Chapter |2|
Conception and implantation
Roger Pepperell
LEARNING OUTCOMES
After studying this chapter you should be able to:
Knowledge criteria
• Describe the basic principles of the formation of the
gametes
• Describe the physiology of the normal menstrual cycle
• Describe the physiology of coitus, fertilization and
implantation  egg by a sperm, the chromosome count is returned to the
Clinical competency
• Counsel a couple about the fertile period
Oogenesis
Primordial germ cells originally appear in the yolk sac and
can be identified by the fourth week of fetal development
(Fig. 2.1). These cells migrate through the dorsal mesen-
tery of the developing gut and finally reach the genital
ridge between 44 and 48 days post-conception. Migration
occurs into a genital tubercle consisting of mesenchymal
cells that appear over the ventral part of the mesonephros.
The germ cells form sex cords and become the cortex of
the ovary.
The sex cords subsequently break up into separate
clumps of cells and, by 16 weeks, these clumped cells
become primary follicles, which incorporate central germ
cells.
These cells undergo rapid mitotic activity, and by 20
weeks of intrauterine life, there are about 7 million cells,
known as oogonia. After this time, no further cell division
occurs and no further ova are produced. By birth, the oogo-
nia have already begun the first meiotic division and have
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become primary oocytes. The number of primary oocytes
falls progressively and by birth is down to about 1 million
and to about 0.4 million by puberty.
Meiosis
The process of meiosis results in 23 chromosomes being
found in each of the gametes, half the number of chromo-
somes found in normal cells. With the fertilization of the
normal count of 46 chromosomes. Fusion of the sperm and
the egg occurs when the first of two meiotic divisions of the
oocyte has already been completed; with the second mei-
otic division occurring subsequently and being completed
prior to the 23 chromosomes of the male gamete joining
those of the female gamete within the nucleus of the cell,
the zygote is formed, which will become the embryo.
In meiosis, two cell divisions occur in succession, each
of which consists of prophase, metaphase, anaphase and
telophase. The first of the two cell divisions is a reduction
division, and the second is a modified mitosis in which the
prophase is usually lacking (Fig. 2.2). At the end of the first
meiotic prophase, the double chromosomes undergo syn-
apsis, producing a group of four homologous chromatids
called a tetrad. The two centrioles move to opposite poles.
A spindle forms in the middle, and the membrane of the
nucleus disappears. During this prophase period of meiosis
I, the double chromosomes, which are closely associated
in pairs along their entire length, undergo synapsis, cross-
ing over and undergoing chromatid exchange, with these
processes accounting for the differences seen between two
same-sex siblings despite the fact that the female gametes
came from the same mother.
The primary oocytes remain in suspended prophase
until sexual maturity is reached, or even much later, with
meiosis I not recommencing until the dominant follicle
is triggered by luteinizing hormone (LH) to commence
   Conception and implantation Chapter |  2 |

Yolk Primordial germ Reach genital Primary follicle with 7 million cells
sac cells identified ridge central germ cell (oogonia)

Sex cords formed = Mitotic

cortex of gonads = activity No further
separate clumps cell division
of cells or ova

0 2 4 6 8 10 12 14 16 18 20
Gestation (weeks)
Fig. 2.1  Embryonic and fetal development of oogonia.

the development of a small second polar body. In the

Prophase
male the original cell containing 46 chromosomes ulti-
mately results in four separate spermatozoa, each being
of the same size but containing only 23 chromosomes
(see Spermatogenesis, later). 

