Ploidy: Number of sets of

chromosomes in a cell
Haploid (n)-- one set chromosomes
Diploid (2n)-- two sets chromosomes
Eggs and sperm are haploid (n)
 Developmental stages of
spermatogenesis
In the course of spermatogenesis the germ cells
move towards the lumen as they mature.

A-spermatogonium
B-spermatogonium
Primary spermatocyte (= spermatocyte order I)
Secondary spermatocyte (= spermatocyte order II)
Spermatid
Sperm cell (= spermatozoon)
 The spermatogenesis can be subdivided into
two successive sections:
The first comprises the cells from the
spermatogonium up to and including the
secondary spermatocyte and is termed
spermatocytogenesis.
The second one comprises the
differentiation/maturation of the sperm cell,
starting with the spermatid phase and is
termed spermiogenesis (or
spermiohistogenesis).
 The approximate 64 day cycle of the
spermatogenesis can be subdivided into four
phases that last differing lengths of time:

Mitosis of the spermatogonia-- 16 days--- Up to the

primary spermatocytes

First meiosis-- 24 days-- For the division of the primary

spermatocytes to form secondary spermatocytes
Second meiosis-- A few hours-- For the spermatids
Spermiogenesis-- 24 days--- Up to the completed
sperm cells
Total~ 64 days
 Spermatocytogenesis
Among the spermatogonia (all in all,
over 1 billion in both testicles) that form
the basal layer of the germinal epithelium,
several types can be distinguished:
certain type A cells are seen as
spermatogonia that divide mitotically and
reproduce themselves where by the
spermatogonia population is maintained.
 The beginning of spermatogenesis in which the
daughter cells (second group of type A cells)
remain bound together by thin bridges of
cytoplasm. Through the preservation of these
cytoplasmic connections, spermatogonia are
inducted into the spermatogenesis process.

After a further mitotic division type B

spermatogonia are engendered that also
divide themselves mitotically into primary
spermatocytes (I).
 The freshly created primary spermatocytes
(I) now enter into the first meiosis.
They then go immediately into the S phase
(that is, into the preleptotene meiosis),
double their internal DNA, leave the basal
compartment and reach the special milieu of
the luminal compartment.
Following the S phase, these cells attain the
complex stage of the prophase of the
meiosis and become thereby noticeably
visible with a light microscope.
 This prophase, which lasts 24 days, can be
divided into five sections:
Leptotene
Zygotene
Pachytene
Diplotene
Diakinesis
 In the prophase in every germ cell a
new combination of maternal and
paternal genetic material occurs.
After the long prophase follow the
metaphase, anaphase and telophase
that take much less time.
One primary spermatocyte yields two
secondary spermatocytes.
 The secondary spermatocytes go directly into
the second meiosis, out of which the spermatids
emerge.
Since in the secondary spermatocytes neither
DNA reduplication nor a recombination of the
genetic material occurs, the second meiosis can
take place quickly.
It lasts only around five hours and for that reason
secondary spermatocytes are rather seldom seen
in a histological section.
Through the division of the chromatids of a
secondary spermatocyte, two haploid spermatids
arise that contain only half the original DNA
content.
 Besides the sperm cells the
spermatids are the smallest cells of
the germinal epithelium. In a process
lasting several weeks (so-called
spermiogenesis or
spermiohistogenesis) they are
transformed into spermatocytes with
the active assistance of the Sertoli's
cells.
spermatogenesis wave
 As in the diagram, spermatogenesis
waves move in spirals - like a
corkscrew - towards the inner part of
the lumen.
Outside, on the edge of the tubule
and at the beginning of the spiral, lie
the spermatogonia; and, at the end of
the spiral, the fully developed sperm
cells are in the lumen.
 Spermiogenesis
(spermatohistogenesis) and
structure of the sperm cell

