Ovarian Follicular dynamics in

domestic animals
• The Female gamete the oocyte originate and
develop from the ovarian follicles. The ovarian
follicles develop from the primordial germ
cells during fetal period and are called
primordial follicles.
• Follicle is the primary functional unit of ovary
that releases the oocyte and produces the
steroid hormones.
• Gestation
• Embryonic sac endoderm primordial germ cells
– germ cell cords migrate
Epithelial cells Gondal Ridge
•
• Condense around
• germ cells Mitosis
Meiosis

Arrested at prophase -I
Primordial follicles.
Fate of follicles and oocytes. 100 Days of Gestation
2.9 million
Total number of follicles
At birth 100,000
12 months 75,000
4-6 years 21,000
Aged cow 2,500

Follicles grow to tertiary characteristic and degenerate.

Maximum number of oocytes ovulated for a cow

10 cycles/year x 12 year = 120 oocytes ovulated/lifespan

It has been estimated that no more than about 300 oocytes are ever likely to
reach the ovulatory stage during the normal reproductive lifespan of the cow.
• Follicular growth can be classified into three
phases according to their developmental stage
and gonadotropin dependence : (1) follicular
growth through primordial, primary, and
secondary stages gonadotropin-independent
phase), (2) transition from preantral to early
antral stage (gonadotropin-responsive phase),
and (3) continual growth beyond the early
antral stage (gonadotropin-dependent phase),
which includes follicle recruitment, selection,
and ovulation.
• Ovaries of cattle contain two different pools of
follicles, the non-growing pool and the
growing pool. The non-growing pool contains
the primordial follicles, whereas the growing
pool contains the primary, secondary and
tertiary follicles. Entry of primordial follicles
into the growth phase occurs throughout the
reproductive life. The primordial follicles
continuously leave the arrested pool and
undergo the primordial to primary follicle
transition.
After the preovulatory LH surge in the early hours of estrus in the cow,
the follicle is known as the preovulatory follicle.
• The growth of follicles from the primordial to
ovulatory follicles is complex and requires a
couple of days to months.

• The sequence of events involves the growth of

primordial follicles into preantral and then antral
follicles (follicles with a central fluid filled cavity-
the antrum). This transformation involves the
proliferation of granulosa cells and reorganization
of cells forming the follicle.
• The growth of the primordial follicles is
resumed at puberty subsequent to which the
follicles grow in a cyclic fashion that is
repeated at defined intervals regularly
throughout the year in non seasonal breeding
species.
• During this transition the oocytes increase in size
and the surrounding squamous pre-granulosa
cells become cuboidal and proliferate to form a
layer of cuboidal cells around the growing oocyte.
The follicle is now called the primary follicle. The
mechanisms responsible for the initiation of
follicular growth during this phase are poorly
understood although some molecules such as
growth factors and gonadotrophins have been
discussed.
 Puberty
Gonadotropin-independent phase