Metaphase
Anaphase
Telophase
Meiosis
Fig. 2.2  Primary oocytes remain in suspended prophase. Mei-
otic division resumes under stimulation by luteinizing hormone.
ovulation. In anaphase, the daughter chromatids separate
and move towards opposite poles. Meiosis II commences
around the time the sperm are attached to the surface of
the oocyte and is completed prior to the final phase of
fertilization.
Thus, the nuclear events in oogenesis are virtu-
ally the same as in spermatogenesis, but the cytoplas-
mic division in oogenesis is unequal, resulting in only
one secondary oocyte. This small cell consists almost
entirely of a nucleus and is known as the first polar body.
As the ovum enters the Fallopian tube, the second mei-
otic division occurs and a secondary oocyte forms, with
Follicular development in the ovary
The gross structure and the blood supply and nerve sup-
ply of the ovary have been described in Chapter 1. How-
ever, the microscopic anatomy of the ovary is important in
understanding the mechanism of follicular development
and ovulation.
The surface of the ovary is covered by a single layer
of cuboidal epithelium. The cortex of the ovary contains
a large number of oogonia surrounded by follicular cells
that become granulosa cells. The remainder of the ovary
consists of a mesenchymal core. Most of the ova in the
cortex never reach an advanced stage of maturation and
become atretic early in follicular development. At any
given time, follicles can be seen in various stages of matu-
ration and degeneration (Fig. 2.3). About 800 primary
follicles are ‘lost’ during each month of life from soon
after puberty until menopause, with only one or two of
these follicles resulting in release of a mature ovum each
menstrual cycle in the absence of ovarian hyperstimula-
tion therapy. This progressive loss occurs irrespective of
whether the patient is pregnant, on the oral contracep-
tive pill, having regular cycles or amenorrhoeic, with
menopause occurring at the same time irrespective of the
number of pregnancies or cycle characteristics. The vast
majority of the follicles lost have undergone minimal or
no actual maturation.
The first stage of follicular development is character-
ized by enlargement of the ovum with the aggregation
of stromal cells to form the thecal cells. When a domi-
nant follicle is selected at about day 6 of the cycle, the

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 Section |  1 | Essential reproductive science
Blood vessels Follicle: double layered
Egg nest Mature follicle
Ovum
Corpus Ruptured
luteum follicle
Fig. 2.3  Development and maturation of the Graafian follicle.
innermost layers of granulosa cells adhere to the ovum
and form the corona radiata. A fluid-filled space devel-
ops in the granulosa cells, and a clear layer of gelatinous
material collects around the ovum, forming the zona pel-
lucida. The ovum becomes eccentrically placed, and the
Graafian follicle assumes its classic mature form. The
mesenchymal cells around the follicle become differen-
tiated into two layers, forming the theca interna and the
theca externa.
As the follicle enlarges, it bulges towards the surface of
the ovary and the area under the germinal epithelium thins
out. Finally, the ovum, with its surrounding investment
of granulosa cells, escapes through this area at the time of
ovulation.
The cavity of the follicle often fills with blood but, at
the same time, the granulosa cells and the theca interna
cells undergo the changes of luteinization to become
filled with yellow carotenoid material. The corpus luteum
in its mature form shows intense vascularization and pro-
nounced vacuolization of the theca and granulosa cells
with evidence of hormonal activity. This development
reaches its peak approximately 7 days after ovulation, and
thereafter the corpus luteum regresses unless implanta-
tion occurs, when human chorionic gonadotropin (hCG)
production by the implanting embryo prolongs corpus
luteum function until the placenta takes over this role at
about 10 weeks of gestation. The corpus luteum degen-
eration is characterized by increasing vacuolization of the
granulosa cells and the appearance of increased quantities
of fibrous tissue in the centre of the corpus luteum. This
finally develops into a white scar known as the corpus albi-
cans (Fig. 2.4).  Blood levels of FSH are slightly higher during menses and
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Zona pellucida Granulosa cells –
corona radiata
Mature
follicle
Antrum formation –
follicular fluid
Ruptured Corpus luteum Primary
follicle follicle
Egg nest
Fig. 2.4  Ovulation and corpus luteum formation.
Hormonal events associated with
ovulation
The maturation of oocytes, ovulation and the endometrial
and tubal changes of the menstrual cycle are all regulated
by a series of interactive hormonal changes (Fig. 2.5).
The process is initiated by the release of the gonadotro-
phin-releasing hormone (GnRH), a major neurosecretion
produced in the median eminence of the hypothalamus.
This hormone is a decapeptide and is released from axon
terminals into the pituitary portal capillaries. It results in
the release of both follicle-stimulating hormone (FSH) and
LH from the pituitary.
GnRH is released in episodic fluctuations, with an
increase in the number of surges being associated with
the higher levels of plasma LH commencing just before
mid-cycle and continued ongoing GnRH action required
to initiate the huge oestrogen-induced LH surge. Kiss-
peptin levels in the serum and urine have been studied
during the menstrual cycle and found to be useful in the
prediction of ovulation. During the first 5 days of the
cycle, it is low. There is a surge around the eleventh day
when the dominant follicle is about 1.2 cm. The second
surge, which is smaller, is around the fourteenth day.
Serum kisspeptin levels correlate well with 17-β oestra-
diol (E2) levels. It appears that kisspeptin surge may be a
good marker of the dominant follicle development prior
to ovulation.
The three major hormones involved in reproduction
are produced by the anterior lobe of the pituitary gland
or adenohypophysis and include FSH, LH and prolactin.
   Conception and implantation Chapter |  2 |

