Fertilization and

Development of the
Fetus and Placenta 2
C H A P T E R

LEARNING OBJECTIVES

At the end of this chapter, you should

be able to:
 Explain the conception process
 Explain the development of the
embryo
 Explain the development of the
placenta
 Explain the development of the fetal
skull
 Explain the disposition of the fetus

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INTRODUCTION
Human conception begins at the time of fertilization and development contin-
ues even after birth. Several factors can affect the growth and development of the
human conceptus, causing abnormalities and birth defects. This chapter deals
with the important aspects of fetal development and the processes involved. The
ovum and the sperm are the major female and male germ cells. A zygote or con-
ceptus results when the two unite.

CONCEPTION
Conception (fertilization) depends upon ovulation, patency of the genital tract
and a proper endometrial growth for the implantation of the fertilized egg in
the uterus. Similarly, the development of spermatid into mature spermatozoa,
a patent passage and proper ejaculation are necessary on the part of the male
(Figure 2.1). Lack of even one of these components can cause infertility, abor-
tion or an abnormal embryo.

Testis Ovary
Primary oocyte
Primary
46, XX
spermatocyte
First 46, XY
meiotic
division
23, X 23, Y
Secondary spermatocytes
Second First meiotic division
meiotic
division
Secondary
23, X 23, X 23, Y 23, Y oocyte 23, X
First polar
body
Second meiotic
division
Sperm
Second polar body
23, X
23, X 23, X 23, Y 23, Y Mature
oocyte 23, X

Figure 2.1 Spermatogenesis and oogenesis

Maturation of the ovum is known as oogenesis, and that of spermatozoa is

known as spermatogenesis.

Oogenesis
In the ovaries, oogenesis occurs before birth and completes only much
later during reproductive years. The primordial germ cells appear in the
yolk sac as early as the fourth week of gestation. The germ cells scatter in
the ovary and the stromal cells surrounding each germ cell form the flat
cells which after puberty develop into granulosa cells. Each unit contain-
ing the germ cells and a surrounding layer of granulosa cells is known as
primordial follicle (Figure 2.2).
The ovary of the fetus is packed with 47 million primordial follicles, but
many degenerate and only 400 000 follicles are present at puberty. During

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31
each menstrual cycle, several follicles start growing, but only one dominant
follicle grows and ovulates. During the entire reproductive period, only about
400 follicles are available for ovulation.
The development and morphology of the Graafian follicle has been described
earlier in Chapter 1.

Oocyte
The primitive ovum or oocyte begins its first meiotic division before birth,
but completes its meiotic division just before ovulation and forms a sec-
ondary oocyte and first polar body. This secondary oocyte is called ovum
after ovulation. The ovum is covered by an acellular structure called the
zona pellucida and scattered group of granulosa cells called corona radiata.
The ovum is picked up by the ovarian fimbria and propelled towards the
ampulla for fertilization.

Figure 2.2 Primordial follicle showing nucleus

Spermatogenesis
Spermatogenesis begins at puberty. Once the embryonic testes are formed, the
multiplication of germ cells ends and they enter into resting phase. At puberty,
these germ cells known as spermatogonia (Figure 2.1) start multiplying and
develop into spermatids and spermatozoa. The sperms develop from Sertoli
cells lining the seminiferous tubules. Spermatogenesis takes 72 days, and 18
days in the transport to the male genital tract. Unlike ovulation which occurs
only once a month, spermatogenesis is a continuous process.

Sperm
The sperm consists of a head and tail. The head contains condensed nucleus
and an acrosomal cap. The tail contains a neck, middle piece and end piece.

Fertilization
Fertilization occurs in the Fallopian tube when the sperm meets the ovum. The
ovum can be fertilized for up to 48 hours after release from the ovary, while the
sperm is viable for 3 days after ejaculation.

