Oogenisis 1

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Year 1 Sem 1 ENG

C4c

1
• Plan

• Gametogenesis in
female (Oogenesis)
Definition
Oogenesis is the process of formation of the female gamete (oocyte).

In embryos with an XX sex chromosome configuration (in absence of Y


chromosome) the primordial germ cells evolve towards the female gamete.

Oogenesis is described in two periods: prenatal and postnatal.

I. Prenatal period of oogenesis

• Primordial germ cells follow


the steps of the indifferent
period:
 formation of the
primordial cells,
 migration,
formation of the
genital (gonadal) ridge.
Prenatal period of oogenesis
• Primordial germ cells arrive at the end
of the 4th week (beginning of the 5th
week) in the genital (gonadal) ridges
and invade it in the 6th week. They are
called now oogonia

• In the 7th week (as cells without


chromosomes Y in their nuclei!), they
induce the development of the genital
(gonadal) ridges into female gonad.

• Primordial germ cells differentiate into


oogonia and evolve in the same time
with the organogenesis of the ovary.

1 Proliferating coelomic epithelium 8 Intestinal tube


2 Thickening of the mesenchyme 9 Dorsal mesentery
3 Germinal cords 10 Gonadal ridge
4 Primordial germ cells (red dots) 11 Nephrogenic cord
5 Mesenchyme 12 Mesonephric (Wolffian) duct
6 Allantois 13 Mesonephric tubule
7 Vitelline 14 Aorta
I. Prenatal period of oogenesis
1. Organogenesis of the ovary

The coelomic epithelium of the genital ridges proliferates and penetrates the
underlying mesechyme and forms the primitive sex cords (indifferent
gonad).
The primitive sex cords dissociate into irregular cell clusters and then
disappear.
I. Prenatal period of oogenesis
1. Organogenesis of the ovary
The surface epithelium of the female gonad continues to proliferate and in the
7th week it gives rise to a second generation of cords that remain in contact with
the surface epithelium and are called cortical cords.

In the 4th month the cortical cords split into cells that surround the primordial
germ cells.

The epithelial cells (of somatic origin) surrounding the oogoniae are called
follicular cells. This cell complex formed by oogonia and follicular cells remain
under the surface epithelium of the female gonad (cortical development).
I. Prenatal period of oogenesis
2. Evolution of the primordial germ cells

Most of the oogonia continue their


divisions by mitosis and their number
increase. Some of them differentiate
into larger cells – primary oocyte. By
the 5th month, precursors of the
female gamete (oogonia and primary
oocytes) are almost 7 million.

Then, they begin to degenerate and


die (apoptosis!) so that, by the 7th
month, only a few cells remain under
the surface on the gonad. They enter
the prophase of the 1st meiotic
division and they remain in this
stage (resting, dormant stage) until
puberty.
I. Prenatal period of oogenesis
2. Evolution of the primordial germ cells
A primary oocyte together with its surrounding epithelial cells (simple
squamous epithelium) is called primordial follicle.
II. Postnatal period of oogenesis
1. Before puberty

Short time before birth all


primary oocytes are in the
prophase of the 1st
meiotic division. They
remain in this stage
(without finishing the 1st
meiotic division) –
diplotene stage, until
puberty.

This resting (dormant)


period is due (probably!) to
an oocyte maturation
inhibitor factor produced
by the follicular cells.
II. Postnatal period of oogenesis
1. Before puberty

At birth, the number of primary oocytes


varies between 700.000 and 2 million. Each
female is born with all of the oocytes she
will ever have in her lifetime!

During childhood, most of them degenerate


(atresia!), so that by the beginning of the
puberty only 40.000 are present in both
gonads. During the reproductive lifetime (25-
30 years), only 500 oocytes will be ejected
from the ovary.
II. Postnatal period of oogenesis
2. Beginning with the puberty
Beginning with the puberty, 5-12(15) primordial follicles start their maturation
each ovarian cycle.