Pituitary gland
GnRH – median eminence
of hypothalamus

Posterior lobe Anterior lobe

Gonadotrophin (mU/mL serum)

LH FSH Prolactin
70 LH
LSH
Oestradiol (nmol/L plasma)
50

1.5 Progesterone (nmol/L plasma)

30 50

0.75
25
10

0 7 14 21 28 0 7 14 21 28 0 7 14 21 28
A Days B Days C Days

Fig. 2.5  The hormonal regulation of ovulation. Gonadotrophin-releasing hormone (GnRH) stimulates the release of gonadotrophins
from the anterior lobe of the pituitary. Blood levels of (A) luteinizing hormone (LH) and follicle-stimulating hormone (FSH); (B) oestra-
diol; and (C) progesterone during a 28-day menstrual cycle. LSH, Lutein-stimulating hormone.

subsequently decline due to the negative feedback effect

of the oestrogen production by the dominant follicle. LH
levels appear to remain at a relatively constant level in the
first half of the cycle; however, there is a marked surge of
LH 35–42 hours before ovulation and a smaller coinciden-
tal FSH peak (see Fig. 2.5). The LH surge is, in fact, made
up of two proximate surges, and a peak in plasma oestra-
diol precedes the LH surge. Plasma LH and FSH levels are
slightly lower in the second half of the cycle than in the pre-
ovulatory phase, but continued LH release by the pituitary
is necessary for normal corpus luteum function. Pituitary
gonadotrophins influence the activity of the hypothalamus
by a short-loop feedback system between the gonadotro-
phins themselves and the effect of the ovarian hormones
produced due to FSH and LH action on the ovaries.
Oestrogen production increases in the first half of the
cycle, then falls to about 60% of its follicular phase peak
following ovulation, and a second peak occurs in the luteal
phase. Progesterone levels are low prior to ovulation but
then become elevated throughout most of the luteal phase.
These features are shown in Fig. 2.5.
Certain feedback mechanisms regulate the release
of FSH and LH by the pituitary. This is principally
achieved by the oestrogens and progesterone produced
by the ovaries. In the presence of ovarian failure, as seen
in menopause, the gonadotrophin levels become mark-
edly elevated because of the lack of ovarian oestrogen
and progesterone production. Inhibin B levels were
thought to predict the ovarian reserve in infertile women
but have been shown not to be that useful.
Prolactin is secreted by lactotrophs in the anterior lobe
of the pituitary gland. Prolactin levels rise slightly at mid-
cycle but are still within the normal range and remain at
similar levels during the luteal phase and tend to follow
the changes in plasma oestradiol-17 β levels. Prolactin
tends to control its own secretion predominantly through
a short-loop feedback system on the hypothalamus, which
produces the prolactin-inhibiting factor, dopamine. Oes-
trogen appears to stimulate prolactin release in addition to

15
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 Section |  1 | Essential reproductive science

the release of various neurotransmitters, such as serotonin,
noradrenaline (norepinephrine), morphine and enkepha-
lins, by a central action on the brain. Antagonists to dopa-
mine such as phenothiazine, reserpine and methyltyrosine
also stimulate the release of prolactin, whereas dopamine
agonists such as bromocriptine and cabergoline have the
opposite effect.
Hyperprolactinaemia prevents ovulation by an
inhibitory effect on hypothalamic GnRH production
and release and is an important cause of secondary
amenorrhoea and infertility.
with only the dominant follicle then getting enough FSH
to continue further development. At the same time, FSH
stimulates receptors for LH.
LH stimulates the process of ovulation, the reactiva-
tion of meiosis I and sustains the development of the cor-
pus luteum; receptors for LH are found in the theca and
granulosa cells and in the corpus luteum. There is a close
interaction between FSH and LH in follicular growth and
maturation. The corpus luteum produces oestrogen and
progesterone until it begins to deteriorate in the late luteal
phase (see Fig. 2.4). 

