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 EXTRAEMBRYONIC
MEMBRANES

Longitudinal view

The folding process not only creates the body form but also the extraembryonic structures of the yolk sac, allantois, amnion and chorion
Embryo proper
Primitive gut

Presumptive amnion

Allantois

Presumptive chorion

Transient chorio-vitelline placenta

Somatopleure

Yolk sac

Splanchnopleure

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

Mesoderm

Somatopleure

Neural tube
Ectoderm

Gut tube

Endoderm

Yolk sac

Splanchnopleure

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
Foetal capillary (from umbilical artery)
Endothelial layer

Connective tissue layer

(may be minimal)
Cellular layer
(may be trophoectoderm + maternal
epithelium or a syncytium of the two,
or solely trophoectoderm)
Connective tissue layer
(may be minimal)

EPITHELIOCHORIAL
Maternal endometrial epithelium intact
(horse,pig)
SYNEPITHELIOCHORIAL
Syncytium of maternal epithelium and
Chorion
(ruminants)
ENDOTHELIOCHORIAL
Removal of endometrial epithelium
And connective tissue
(dogs, cats)
HAEMOCHORIAL
Removal of maternal endothelium
(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
Uniform distribution of chorionic villi
over contact surface (horse, pigs)

COTYLEDONARY
Villi restricted to defined area
(cotyledons) (ruminants)

ZONARY
Girdle of chorionic villi around middle
of chorionic sac (dogs,cats)

DISCOIDAL
Disc-shaped area on chorionic sac
(humans, rodents)

The haemochorial placenta shows the intimate juxtaposition

of foetal and maternal blood allowing efficient exchange

Foetal capillaries

The haemochorial placenta

Umbilical vein

Umbilical arteries

Chorionic
villi

Maternal
blood pool

Maternal
venule
Maternal
arteriole

Notice the expansions

at theturnaround to
allow slower blood flow
and better equilibration
with maternal blood

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
SYNTHESIS

Maternal adipose
FATTY
ACIDS

LIPOPROTEINS
LPL
Maternal liver

OXIDATION

FATTY
ACIDS

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 (9,12,15)
18:2 (9,12)
20:4 (5,8,11,14)

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

AMINO
ACIDS

Maternal liver

UREA

OXIDATION
GROWTH

PROGESTERONE

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
25
14

To
lungs

Brachycephalic vessels
19

DA

From lungs

FO
25

19

14 Liver
30

Placenta

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

25

Trunk

1. Numbers are partial pressures of oxygen in

mm Hg

22

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

Hindlimb

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
MATERNAL

75% saturation
at 30 mm

Hb as
HbO2

50% saturation
at 30 mm

50%

30

pO2 (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)