One can distinguish a follicular cycle and an oogenic cycle both under the
control of a hormonal cycle.
II. Postnatal period of oogenesis
2. Beginning with the puberty
a) The follicular cycle (folliculogenesis)

During this process, the primordial follicle


passes through successive stages:

•primary follicle,
•secondary follicle,
•tertiary (Graafian) follicle.
II. Postnatal period of oogenesis
2. Beginning with the puberty
A. The follicular cycle (folliculogenesis)
a) The primary follicle

During this stage, the follicular cells


of the primordial follicle surrounding
the primary oocyte transform from
flat to cuboidal, proliferate and
produce a stratified epithelium of
granulosa cells. These cells lay on
a membrane called theca folliculi
(Köllicker-Slavianski).
II. Postnatal period of oogenesis
2. Beginning with the puberty
A. The follicular cycle (folliculogenesis)
a) The primary follicle

Both granulosa cells and the


oocyte secrete a layer of
glycoproteins that surrounds
the oocyte and is called zona
pellucida. It is an incomplete
barrier between the follicular
cells and the oocyte, that the
sperm can penetrate only
due to its acrosomal
enzymes.
II. Postnatal period of oogenesis
2. Beginning with the puberty
A. The follicular cycle (folliculogenesis)
a) The primary follicle

Cells of theca folliculi of the


continuing growing follicle differentiate
into two layers:

•theca interna, the inner layer, formed


of secretory cells,
•theca externa, the outer layer,
formed of connective tissue containing
fibroblast-like cells.

They also send finger-like processes


that cross zona pellucida and
interdigitate with the microvilli of the
plasma membrane of the oocyte.
II. Postnatal period of oogenesis
2. Beginning with the puberty
A. The follicular cycle (folliculogenesis)
b) The secondary follicle

In one side of the follicle, fluid-filled spaces


appear between granulosa cells. They unite
and form an eccentric cavity called antrum.

Granulosa cells surrounding the oocyte form


cumulus oophorus.
II. Postnatal period of oogenesis
2. Beginning with the puberty
A. The follicular cycle (folliculogenesis)
b) The tertiary follicle

The antrum enlarges and the


follicle in this stage is called
antral, vesicular or Graafian

(de Graaf, a Dutch anatomist and physician who described the maturing ovarian
follicle in a major work on the female reproductive cycle published in 1672 — a
year before he died of the plague aged 32) who made key discoveries
in reproductive biology.
II. Postnatal period of oogenesis
2. Beginning with the puberty
B. The oogenic cycle

As soon as the follicle is


mature, shortly before
ovulation, the primary oocyte
resumes its 1st meiotic
division and forms two
daughter-cells with 23
chromosomes each (result
of the meiosis!) (22+X), but
of unequal size and
importance:
•the secondary oocyte
(containing all
cytoplasm of the
primary oocyte),
•the 1st polar body.
II. Postnatal period of oogenesis
2. Beginning with the puberty
B. The oogenic cycle

Shortly after the 1st meiotic division is


completed, the secondary oocyte enters the
2nd meiosis without DNA replication.

During this stage, ovulation occurs and the


secondary oocyte remain in the metaphase of
the 2nd meiosis. It is completed only if the
oocyte is fertilized. Otherwise, the
secondary oocyte degenerates in almost 24
hours.
II. Postnatal period of oogenesis
2. Beginning with the puberty
C. The hormonal cycle
Organs involved:
•hypothalamus,
•pituitary gland,
•ovaries.

It consists in two phases:


•follicular,
•luteal

Oogenic and follicular cycles are controlled


by the hypothalamus.
By means of the gonadotropin-releasing
hormone (GnRH) it acts on the anterior
pituitary gland which secretes
gonadotropin hormones:
•follicle-stimulating hormone (FSH)
• luteinizing hormone (LH).
II. Postnatal period of oogenesis
2. Beginning with the puberty
C. The hormonal cycle
a) Follicular phase

At the beginning of each


ovarian cycle primordial
follicles begin to develop
under the influence of FSH.
Only one of them reaches
the full maturity. Most of
them degenerate and
transform in corpus
atreticum.
II. Postnatal period of oogenesis
2. Beginning with the puberty
B. The hormonal cycle
a) Follicular phase