The endometrial cycle

The action of gonadotrophins
FSH stimulates follicular growth and development and
binds exclusively to granulosa cells in the growing follicle.
Of the 30 or so follicles that begin to mature in each men-
strual cycle, one becomes pre-eminent and is called the
dominant follicle. The granulosa cells produce oestrogen,
which feeds back on the pituitary to suppress FSH release,
The normal endometrium responds in a cyclical manner
to the fluctuations in ovarian steroids. The endometrium
consists of three zones, and it is the two outer zones that are
shed during menstruation (Fig. 2.6).
The basal zone (zona basalis) is the thin layer of the com-
pact stroma that interdigitates with the myometrium and

$ %

&

Fig. 2.6  Cyclical changes in the normal menstrual cycle. (A) Proliferative phase. (B) Mid-luteal phase. (C) Menstrual phase.

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   Conception and implantation Chapter |  2 |

shows little response to hormonal change. It is not shed
at the time of menstruation. The next adjacent zone (zona
spongiosa) contains the endometrial glands, which are lined
by columnar epithelial cells surrounded by loose stroma.
The surface of the endometrium is covered by a compact
layer of epithelial cells (zona compacta) that surrounds the
ostia of the endometrial glands. The endometrial cycle is
divided into four phases:
1. Menstrual phase. This occupies the first 4 days of the cy-
cle and results in shedding of the outer two layers of the
endometrium. The onset of menstruation is preceded
by segmental vasoconstriction of the spiral arterioles.
This leads to necrosis and shedding of the functional
layers of the endometrium. The vascular changes are
associated with a fall in both oestrogen and progester-
one levels, but the mechanism by which these vascular
changes are mediated is still not understood. What is
clear clinically is that the menstruation due to the shed-
ding of the outer layers of the endometrium occurs
whether oestrogen or progesterone, or both, fall, with
the loss generally being less if both the oestrogen and
progesterone levels fall (as at the end of an ovulatory
cycle), and heavier when only the oestrogen level falls
(as in an anovulatory cycle).
2. Phase of repair. This phase extends from day 4 to day 7
and is associated with the formation of a new capillary
bed arising from the arterial coils and with the regenera-
tion of the epithelial surface.
3. Follicular or proliferative phase. This is the period of
maximal growth of the endometrium and is associated
with elongation and expansion of the glands and with
stromal development. This phase extends from day 7
until the day of ovulation (generally day 14 of the cy-
cle).
4. Luteal or secretory phase. This follows ovulation and
continues until 14 days later when menstruation starts
again. During this phase, the endometrial glands be-
come convoluted and ‘saw-toothed’ in appearance. The
epithelial cells exhibit basal vacuolation, and by the
mid-luteal phase (about day 20 of a 28-day cycle), there
is visible secretion in these cells. The secretion subse-
quently becomes inspissated and, as menstruation ap-
proaches, there is oedema of the stroma and a pseudo-
decidual reaction. Within 2 days of menstruation, there
is infiltration of the stroma by leukocytes.
It is now clear that luteinization of the follicle can
occur in the absence of the release of the oocyte, which
may remain entrapped in the follicle. This condition is
described as entrapped ovulation or luteinized unruptured
follicle (LUF) syndrome and is associated with normal pro-
gesterone production and an apparently normal ovulatory
cycle. Histological examination of the endometrium gen-
erally enables precise dating of the menstrual cycle and is
particularly important in providing presumptive evidence
of ovulation. 
Production of sperm
Spermatogenesis
The testis combines the dual function of spermatogenesis
and androgen secretion. FSH is predominantly responsible
for stimulation of spermatogenesis and LH for the stimula-
tion of Leydig cells and the production of testosterone.
The full maturation of spermatozoa takes about 64–70
days (Fig. 2.7). All phases of maturation can be seen in the
testis. Mitotic proliferation produces large numbers of cells
(called spermatogonia) after puberty until late in life. These
spermatogonia are converted to spermatocytes within the
testis, and then the first meiotic division commences. As in
the female, during this phase, chromatid exchange occurs,

Spermatogenesis
≡ 64 days
Spermatogonia 2N diploid 2N haploid 1N haploid
Diploid Diploid Primary Secondary Spermatids Spermatozoon
FSH spermatocyte spermatocyte
+
LH

Mitotic First meiotic Second meiotic

division division division Spermiogenesis

42 days 22 days

Fig. 2.7  The maturation cycle of spermatozoa.