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Sperm First polar

body
Zona
pellucida

Figure 2.3 Fertilization

The sperms travel up the reproductive tract of the female to reach the tubes.
Release of prostaglandin from the sperm cap helps contract the uterine muscle
and makes the sperm reach the ampulla of the tube. Of the 500600 million
sperms released, only one penetrates the zona pellucida of the ovum (Figure
2.3), following which a chemical reaction occurs, making it impossible for other
sperms to penetrate the ovum.
In absence of fertilization, the ovum fails to undergo second meiotic division
and degenerates.
After the fertilization, the chromosome number is restored to 46, and the
zygote (fertilized egg) contains a mixture of paternal and maternal chromo-
somes.
The gonadal sex of the human being is determined by the sex chromosome
of the sperm. Y sex chromosome induces the development of the testes by the
The window of seventh or eighth week of embryonic life. Absence of Y chromosome results in
opportunity for a the development of the ovaries and a female baby by the tenth week. Abnor-
woman to get
pregnant is three
mal chromosome in either ovum or sperm results in infertility, abortion or
days before and 2 abnormal baby.
days after ovulation In an in vitro fertilization (test tube baby), capacitation and fertilization are
because of the developed in a culture medium and the early embryo is transferred into the
lifespan of the
sperms and egg.
uterine cavity during the late secretory phase of the menstrual cycle.

DEVELOPMENT OF THE EMBRYO

The fetal development is categorized into three stages:
1. Ovular stage from the time of fertilization to blastocyst formationlasts
for 3 weeks (up to 5 weeks amenorrhoea).
2. Embryonic stage from 3 weeks to 8 weeks when the embryo forms (10
weeks amenorrhoea).
3. Fetal development from 8 weeks post-fertilization. The development and
growth of the fetus occurs up to term (from 10 weeks amenorrhoea).
After its formation, the zygote undergoes mitotic division into two blastomeres,
approximately 30 hours after fertilization. Subsequent division occurs every
24 hours resulting in a cluster of smaller blastomeres known as morula around
the third post-fertilization day, when the morula enters the uterine cavity. On
the fourth post-fertilization day, the cells of the morula arrange themselves
into an inner cell mass of large cells and a peripheral layer of small cells, which
is known as trophoblast and is responsible for implantation of the zygote into
the endometrial line but takes no part in the development of the embryo. The

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Table 2.1 Post-fertilization growth
Time Event
0 hour Fertilization
30 hours 2-cell stage (blastomere)
50 hours 4-cell stage
96 hours 16-cell stagemorula enters the uterine cavity
5th day Blastocyst formation
7th day Zona pellucida degenerates and implantation occurs

inner cell mass develops into an embryo, yolk sac and amnion, and is referred
as embryonic cell mass (Table 2.1). The trophoblastic layer splits into an outer
layer of nucleated protoplasm without cell outline called the syncytium and
inner layer of large cells called cytotrophoblast or Langhans layer, and converts
a morula into a blastocyst (Figure 2.4).

Polar body

Zona pellucida

Blastomere

A. Two-cell stage B. Four-cell stage

C. Eight-cell stage D. Morula

Inner cell mass
Degenerating
zona pellucida

Trophoblast

Blastocyst cavity
E. Early blastocyst F. Late blastocyst

Figure 2.4 Early development of a fertilized egg

Implantation
The blastocyst contains 100 cells, each cell measuring 300 microns. At first,
it lies free in the uterine cavity for 2 days. During that period the zona pel-
lucida degenerates. On the seventh post-fertilization day, the blastocyst gets
implanted and gets embedded in the endometrium.
Following implantation, the trophoblast proliferates to form villi. The syn-
cytial layer penetrates the decidual lining (endometrium) and creates spaces
between the decidual cells and the trophoblastic layer. It also penetrates the
wall of the spiral arteries in the decidua and establishes a communication with
the maternal circulation.

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Functions of the trophoblastic layer are implantation of the zygote, provision
of oxygen and nutrition to the embryo, and formation of hormones like beta-
HCG, which maintains pregnancy till 12 weeks.

Ovular stage/Pre-embryonic stage

By the eighth day of fertilization, the inner cell mass is converted into a bilam-
inar germ disc, the dorsal ectoderm of columnar cells and the ventral endo-
derm of flattened polygonal cells. The endoderm speedily lines the trophoblast
to enclose a primitive yolk sac. The ectodermal layer encloses the amniotic
cavity on the dorsal aspect of the bilaminar disc.
The mesoderm is the last to form between the ectoderm and endoderm by
proliferation of cells at the ectoderm facing the amniotic cavity. The bilam-
inar disc is connected to the trophoblast by the mesenchymal connective stalk
which later forms the umbilical cord. The mesoderm splits into two layers. The
outer layer lines the trophoblast to form the chorion and inner layer goes with
the amnion and the endoderm.