Follicular and thecal cells


secrete estrogens:
•they act on the uterine
endometrium and induce
the proliferative phase.
•they stimulate the pituitary
gland to secrete LH.
II. Postnatal period of oogenesis
2. Beginning with the puberty
B. The hormonal cycle
b) Luteal phase

On about day 13-14 of the


menstrual cycle levels of FSH
and LH rise suddenly.
II. Postnatal period of oogenesis
2. Beginning with the puberty
B. The hormonal cycle
b) Luteal phase

Coincident with final


development of the Graafian
follicle, the primary oocyte (in
the dormant stage) finishes
its 1st meiotic division, begins
promptly the 2nd meiotic
division, but stops its
evolution in the metaphase (3
hours before ovulation).
II. Postnatal period of oogenesis
2. Beginning with the puberty
B. The hormonal cycle
b) Luteal phase
The cumulus cells surrounding the
oocyte loose their cell-to-cell
connections and the oocyte together with
some cumulus cells detach into the
antral cavity.

Cumulus cells accompanying the oocyte


have a role in:
•transport of the oocyte,
•fertilization,
•early development of the zygote.

The follicle forms a bulge on the surface


of the ovary and a small protrusion
called stigma, appears.
II. Postnatal period
of oogenesis
2. Beginning with the puberty
B. The hormonal cycle
b) Luteal phase

Factors producing the extrusion of


the oocyte:
•tension produced by the
smooth muscle cells in the
follicular wall
•release of collagen-
degrading enzymes by
fibroblasts
•increasing of intrafollicular
pressure
•degeneration of the ovarian
surface
II. Postnatal period of oogenesis
2. Beginning with the puberty
B. The hormonal cycle
b) Luteal phase

Cumulus oophorus cells


rearrange around the zona
pellucida to form the corona
radiata.
II. Postnatal period of oogenesis
2. Beginning with the puberty
B. The hormonal cycle
b) Luteal phase

After ovulation, granulosa cells of the


ruptured follicle together with cells from
theca interna, under the influence of
LH, develop a yellowish pigment and
change into the luteal cells that form
corpus luteum.
It secretes progesterone that prepares
the endometrium (secretory stage) for
an eventual implantation of the embryo.
It secretes also estrogene.
II. Postnatal period of oogenesis
2. Beginning with the puberty
B. The hormonal cycle
b) Luteal phase

If fertilization fails to occur, after


9 days corpus luteum
decreases in size because of
the degeneration of luteal cells
and transform in corpus
albicans.
II. Postnatal period of oogenesis
2. Beginning with the puberty
B. The hormonal cycle
b) Luteal phase

Levels of progesterone and


estrogens decrease
precipitating menstrual
bleeding.
II. Postnatal period of oogenesis
2. Beginning with the puberty
B. The hormonal cycle
b) Luteal phase

If fertilization takes place


degeneration of the corpus
luteum is prevented by
chorionic gonadotropin (hCG)
secreted by the trophoblast of
the developing embryo.
Corpus luteum transforms
into corpus luteum of
pregnancy. It secretes
progesterone until the 4th
month when placenta replace
its function.
Normal gametogenesis: conversion of germ
cells into gametes. The illustrations compare
spermatogenesis and oogenesis. Oogonia are
not shown in this figure because they
differentiate into primary oocytes before birth.
The chromosome complement of the germ cells
is shown at each stage. The number
designates the total number of chromosomes,
including sex chromosome(s) (shown after the
comma). Note:

(1) After the two meiotic divisions, the diploid


number of chromosomes, 46, is reduced to the
haploid number, 23;
(2) four sperms form from one primary
spermatocyte, whereas only one secondary
oocyte results from maturation of a primary
oocyte;
(3) the cytoplasm is conserved during
oogenesis to form one large cell, the oocyte.
Polar bodies are small, non‐functional cells. They receive very little of the available cytoplasm and
degenerate soon after division. In this way the oocyte is able to retain its size but discard
chromosomal material to become a haploid cell ready for fertilisation. One polar body is formed with
meiosis I and two polar bodies are formed with meiosis II.

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