17
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 Section |  1 | Essential reproductive science
resulting in all gametes being different despite coming from
the same original cell. Spermatocytes and spermatids are
produced from the spermatogonia. Spermatozoa are finally
produced and released into the lumen of the seminiferous
tubules and then into the vas deferens. At the time of this
final release, meiosis II has been completed. Full capacitation
of the sperm, to enable fertilization to occur, is not achieved
until the sperm have passed through the epididymis and
seminal vesicles, augmented by a suitable endocrine envi-
ronment in the uterus or Fallopian tube and finally when the
spermatozoon becomes adherent to the oocyte. 
Structure of the spermatozoon
The spermatozoon consists of a head, midpiece and tail
(Fig. 2.8). The head is flattened and ovoid in shape and is
covered by the acrosomal cap, which contains several lysins.
The nucleus is densely packed with the genetic mate-
rial of the sperm. The midpiece contains two centrioles,
proximal and distal, which form the beginning of the tail.
The distal centriole is vestigial in mature spermatozoa but
is functional in the spermatid. The body contains a coiled
helix of mitochondria that provides the ‘powerhouse’ for
sperm motility.
The tail consists of a central core of two longitudinal
fibres surrounded by nine pairs of fibres that terminate at
various points until a single ovoid filament remains. These
contractile fibres propel the spermatozoa.  sperm migrate at a rate of 6 mm/min. This is much faster
Seminal plasma
Spermatozoa carry little nutritional reserve and therefore
depend on seminal plasma for nutritional support. Seminal
plasma originates from the prostate, the seminal vesicles,
Head Acrosomal cap
Neck Basal body
Mitochondrial
Midpiece sheath
Annulus
Tail Fibrous sheath
Fig. 2.8  Structure of the mature spermatozoon.
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the vas deferens and the bulbourethral glands. There is a
high concentration of fructose, which is the major source of
energy for the spermatozoa. The plasma also contains high
concentrations of amino acids, particularly glutamic acid,
and several unique amines such as spermine and spermidine.
Seminal plasma also contains high concentrations of
prostaglandins, which have a potent stimulatory effect on
uterine musculature. Normal semen clots shortly after ejac-
ulation but liquefies within 30 minutes through the action
of fibrinolytic enzymes. 
Fertilization
The process of fertilization involves the fusion of the male
and female gametes to produce the diploid genetic comple-
ment from the genes of both partners.
Sperm transport
Following the deposition of semen near the cervical os,
migration occurs rapidly into the cervical mucus. The speed
of this migration depends on the presence of receptive mucus
in mid-cycle. During the luteal phase, the mucus is not recep-
tive to sperm invasion and, therefore, very few spermatozoa
reach the uterine cavity. Under favourable circumstances,
than could be explained by the motility of the sperm and
must therefore also be dependent on active support within
the uterine cavity. Only motile spermatozoa reach the fim-
briated end of the tube, where fertilization occurs. 
Capacitation
During their passage through the Fallopian tubes, the
sperm undergo the final stage in maturation (capacitation),
which enables penetration of the zona pellucida. It seems
likely that these changes are enzyme induced, and enzymes
such as β-amylase or β-glucuronidase may act on the mem-
branes of the spermatozoa to expose receptor sites involved
in sperm penetration. In addition, various other factors
that may be important in capacitation have been identi-
fied, such as the removal of cholesterol from the plasma
membrane and the presence of α- and β-adrenergic recep-
tors on the spermatozoa. Until recently, it was thought that
capacitation occurred only in vivo in the Fallopian tubes.
However, it can also be induced in vitro by apparently non-
specific effects of relatively simple culture solutions.
Inhibitory substances in the plasma of the cauda epidid-
ymis and in seminal plasma can prevent capacitation, and
these substances also exist in the lower reaches of the female
genital tract. It seems likely that these substances protect the
sperm until shortly before fusion with the oocyte. 