The differentiation of the spermatids into

sperm cells is called spermiogenesis. It
corresponds to the final part of
spermatogenesis and comprises the
following individual processes that partially
proceed at the same time:
 Nuclear condensation: thickening and
reduction of the nuclear size,
condensation of the nuclear contents into
the smallest space.
Acrosome formation: Forming a cap
(acrosome) containing enzymes that play
an important role in the penetration
through the pellucid zone of the oocyte.
Flagellum formation: generation of the
sperm cell tail.
Cytoplasma reduction: elimination of all
unnecessary cytoplasm.
 1.Axonemal structure,
first flagellar
primordium
2.Golgi complex
3.Acrosomal vesicle
4.Pair of centrioles
(distal and proximal)
5;Mitochondrion
6.Nucleus
7.Flagellar primordium
8.Microtubules
9.Sperm cells tail
10.Acrosomal cap
 Nuclear condensation
The nucleus becomes smaller, denser and
takes on a characteristic, flattened form.
Seen from above, the nucleus is oval and,
from the narrow side, is pear-shaped.
The acrosome lies over the tip. Nucleus and
acrosome form the sperm cell's head that is
bound to the mid-piece by a short neck.
 Acrosome formation
The Golgi complex engender the
vesicles, which then merge into a
larger formation that settles close to
the cell nucleus and finally inverts itself
like a cap over the largest part of the
nucleus. The acrosome corresponds
functionally to a lysosome and thus
contains lysosomal enzymes
(hyaluronidase among others).
 Development of the
flagellum
The future axonemal structure
grows out of one centriole
(distal). This consists of a
bundle of nine peripheral
double microtubules and two
single ones in the center.
 Four parts of the finished flagellum can be
distinguished:
The neck contains the two centrioles (proximal
and distal) among other things.
The mid piece consists of a sheath of ring-
shaped mitochondria grouped around the
axoneme to provide the energy for the flagellar
movement.
The principle piece has a sheath of ring fibers
around the axoneme.
The tail consists of only the 9+2 structure of the
axoneme
The mature sperm cell is approximately 60 mm
long and completely enveloped by the plasma
membrane.
1. 1. Plasma membrane
2.Outer acrosomal membrane
3.Acrosome
4.Inner acrosomal membrane
5.Nucleus
6.Proximal centriole
7.Rest of the distal centriole
8.Thick outer longitudinal fibers
9.Mitochondrion
10.Axoneme
11.Anulus
12.Ring fibers
A.Head
B.Neck
C.Mid piece
D.Principal piece
E.Endpiece
The mature sperm cell is slender; in the middle part, the mitochondria are thick and
ring-shaped. The DNA in the nucleus is maximally condensed.
 Leydig's interstitial cells and
hormonal regulation
Between the seminal canals lie Leydig's
interstitial cells. These are endocrine cells that
mainly produce testosterone, the male sexual
hormone, and release it into the blood and into
the neighboring tissues.
An initial active stage of these cells occurs during
the embryonic development of the testis. Later in
juvenile life, due to the influence of the LH
(luteinizing hormone) secreted by the anterior
hypophysis (pituitary gland), Leydig's interstitial
cells enter a second, long lasting stage of activity.
Together with the hormones secreted by the
adrenal cortex, testosterone initiates puberty and
thus the maturation of the sperm cells.
 Testosterone production is directed by LH
(luteinizing hormone), secreted by the anterior
lobe of the hypophysis.

There by testosterone can be transported by

Sertoli's cells into the luminal compartment
and there be concentrated. Testosterone is
decisive for spermatogenesis.