Embryonic stage
During this phase, all the organ systems and the major structures develop,
making the embryo take a recognizable human form. Following the formation
of mesoderm in the third week of development, the bilaminar disc becomes
trilaminar and from it various tissues are formed. The ectoderm develops into
nervous system, skin and its appendages, and the pituitary gland.
The mesoderm forms the bones, cartilages, muscles, cardiovascular system,
suprarenal, kidneys, gonads, salivary glands, pleura, peritoneum and pericar-
dium.
The endoderm forms the epithelial lining of the gastrointestinal tract,
respiratory tract, bladder mucosa, liver, gall bladder and pancreas.
This is the phase of rapid cell division and differentiation. Exposure to terato-
genic agents in this period leads to the maximum fetal defects.

Fetal development
By the eighth post-fertilization week or tenth week from the last menstrual
period, the embryo has developed into a fetus with all identifiable human
features.
Thereafter, the fetus continues its growth in length and weight in a fixed
manner as shown in Table 2.2. The growth is characterized by cellular hyper-
plasia initially followed by hyperplasia and hypertrophy of the organs.
By the 32nd day, the sitting height of the embryo, i.e. crown-rump (CR)
length, is 5 mm. Up to the 55th day, the embryo grows by 1 mm per day and
thereafter.
CR length of 1.5 mm per day can be measured ultrasonically and duration of
gestation gauged. The amniotic sac measures 1 cm and a large yolk sac is seen
by the fifth week.
By the 12th week, the fetus measures 89 cm. During the fourth and fifth month,
the crown-heel (CH) length is gauged by Haases rule, i.e. CH length equals the
square of lunar months. CR length at the fourth month is 4 4 = 16 cm and
25 cm at the fifth month. From the seventh lunar month onwards, the CH length
in centimetres is the number of lunar months multiplied by 5 (Table 2.3).

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Table 2.2 Gestational age and fetal characteristics
Characteristic Gestational age
2837 weeks (premature) 3740 weeks (mature) Over 40 weeks (postmature)
Skin Skin is dull red in colour, lax Skin pink and smooth Skin wrinkled, cracks easily
and wrinkled due to absence and peels off
of subcutaneous fat
Subcutaneous Subcutaneous fat absent or Plenty of subcutaneous fat Scanty fat
fat scanty
Vernix Vernix caseosa scanty Long hair Long hair
Hair Short scalp hair Long hair Long hair
Nails Short nails Long nails Long nails
Lanugo Lanugo present Lanugo absent Lanugo absent
Skull sutures Skull sutures open Sutures closed except at the Sutures closed, hard skull
fontanelles
Crying Cries feebly Cries vigorously Asphyxia may be present at
birth

Table 2.3 Fetal age and size

Menstrual weeks Fetal age by CR length (mm) Fetal length (mm) Estimated fetal
(LMP) fertilization weight (g)
12 10 60 80 60
16 14 120 150 150180
20 18 160 200 300320
24 22 210 300 680
28 26 250 350 1000
32 30 280 400 15001800
34 32 300 420 2000
36 34 320 450 2500
38 36 340 480 2900
40 38 360 500 32003400

Weight
The fetal weight increases in a linear fashion up to about 20 weeks. After
that, the fetal weight is influenced by extraneous environment and maternal
nutrition. The average fetal weight at term varies between 2.5 and 3.5 kg and
increases with maternal age and parity. Diabetes causes increased weight and
the baby weighs more than 4.5 kg, whereas multiple pregnancy and hyperten-
sion causes growth-retarded fetus.
The fetal characteristics at various gestational ages are shown in Table 2.2.
At fifth week of amenorrhoea the gestation sac measures 1 cm as seen on ultra-
sound. At 6 weeks, it measures 23 cm and the embryo measures 45 mm. At
8 weeks amenorrhoea, the embryo has grown to 2024 mm.