Spermatozoon
Zona pellucida
A Nucleolus
Spermatozoon
B large and pale; this is known as the decidual reaction.
C
Fig. 2.9  (A) Adherence of the sperm to the oocyte initiates the
acrosome reaction. (B, C) Syngamy involves the passage of the
nucleus of the sperm head into the cytoplasm of the oocyte
with the formation of the zygote.
Fertilization and implantation
Only a small number of spermatozoa reach the oocyte in
the ampulla of the tube and surround the zona pellucida.
The adherence of the sperm to the oocyte initiates the acro-
some reaction, which involves the loss of plasma membrane
over the acrosomal cap (Fig. 2.9A).
The process allows the release of lytic enzymes, which
facilitate penetration of the oocyte membrane. Generally,
only one sperm head fuses with the oocyte plasma mem-
brane, and by phagocytosis the sperm head and midpiece
are engulfed into the oocyte.
The sperm head decondenses to form the male pro-
nucleus and eventually becomes apposed to the female
pronucleus in the female egg to form the zygote. The mem-
branes of the pronuclei break down to facilitate the fusion
of male and female chromosomes. This process is known
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  Conception and implantation Chapter |  2 |
as syngamy (see Fig. 2.9B, C) and is followed almost imme-
diately by the first cleavage division.
During the 36 hours after fertilization, the conceptus
is transported through the tube by muscular peristaltic
action. The zygote undergoes cleavage and, at the 16-cell
stage, becomes a solid ball of cells known as a morula. A
fluid-filled cavity develops within the morula to form the
blastocyst (Fig. 2.10). Six days after ovulation, the embryonic
pole of the blastocyst attaches itself to the endometrium,
usually near to the mid-portion of the uterine cavity. By the
seventh post-ovulatory day, the blastocyst has penetrated
deeply into the endometrium.
Endometrial cells are destroyed by the cytotrophoblast,
and the cells are incorporated by fusion and phagocytosis
into the trophoblast. The endometrial stromal cells become
The processes of fertilization and implantation are now
complete. 
The physiology of coitus
Normal sexual arousal has been described in four levels
in both the male and the female. These levels consist of
excitement, plateau, orgasmic and resolution phases. In
the male, the excitement phase results in compression of the
venous channels of the penis, resulting in erection. This
is mediated through the parasympathetic plexus through
S2 and S3. During the plateau phase, the penis remains
engorged and the testes increase in size, with elevation of
the testes and scrotum. Secretion from the bulbourethral
glands results in the appearance of a clear fluid at the ure-
thral meatus. These changes are accompanied by general
systemic features, including increased skeletal muscle ten-
sion, hyperventilation and tachycardia.
Erectile dysfunction may result from neurological
damage to the spinal cord or the brain and is seen
as a result of spina bifida, multiple sclerosis and diabetic
neuropathy. However, over 200 prescription drugs are
known to cause impotence, and these account for some
25% of all cases. Recreational drugs such as alcohol,
nicotine, cocaine, marijuana and LSD may also cause
impotence; however, this can usually be improved by the
male taking the pharmacological preparation sildenafil
citrate (Viagra). Sexual stimulation causes local release
of nitric oxide. Inhibition of phosphor diesterase type
5 (PDE5) results in increased levels of cyclic guanosine
monophosphate (cGMP) in the corpus cavernosum.
This leads to smooth muscle relaxation and inflow of
blood to the corpus cavernosum, causing the erection.
Hence the drug has no effect in the absence of sexual
stimulation.
19
 Section |  1 | Essential reproductive science
6th day
post-ovulation
Blastocyst
Fig. 2.10  Stages of development from fertilization to implantation.
The orgasmic phase is induced by stimulation of the glans
penis and by movement of penile skin on the penile shaft.
There are reflex contractions of the bulbocavernosus and
ischiocavernosus muscles and ejaculation of semen in a
series of spurts. Specific musculoskeletal activity occurs that
is characterized by penile thrusting. The systemic changes
of hyperventilation and rapid respiration persist.
Seminal emission depends on the sympathetic nervous
system. Expulsion of semen is brought about by contrac-
tion of smooth muscle within the seminal vesicles, ejacula-
tory ducts and prostate.
During the resolution phase, penile erection rapidly sub-
sides, as do the hyperventilation and tachycardia. There is a
marked sweating reaction in some 30–40% of individuals.
During this phase, the male becomes refractory to further
stimulation. The plateau phase may be prolonged if ejacu-
lation does not occur.
In the female, the excitement phase involves nipple
and clitoral erection, vaginal lubrication (resulting partly
from vaginal transudation and partly from secretions
from Bartholin’s glands), thickening and congestion of
the labia majora and the labia minora and engorgement
of the uterus. Stimulation of the clitoris and the labia
results in progression to the orgasmic platform, with nar-
rowing of the outer third of the vagina and ballooning
of the vaginal vault. The vaginal walls become congested
and purplish in colour, and there is a marked increase in
20
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Morula
Ovum
vaginal blood flow. During orgasm, the clitoris retracts
below the pubic symphysis and a succession of contrac-
tions occurs in the vaginal walls and pelvic floor approx-
imately every second for several seconds. At the same
time, there is an increase in pulse rate, hyperventilation
and specific skeletal muscular contractions. Blood pres-
sure rises, and there is some diminution in the level of
awareness. Both intravaginal and intrauterine pressures
rise during orgasm.
The plateau phase may be sustained in the female and
result in multiple orgasms. Following orgasm, resolution of
the congestion of the pelvic organs occurs rapidly, although
the tachycardia and hypertension, accompanied by a sweat-
ing reaction, may persist.
Factors that determine human sexuality are far more
complex than the simple process of arousal by clitoral or
penile stimulation. Although the frequency of intercourse
and orgasm declines with age, this is in part mediated by
loss of interest by the partners. The female remains capable
of orgasm until late in life, but her behaviour is substan-
tially determined by the interest of the male partner. Sexual
interest and performance also decline with age in the male,
and the older male requires more time to achieve excite-
ment and erection. Ejaculation may become less frequent
and forceful.
Common sexual problems are discussed in Chapter 19.
   Conception and implantation Chapter |  2 |