Testosterone is also carried away via blood

and lymph fluid. Testosterone has effects on all
tissues, especially also on the brain during
development as well as on the sexual organs.
 Oogenesis
Following the immigration of the primordial germ
cells into the gonadal ridge, they proliferate, are
enveloped by coelomic epithelial cells, and form
germinal cords that , though, keep their connection
with the coelom epithelium.
Now a cortical zone (cortex ovarii) and a medulla can
be distinguished, whereby it should be mentioned that
in females the germinal cords never penetrate into the
medullary zone.
In the genital primordium the following processes then
take place:
 A wave of proliferation begins that lasts
from the 15th week to the 7th month:
primary germ cells arise in the cortical
zone via mitosis of oogonia clones,
bound together in cellular bridges, that
happens in rapid succession.
The cell bridges are necessary for a
synchronous onset of the subsequent
meiosis.
 With the onset of the meiosis (earliest
onset in the prophase in the 12th week)
the designation of the germ cells changes.
They are now called primary oocytes.
The primary oocytes become arrested in
the diplotene stage of prophase I (the
prophase of the first meiotic division).
 Shortly before birth, all the fetal oocytes in the
female ovary have attained this stage. The
meiotic resting phase that then begins is called
the dictyotene and it lasts till puberty, during
which each month (and in each month thereafter
until menopause) a pair of primary oocytes
complete the first meiosis.
Only a few oocytes (secondary oocytes plus one
polar body), though, reach the second meiosis
and the subsequent ovulation. The remaining
oocytes that mature each month become atretic.
The primary oocytes that remain in the ovaries
can stay in the dictyotene stage up to
menopause, in the extreme case, without ever
maturing during a menstrual cycle.
 While the oogonia transform into primary
oocytes, they become restructured so that at
the end of prophase I (the time of the
dictyotene) each one gets enveloped by a
single layer of flat, follicular epithelial cells
(descendents of the coelomic epithelium).
(oocyte + follicular epithelium = primordial
follicle).
 On the one hand, the female germ cell
that at birth is called the primary oocyte,
and which can develop further only during
(and after) puberty (hormonal cycle is
necessary).
On the other hand, the follicular epithelium
that can develop further from the
primordial follicle via several follicle
stages while oocytes remain in their
primary state.
 The developmental sequence of
the female germ cells is as
follows:
Primordial germ cell -
oogonium - primary oocyte -
primary oocyte in the
dictyotene
 The developmental sequence of a
follicle goes through various follicle
stages:
Primordial follicle - primary follicle -
secondary follicle - tertiary follicle
(graafian follicle)
Since a follicle can die at any moment in
its development (= atresia), not all reach
the tertiary follicle stage.
 Structure of the ovary

An ovary is subdivided into cortical

(ovarian cortex) and medullary
compartments (ovarian medulla).
Both blood and lymph vessels are found in
the loose connective tissue of the ovarian
medulla.
In the cortical compartment the oocytes
are present within the various follicle
stages.
 The sex hormones influence the primordial follicles to
grow and a restructuring to take place.
From the primordial follicles the primary follicles,
secondary follicles, and tertiary follicles develop in turn.
Only a small percentage of the primordial follicles reach
the tertiary follicle stage - the great majority meet their
end beforehand in the various maturation stages.
Large follicles leave scars behind in the cortical
compartment and the small ones disappear without a
trace.