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In early weeks, ultrasound recognizes the gestational sac and yolk sac, and
measures CR length.
Later in pregnancy, the femoral length, the biparietal diameter, head cir-
cumference and abdominal circumference indicate the gestational age of the
fetus. The fetal growth can be measured serially every fortnight. The fetal sex
and most of the structural abnormalities can be detected by the 16th week.
The fetal weight can be gauged by studying the fetal volume. After 32 weeks,
the physiological activities and its well-being can be observed by Manning score,
which comprises parameters such as fetal tone, limb movements, fetal breathing,
amniotic fluid volume and non-stress test. Doppler ultrasound studies the fetal
blood flow in various vessels such as umbilical artery, aorta and cerebral vessels.

DEVELOPMENT OF THE PLACENTA (DECIDUA)

Soon after fertilization and implantation, the endometrium grows into decidua
under the stimulus of progesterone secreted by the persistent corpus luteum
of pregnancy. The decidual reaction first appears at the implantation site, then
spreads over the entire surface of the endometrium.
Decidua is categorized as follows (Figure 2.5):
1. Decidua basalis at the implantation site, where the placenta develops.
2. Decidua capsularis which covers the blastocyst.
3. Decidua vera which lines the uterine cavity.

Decidua vera
Decidua
capsularis

Embryo

Decidua
basalis
Decidual
space

Figure 2.5 Early pregnancy showing an embryo

Decidua

Knowledge of the Amniotic

developmental stages cavity
helps to ascertain
gestational age
during pregnancy,
identify normal and
abnormal fetus by
its size and establish
norms of fetal growth
in different ethnic
groups.
Figure 2.6 Decidual space is obliterated: fetus and deciduas 12 weeks pregnancy

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The space between the decidua capsularis and decidua vera persists until 12
weeks of pregnancy. As the fetal sac expands, this space gets obliterated by the
fusion of two deciduae (Figure 2.6).
Morphology: The decidua consists of a deep layer of endometrial glands
called cavernous layer and a superficial layer of compact stromal cells contain-
ing dilated spiral arteries.
The placenta/decidua provides:
 A nidus for implantation.
 Nutrition and oxygen to the embryo.
 Prevents deep penetration by the trophoblast.
 It secretes prolactin, relaxin and prostaglandin, which is responsible for
initiation of labour at term.

Hormonal control
As mentioned earlier, fertilization causes the corpus luteum to persist and con-
tinue secreting progesterone. Progesterone is responsible for decidual growth
and maintenance of pregnancy up to the 10th12th week when the placenta
develops and carries on with the pregnancy.

DEVELOPMENT OF THE FETAL SKULL

The fetal skull comprises the base of the skull and face which are firm and non-
compressible. The vault is made of pliable bones joined together by unossified
membranes which form the sutures and fontanelles. The vault, therefore, can
undergo a change in shape and size called moulding during labour under the
pressure of uterine contractions and resistance in the bony pelvis.
The vault consists of two parietal bones on either side, the frontal bones in
front and the occipital bone behind. The sagittal suture crosses the vault in the
midline between the two parietal bones. In the same plane in front of the ante-
rior fontanelle lies the frontal suture between the two frontal bones. The suture
separating the frontal bones from the parietal bones is known as the coronal
suture (Figure 2.7). The lambdoid suture separates the parietal bones from the
occipital bone. These sutures allow the overlapping of the bones during labour

Suboccipito-bregma
tic
Flexed diameter 9.5 cm
Submento-
vertex
bregmatic l
ca
diameter rti
9.5 cm - ve
to r
en te Brow
M ame
Occipito-frontal d i cm
space 11 cm 14 Deflexed
vertex

Partially
Face deflexed vertex

Figure 2.7 Normal fetal skull with diameters

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Anterior fontanelle
(bregma)
SINCIPUT VERTEX