Essential information

Oogenesis Seminal plasma

• Primordial germ cells appear in the yolk sac • Originates from the prostate, seminal vesicles and
• By 20 weeks, there are 7 million oogonia bulbourethral glands
• Number of oocytes falls to 1 million by birth • High concentration of fructose provides energy for sperm
• Number falls to about 0.4 million by puberty motility
• Chromosome number in gametes is half that of normal • High concentration of prostaglandins 
cells
• Primary oocyte remains in suspended prophase for Sperm transport
10–50 years • Rapid migration into receptive cervical mucus
• The second meiotic division commences as the ovum • Sperm migrate at 6 mm/min
enters the tube  • Only motile sperm reach the fimbriated ends of the
tubes 
Follicular development in the ovary
• Most ova never reach advanced maturity, and about 800 Capacitation
are lost each month • Final sperm maturation occurs during passage through
• Aggregation of stromal cells around follicles become the oviduct
thecal cells • Inhibitory substances produced in caudoepididymis and
• Innermost layers of granulosa cells form the corona in seminal plasma 
radiata
• After ovulation, the corpus luteum is formed  Fertilization
• Small number of sperm reaches oocyte
Hormonal events and ovulation • Adherence of sperm initiates the acrosome reaction
• FSH stimulates follicular growth • Sperm head fuses with oocyte plasma membrane
• FSH stimulates LH receptor development • Sperm head and midpiece engulfed into oocyte
• LH stimulates ovulation and stimulates and sustains • Fusion of male and female chromosomes is known as
development of the corpus luteum syngamy
• Follicles produce oestrogen • Thirty-six hours after fertilization, the morula is formed
• Corpus luteum produces oestrogen and progesterone  • Six days after fertilization, implantation occurs 

The endometrial cycle Physiology of coitus

• Menstrual phase – shedding of functional layer of
endometrium
• Phase of repair – day 4–7 of cycle
• Follicular phase – maximum period of growth of
endometrial glands due to oestrogen
• Luteal phase – ‘saw-toothed’ glands, pseudodecidual
reaction in stroma  glands secretions
Spermatogenesis
• Full maturation takes 64–70 days
• Mature sperm arise from haploid spermatids 
• Penile erection results from compression of venous
channels
• Ejaculation mediated by contractions of bulbocavernosus
and ischiocavernosus
• Female excitation results in nipple and clitoral erection
• Lubrication comes from vaginal transudation, Bartholin’s
• Orgasm results in clitoral retraction and contractions of
pelvic floor muscles

Structure of spermatozoon
• Head is covered by acrosomal cap
• Body contains helix of mitochondria
• Tail consists of two longitudinal fibres and nine pairs of
fibres 

21
Descargado para DIANA ESTEFANY VARAS MAGUIÑA (gabi_dev2009@hotmail.com) en Universidad Cesar Vallejo de ClinicalKey.es por Elsevier en septiembre 04,
2020. Para uso personal exclusivamente. No se permiten otros usos sin autorización. Copyright ©2020. Elsevier Inc. Todos los derechos reservados.