRA-
EMBRYONIC
MEMBRANES Longitudinal view
The folding process not only creates the body form but also the extra-
embryonic structures of the yolk sac, allantois, amnion and chorion
Embryo proper
Primitive gut

Presumptive amnion Allantois

Presumptive chorion

Transient chorio-vitelline placenta

Yolk sac Splanchnopleure

Somatopleure Longitudinal view
 The chorio-vitelline placenta is temporary (or absent)
and is replaced by the chorio-allantoic placenta

Chorio-allantoic placenta
Amnion
Chorion

Allantois
Yolk sac

Longitudinal view
 Folds also undercut the sides of the body

Somatopleure Gut tube

Neural tube
Mesoderm
Ectoderm
Endoderm

Splanchnopleure Yolk sac

Transverse view
 Video of chick embryo (50h)
showing body folds and amnion
In the chick, the formation of the head fold
precedes that of the tail and the formation
of the body sides progresses caudally

https://www.eevec.vet.ed.ac.uk/vc/node.asp?ID=vcembr02
 Histological classification of placentas is based on the degree
of removal of the maternal layers
KEY CHARACTERISTICS OF MAMMALIAN PLACENTA EPITHELIOCHORIAL
Maternal endometrial epithelium intact
(horse,pig)
Foetal capillary (from umbilical artery)

SYNEPITHELIOCHORIAL
Endothelial layer Syncytium of maternal epithelium and
Chorion
(ruminants)
Connective tissue layer
(may be minimal) ENDOTHELIOCHORIAL
Removal of endometrial epithelium
And connective tissue
Cellular layer
(may be trophoectoderm + maternal (dogs, cats)
epithelium or a syncytium of the two,
or solely trophoectoderm) HAEMOCHORIAL
Connective tissue layer Removal of maternal endothelium
(may be minimal) (human, some rodents)

Maternal capillary
(in haemochorial placenta of primates
the endothelium is degraded)
Gross anatomical classification of placentas is based on the pattern
of contact between chorion and endometrium

DIFFUSE COTYLEDONARY
Uniform distribution of chorionic villi Villi restricted to defined area
over contact surface (horse, pigs) (cotyledons) (ruminants)
ZONARY DISCOIDAL
Girdle of chorionic villi around middle Disc-shaped area on chorionic sac
of chorionic sac (dogs,cats) (humans, rodents)
 The haemochorial placenta shows the intimate juxtaposition
of foetal and maternal blood allowing efficient exchange
Foetal capillaries
The haemochorial placenta
A B

Umbilical vein

Umbilical arteries Notice the expansions

at theturnaround to
Maternal
allow slower blood flow
Maternal venule
Maternal and better equilibration
blood pool with maternal blood
Chorionic arteriole
villi

A http://instruct1.cit.cornell.edu/courses/biog105/pages/demos/105/unit8/ovaryplacenta.html
B from Johnson, Essential Reproduction
 Glucose is the dominant energy yielding substrate
for the foetus with little use of fatty acids

ENERGY SUBSTRATES
Glucose oxidation accounts from 50% oxygen use

Most of the rest is due to amino acid and lactate oxidation

Rather little is from fatty acid oxidation

(Notice that maternal energy metabolism is almost the mirror of this

with a shift to fatty acid oxidation and a shift away from amino acids
and glucose)
 To some extent the foetal-placental unit programmes maternal
metabolism to ensure that it meets the needs of the foetus

FOETAL INFLUENCES ON MATERNAL METABOLISM 1

Maternal tissues

CSM
GLUCOSE
GLUCOSE
Maternal liver

MOTHER FOETUS

CSM = CHORIONIC SOMATOMAMMOTROPHIN (also know as placental lactogen)

CSM secreted in increasing amounts during gestation

CSM suppresses insulin action

Therefore depresses glucose use by the mother

'Directs' glucose to the foetus

Maternal insulin resistance can precipitate maternal type 2 Diabetes mellitus

 Although fatty acids are little used by the foetus for energy they are
essential for growth and also for laying down fat reserves
LIPID METABOLISM IN THE FOETUS

TAG 3
Maternal adipose SYNTHESIS

FATTY 1 2 OXIDATION
ACIDS
LIPOPROTEINS FATTY
ACIDS
LPL 4
Maternal liver CELL
MEMBRANES

MOTHER FOETUS

1. Fatty acids transported via maternal (or foetal) serum albumin

2. (a) Triacylglcyerols contain mostly palmitate

(b) Palmitate will also be formed from excess glucose
(c) Epitheliochorial placentas have poor rates of diffusion of fatty acids and neonates (eg calf and piglet)
have little body fat compared to the haemochorial model (human)

3. TAG deposits in both white and brown adipose tissue. Brown fat essential for thermogenesis in neonate

4. Crucial here are the essential fatty acids

18:3 (D9,12,15)
18:2 (D9,12)
20:4 (D5,8,11,14)
As with glucose, the foetal-placental unit programmes mammalian
metabolism to ensure that it meets the Nitrogen needs of the foetus

FOETAL INFLUENCES ON MATERNAL METABOLISM 2

Maternal tissues Maternal liver 1

UREA
OXIDATION
GROWTH
PROGESTERONE
AMINO
ACIDS
AMINO
ACIDS

MOTHER FOETUS
Notes:
1. An added benefit of the redirection of amino acids from
the maternal liver is that maternal urea production is low
thus favouring urea return across the placenta
Several foetal adaptations contribute to the ability of
the foetus to deliver sufficient oxygen to its tissues

OXYGEN SUPPLY - FOETAL ADAPTATIONS 1

Cardiac anatomy limits intermixing of oxygenated blood

and venous return from the head

Foetal haemoglobin has a high affinity for oxygen

There is a double Bohr effect acting on the placental

transfer of oxygen

Cardiac output is high

Haemoglobin concentration is 50% higher than maternal

The foetal cardiovascular system is adapted to providing well-oxygenated blood to the
brain in spite of intermixing of venous return and an incompletely divided heart

OXYGEN SUPPLY - FOETAL ADAPTATIONS 2

1. Numbers are partial pressures of oxygen in

mm Hg

2. Low vascular resistance in placenta takes

45% of cardiac output

3. Blood returning to right atrium is a mixture

of oxygenated umbilical blood and
venous return from trunk and limbs

4. Crista dividens directs this better

oxygenated blood through foramen
ovale for preferential delivery to brain via
left ventricle

5. The poorly oxygenated blood from the brain

is directed to the right ventricle and then
via ductus arteriosis to the dorsal aorta
 The haemogobin of foetal red blood cells has a higher
affinity for oxygen than that in maternal blood

OXYGEN SUPPLY - FOETAL ADAPTATIONS 3

100%
FOETAL
75% saturation MATERNAL
Hb as at 30 mm
HbO 2

50% 50% saturation

at 30 mm

30
pO 2 (mm Hg)

P50 maternal
P50 foetal
 REFERENCES

Cunningham JGC (2002) Textbook of Veterinary Physiology (Saunders)

Guyton and Hall (2005) Textbook of Medical Physiology (Elsevier)

Johnson MH (2007) Essential Reproduction (Blackwells)