The tertiary follicles get to be the largest and, shortly

before ovulation, can attain a diameter up to 2.5 mm
through a special spurt of growth. They are then termed
graafian follicles.
 1.Primordial follicle
2.Primary follicle
3.Secondary follicle
4.Tertiary follicle
5.Antrum folliculi
6.Cumulus oophorus
 The follicle stages from primordial
follicle to tertiary follicle
Primordial follicle
At the time of birth all the surviving primary
oocytes are surrounded by thin, single layers
of so-called follicular epithelial cells. These are
delimited from the rest of the ovarian stroma by
a thin basal lamina.
Follicular epithelial cells are former coelomic
epithelial cells. The primordial follicles always
form the majority of the follicles in the ovary.
 Under the influence of the sex hormones
some of them are able to develop
further to one or more.
Although this further development can
already take place sporadically in the time
before birth and up to puberty, the main
part occurs as soon as a regular hormonal
cycle is established.
Particularly the last phase of the
maturation of a tertiary follicle to become a
large follicle, ready to rupture, remains
reserved for the time of regular cycles.
 Primary follicle
In the transition of the
primordial follicles into
primary follicles.
 A.Primordial follicle
B.Primary follicle
1.Oocyte
2.Follicular epithelium
 Secondary follicle
When primary follicles survive, secondary follicles
with follicular epitheliums encompassing multiple
rows are engendered.
Their identifying characteristic is a fluid-filled cavity,
the antral follicle.
This is now called the stratum granulosum. In the
secondary follicles a glycoprotein layer, the pellucid
zone, between the oocyte and follicular epithelium
becomes visible.
Cytoplasmic processes of the granulosa cells that lie
upon it reach the oocyte through the pellucid zone
and thereby assure their maintenance function.
Outside the basal lamina the stroma ovarii organizes
itself to become theca folliculi cells.
Secondary follicle
 1.Oocyte
2.Pellucid zone
3.Stratum granulosum
4.Theca folliculi cells
 Tertiary follicle
If the secondary follicles survive, tertiary follicles
are engendered.

The oocyte lies at the edge in a mound made of

granulosa epithelial cells, the cumulus
oophorus.
In the meantime it has grown so large that its
cellular nucleus has attained the size of a whole
primordial follicle.
The connective tissue around the follicle has
already clearly differentiated itself into a theca
interna, well supplied with capillaries, out of
large, lipid-rich cells (hormone production) and a
theca externa, which forms a transition to the
stroma ovarii and contains larger vessels.
 1.Oocyte
2.Pellucid zone
3.Stratum granulosum
4.Theca interna
5.Theca externa
6.Antral follicle
7.Cumulus oophorus (Granulosa cells,
together with the oocyte)
8.Basal lamina between theca and
stratum granulosum
 Decisive for a successful follicle
growth is a well-developed net of
capillaries in the theca interna.
The precise steering mechanism that
leads to the selection of a follicle and
its subsequent maturation to become
a graafian follicle is still unknown.
Before ovulation a growth spurt of the
tertiary follicles takes place.
 Graafian follicle
This corresponds to an
especially large tertiary follicle
that can be expected to suffice
for ovulation.
 Phase A:
Primordial germ cells grow, proliferate and become sheathed with coelomic
epithelial cells. Gonadal cords arise; 6th to 8th week.

Phase B:
Spurt of growth: cellular clones of the oogonia are formed, whereby the
cells remain connected with each other through cellular bridges; 9th to the
22nd week.

Phase C:
The oogonia become primary oocytes that enter the prophase of the first
meiosis; 12th to the 25th week.

Phase D:
The primary oocytes become arrested in the dictyotene stage of the
prophase: the primordial follicles are engendered; 16th to the 29th week.

Phase E:
At around the 14th week a quantitatively increased decline in the number of
germ cells commences as well as atresia in all of the follicle stages.
 One terms the decline or the
regression of follicles of each stage at
every time in the life of a woman
follicular atresia.
These follicles do not ovulate and the
name is derived from that fact.
Follicle atresia occurs more
intensely, though, at certain moments
(fetal period, early postnatal, begin of
the menarche).
 Explanations for the onset of heightened atresia
are:
a) in the 14th week:
Already during the meiotic prophase, mainly in the
pachytene stage (lasts the longest, around 3
weeks), the cells are especially susceptible and
succumb.
With the formation of the primordial follicle in the
16th week the follicular atresia also begins as an
additional reason for the decline in the number of
germ cells.
Both processes together cause the germ cell count
to shrink to a third (somewhat over 2 million / ovary)
of the maximum number.
 postnatal:
During the fetal period sex hormones are produced
in the placenta. This results in a high estrogen level
in the blood of the mother and of the fetus.
This gives rise to a considerable maturation of the
primordial follicles up to the tertiary follicle phase in
the female fetus.
When the sex hormone level in the fetus sinks
after birth all of the previously matured follicles
become atretic (a slight withdrawal bleeding can
even occur in the newly born from the 5th to the
10th day).
Per ovary there are then less than 2 million germ
cells present afterwards.
 during puberty:
With the onset of puberty (at around 12 years of age) an
elevated production of estrogen occurs again, which
leads to a maturation of the inner and outer gender
attributes.
After puberty is past, around 250'000 germ cells pro
ovary remain. With the onset of a regular cycle a nearly
linear decline commences that with 40 years of age
increases.
With the continuous decrease of the follicle cell count the
production of estrogen is also reduced constantly. With
roughly 50 years of age when follicles are no longer
present, the estrogen production ceases and
menopause ensues
It is assumed the dictyotene stage of meiosis represents
a special condition for oocytes that exhibits great stability
with regard to outer physical and chemical influences. In
contrast, the earlier stages, especially the pachytene,
are more sensitive.
 The ovarian cycle