Occipito-frontal
Glabella diameter

Posterior
Nasion fontanelle

OCCIPUT

Suboccipito-
bregmatic
diameter

Supraoccipito-
mental diameter
Submento-bregmatic
diameter

Figure 2.8 Anterior fontanelle

when the head meets the resistance by the pelvic bones in the pelvis. Thus,
the size of the fetal head is reduced, and this allows an easy delivery. This
physiological moulding of up to 5 mm is beneficial for a safe vaginal delivery.
A normal moulding disappears in a few hours after birth.
Excessive overlapping or moulding can cause harm to the fetal brain, internal
cerebral haemorrhage and fetal death. In a preterm baby, even a slight moulding
can inflict intracranial injury. Therefore, utmost care is required during a vaginal
delivery of a preterm baby.
The anterior fontanelle is called bregma which is a kite-shaped unossified
membrane lying between the two frontal bones in front and the parietal bones
behind. The anterior fontanelle is felt as a depression about 3 2 cm (Figure
2.8). The posterior fontanelle is a depression produced by the occiput bone
being depressed below the posterior borders of the parietal bones (Figure 2.9).
These two fontanelles are of clinical importance. They can be identified during
vaginal examination during labour, and from their position, the presentation

Frontal eminence
The posterior
fontanelle closes at
the third month in
the newborn. The Anterior fontanelle
anterior
fontanelle closes at
about 18 months.
In the newborn, an
increased size of
the fontanelle can
raise the suspicion Parietal eminence
of hydrocephalus. Posterior fontanelle
Whereas, depressed
fontanelle is seen in
dehydration. Figure 2.9 Posterior fontanelle

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and the fetal attitude can be accurately gauged. The moulding will also indicate
the degree of obstruction in the cephalopelvic relationship.
The occiput is the area behind the occipital bone and the sinciput is the
frontal portion of the vault consisting of frontal bones. The occiput is broader
compared to the sinciput, and this can be appreciated during abdominal palpa-
tion of the fetal head.
Vertex is the area on the vault of the skull which is bounded anteriorly by
the anterior fontanelle and coronal sutures, posteriorly by the posterior fonta-
nelle and the lambdoid suture, and laterally by the parietal eminence (Figure
2.10). Depending upon the degree of flexion of the head over its body, shape
of the head, its circumference and its diameter of contact with the pelvic brim,
the diameter of engagement varies and can affect the mode of delivery.

Occipital bone

Lambdoid suture

Posterior fontanelle

Biparietal diameter

Parietal eminence

Sagittal suture

Coronal suture Bitemporal diameter

Frontal bone

Anterior fontanelle
(bregma)
Frontal suture

Figure 2.10 Vertex

Table 2.4 Diameters of engagement in a term fetus

Diameter of engagement Length Presentation
Suboccipito-bregmatic diameter (nape of the neck to the 9.4 cm (3 inches) Completely flexed vertex
centre of bregma)
Suboccipito-frontal diameter (nape of the neck to the 10.0 cm (4 inches) Incompletely flexed head
anterior end of bregma)
Occipito-frontal diameter (occipital protuberance to the 11.3 cm (4 inches) Occipito-posterior-
root of the nose) extended vertex
Mento-vertical diameter (point of chin to a point 1 inch in 13.8 cm (5 inches) Brow presentation
front of posterior fontanelle on the sagittal suture)
Submento-vertical diameter (angle between the chin and 11.2 cm (4 inches) Incompletely extended face
neck to the centre of sagittal suture)
Submento-bregmatic diameter (angle between the chin 9.4 cm (3 inches) Complete extension with
and neck to the centre of bregma) face presentation

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In passing from complete flexion to complete extension of the head, the
diameters of engagement of a term fetus are described in Table 2.4.
The three transverse diameters of the fetal head are given below.
 Biparietal diameter (9.4 cm): It is the distance between the two parietal
eminences. When this diameter enters the pelvic brim, the head is said to
be engaged.
 Bitemporal diameter (8 cm): It is the distance between the antero-inferior
ends of the coronal sutures.
 Bimastoid diameter (7.5 cm): It is the distance between the tips of mastoid
process.
A successful outcome in vaginal delivery depends upon:
 Size of the fetal head
 Diameter of engagement
 Size of the fetus and its disposition
 Amount of amniotic fluid
 Pelvic size
 Strength of the uterine contractions

THE DISPOSITION OF THE FETUS

In the last few weeks of pregnancy, the fetus is packed in the smallest
possible space in an orderly manner, and this is termed as the fetal atti-
tude (Figure 2.11). In a normal pregnancy, this disposition is the attitude of
flexion. In this attitude, the head is flexed, so the chin touches the thorax,
the spine is flexed, the thighs are flexed over the trunk and the legs over
the thighs. The limbs however exhibit movements every now and then and
these are perceived both by the pregnant woman and the obstetrician. The
attitude of flexion enables the fetus to adopt and accommodate itself to the
shape and size of the uterus.