Of the roughly 500'000 follicles that are

present in the two ovaries at the beginning
of sexual maturity, only around 480 reach
the graafian follicle stage and are thus
able to release oocytes (ovulation). This
number is simply derived by multiplying
the number of cycles per year (12) and the
number of years in which a woman is
fertile (40).
 The hormonal cycle:
Cyclic changes in the hormone household (hormonal
cycle), governed by the hypothalamic-pituitary
system, are responsible for the periodicity of the
ovulation. In a woman, the rhythmic hormonal
influence leads to the following cyclic events:
the ovarian cycle (follicle maturation) that peaks in
the ovulation and the subsequent luteinization of the
granulose cells
cyclic alterations of the endometrium that prepare
the uterine mucosa so fertilized oocytes can "nest"
there. In the absence of implantation, the mucosa
will be eliminated (menstrual bleeding)
 In the center of this hormonal control is the
hypothalamamics-hypophysial (pituitary
gland) system with the two hypophysial
gonadotropins FSH and LH.
The pulsating liberation of GnRH by the
hypothalamus is the fundamental precondition
for a normal control of the cyclic ovarian
function.
This cyclic activity releases FSH and LH, both of
which stimulate the maturation of the follicles in
the ovary and trigger ovulation.
During the ovarian cycle, estrogen is produced
by the theca interna and follicular cells (in the
so-called follicle phase) and progesterone by the
corpus luteum (so-called luteal phase).
 The control circuit of the hormonal cycle
has two essential control elements:
The pulsatile liberation of GnRH, as well
as FSH and LH
 As a rule, the ovarian cycle lasts 28
days. It is subdivided into two phases:
Follicle phase: recruitment of a so-called
follicle cohort and, within this, the selection
of the mature follicle. This phase ends
with ovulation.
Estradiol is the steering hormone.
Normally, it lasts 14 days, but this can
vary considerably!
Luteal phase: progesteron production
by the "yellow body" (= corpus luteum)
and lasts 14 days (relatively constant).
 A.Follicle phase
B.Luteal phase
C.Primary follicle
D.Secondary follicle
E.Tertiary follicle
F.Graafian follicle
E2 Estradiol
Pr Progesterone
LH Luteinizing hormone
FSH Follicle stimulating hormone
 Cell Type Cell Division Product
Spermatagonium (2N)Undergoes Mitosis
Primary Spermatocyte (2N)
Primary Spermatocyte (2N)Undergoes
Meiosis I & results in
2 Secondary Spermatocytes (1N)
Secondary Spermatocyte (1N)Undergoes
Meiosis II, results in Spermatid (1N).
Spermatid (1N)--Maturation - no cell division--
Sperm cell (1N)
 Oogonium (2N)development - no cell division--
Primary Oocyte (2N).
Primary Oocyte (2N)Undergoes Meiosis I ---
Secondary Oocyte (1N) and First Polar Body
(1N).
Secondary Oocyte (1N)Undergoes Meiosis II
(stimulated by fertilization from sperm)-----
Ovum (1N) and Second Polar Body (1N)