The estimation of Figure 2.11 Attitude of flexion

fetal head size before
labour is done
clinically as well In a vertex presentation normally, the fetus remains in the attitude of flexion
as by ultrasound and forms an ovoid mass with the head occupying the lower uterine segment
measurements of and the body lying and fitting appropriately in the pear-shaped uterine cavity
the biparietal and
occipito-frontal
with a broad fundus. The diameters of the fetal ovoid are:
diameters, to decide  Verticopodalic: 2425 cm
if vaginal delivery is  Bisacromial: 12 cm
possible.
 Bitrochanteric: 10 cm

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The smallest circumference of the fetal head in a fully flexed fetus is
2728 cm.

Caput succedaneum
In the first stage of labour, the pressure of the uterine contractions and the
bony pelvis on the fetal head is through the bag of water, so there is no change
over the scalp. Once the membranes rupture and amniotic fluid is drained, the
fetal scalp comes under the direct compressive forces. The fetal scalp below
the compression develops an effusion or oedema due to obstruction to the
venous blood flow. This swelling is known as caput succedaneum. The size
of the caput depends upon the duration of the pressure. It can be huge in
prolonged labour and premature rupture of membranes. The site of the caput
is related to the fetal presentation. In the left occipito-anterior presentation,
caput forms over the right parietal bone and in the right occipito-anterior pres-
entation, it forms over the left parietal bone. In face presentation, it presents
as a bluish swelling over the face. The caput disappears in 12 days, and leaves
no trace of its ugly appearance.
Caput is recorded as zero when absent, one plus when minimal and two plus
when moderate. The caput is considered excessive when it is three plus and is
a warning sign of prolonged labour.

KEY POINTS
 Fertilization occurs in the ampullary portion of the Fallopian tube when the sperm fuses with the ovum.
 Implantation occurs around the seventh post-fertilization day.
 The endometrium develops into decidua.
 The fertilized egg develops into an embryo through the ovular stage which lasts 3 weeks after fertilization.
 The embryonic stage extends from the third to eighth week of post-fertilization period. During this period,
inner cell mass and trophoblast develop. The inner cell mass differentiates into trilaminar plate and forms a
fetus.
 In the fetal stage, by the tenth week, the fetus continues to grow in length and weight till term.
 The trophoblast invades the decidua, creates a choriodecidual space and establishes feto-maternal
circulation by penetrating into the spiral arteries.
 The body stalk develops into the umbilical cord.
 The fetal skull is the most important structure as it is the largest and least compressible.
 The vertex is the leading point on the fetal head.
 The diameters of the fetal head are important in determining whether a vaginal delivery is possible.
 In the cephalic presentation with a flexed fetal head, the suboccipito-bregmatic diameter is the engaging
diameter and is 9.4 cm.
 The flexed fetal attitude presents the smallest diameter of the fetal head to the maternal pelvis.
 The part of the fetus closest to the cervix is the presenting part.
 The relationship of the fetal axis to the maternal axis is called the lie, and the commonest fetal lie is the
longitudinal lie.
 The caput succedaneum forms after the membranes have ruptured.
 A large caput is a warning sign of prolonged labour.

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MULTIPLE-CHOICE QUESTIONS
1 Which is the process of maturation of the female egg?
A Spermatogenesis
B Conception
C Oogenesis
D Gestation
2 Which is the process of maturation of the sperm?
A Conception
B Gestation
C Oogenesis
D Spermatogenesis
3 The initial mitotic division of the zygote after fertilization results in the formation of which of the following
cells?
A Follicles
B Blastomeres
C Oocytes
D Spermatocytes
4 Which layer helps in the implantation of the zygote?
A Myometrium
B Cortex
C Medulla
D Trophoblast
5 The bones on either side of the vault of the fetal skull are called
A parietal bones
B occipital bones
C frontal bones
D maxilla bones
6 Which suture is situated between the two parietal bones?
A Coronal suture
B Lambdoid suture
C Sagittal suture
D Frontal suture

ESSAY QUESTIONS
1 List the stages of fetal development highlighting the major events.
2 Discuss in brief the process of placental development and functions.

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