Primordial follicle activated Primary and Sec follicle

Gonadotropin-dependent phase

• Preantral Antral follicles

Gonadotropin-dependent phase

• Early antral Follicle recruitment, selection,

and ovulation-Follicular waves
Mucopolysaccharide
layer Zona pellucida

Tertiary follicles also

called small antral
follicles
 Antral Follicle
Theca Internal

Granulosa
It is now known that the antral follicles
are selected to grow and this growth
occurs in a wave like fashion in most
domestic mammalian species. The
follicular growth waves lead to growth in
the size and fluid secretions within the
follicle, expression of estrus, and
ovulation with release of mature oocyte.
• Normally three to six follicles with a diameter
of 4 to 5 mm occur after recruitment of follicle
into a follicular wave.
• From the cohort of the growing follicles one
follicle is selected for continued growth and
becomes dominant. If luteolysis occurs during
the growth phase of the dominant follicles,
final maturation and ovulation occurs. If
luteolysis does not occur during the growing
and maintenance phase of follicles, the fate is
atresia
• Follicle recruitment
• The concept recruitment is used for the entrance
of follicles in the growing pool, but also for the
processes associated with the entrance of
follicles in a wave like growth. FSH appears to
be the key hormone for the initiation of the
follicular wave. The surges in FSH precede the
emergence of a wave. The FSH surges begin 2-
4 days before the detectable (ultrasound)
emergence of a follicular wave (follicles of 4
and 5 mm), peaked 1 or 2 days and started
declining nearly when the follicles of a wave
begin to diverge into a dominant follicle and
subordinate follicles (follicles 6 to 8 mm).
• Recruitment of the cohort containing the
future preovulatory follicle occurs during a
recruitment window which lasts 1, 2 or 3 days
in sheep, cattle or horses respectively. Only
gonadotrophin dependent healthy follicles
are recruited. The number of recruited
follicles growing in the cohort appears to be
highly variable between species. It ranges
from over 50 in pigs, to 5 to 10 in cattle and
1 to 4 in horses. All follicles of the cohort are
capable of ovulating, since selective ablation
of all except one will not postpone ovulation
in sheep and cattle.
• Selection
• Selection means that the number of
growing follicles is brought into line with the
species-specific ovulation number. After
recruitment fewer and fewer recruited
follicles continue in growth and one follicle
is selected to become dominant while the
remaining members of the recruited
follicles become static and eventually
undergo atresia via apoptosis.
• Dominance
• Follicles are functionally dominant (capable of
ovulating after luteal regression) while they are still
growing. Follicles acquired ovulatory capacity at
about 10 mm, corresponding to about 1 day after the
start of follicular deviation.
• Dominance negative feedback effects of products of
the dominant follicle on circulating FSH.
• Dominance progressive increases in the ability of
theca cells to produce androgen and granulosa cells
to aromatize androgen to estradiol.
• Dominant follicles grow to a much larger size (from
8.5 mm at the end of selection to 12-20 mm) in 3 to 4
days.
• Dominant follicle . Progesterone LH Ovulation
• Dominant follicle No follicular wave
Initial Follicular Growth
Small Antral
Follicle

50 - 60 days

Gonadotropin FSH
Independent

Recruitment
 Follicular Waves in Cattle
Progesterone

Atresia
Recruitment
Follicular Size

Dominance
Ovulation

Selection

FSH Sensitive Pool

Ovulation 9 16 21

Day After Ovulation

 Follicular Waves in Cattle
Progesterone
estradiol
Atresia inhibin
Follicular Size

Dominance FSH estradiol (-)

inhibin (-)

Selection FSH estradiol (-)

inhibin (-)

Recruitment FSH estradiol (-)

Ovulation 9 16 21

Day After Ovulation

 Three Follicular Waves in Cattle
Progesterone
Ovulation
Follicular Size

Dominance
FSH, E2
inhibin, LH
Selection
FSH, E2
inhibin, LH

Recruitment
FSH, E2

Ovulation 9 16 21

Day After Ovulation

 Two Follicular Waves

Dominance
Follicular Size

Atresia Ovulation

Selection

Recruitment

Ovulation 10 21

Day After Ovulation

 Species Variation in Follicular Waves
• Cattle - 2 or 3 / cycle
• Sheep - 4 or 5 / cycle Cl-progesterone

• Pigs - 1 / cycle Cl-progesterone, inhibin

• Horses - 1 / cycle
 Theca Interna
LH
Selection
Receptor
Dominant Follicle
cAMP Express LH Receptor
Cholesterol

PKA

Testosterone
LH
FSH
Receptor
Receptor

cAMP cAMP
Testosterone

PKA

Estradiol
Granulosa
 Preovulatory LH Surge
Cumulus Increased Blood
Expansion Flow to Ovary and
Protein Synthesis
Follicle
in and around Follicle

Progesterone Plasminogen Prostaglandin Vascular

Activator Synthesis Permeability
Collagenase (inactive) PGE and PGF
Plasmin Plasminogen
Oocyte Collagenase (active)
Separates Contraction
From Follicular Wall
of Smooth
Follicular Weakens
Muscle
Wall

Ovulation
 Follicular dynamics in buffaloes
Buffaloes evidence 1-, 2- or 3- wave cycles with
the 2-wave cycle being the most common pattern
of follicular growth Size of ovulatory follicle above
8.5mm

In some breeds one wave cycle is common for

example the Surti breed

During hot summer months there is variable degree of

ovarian follicular turnover in one or both ovaries
without expression of estrus signs and regression of
dominant follicles instead of their ovulation.
Follicular dynamics in mares
In the mare ovary there are approximately
40,000 primordial follicles and 100 growing follicles
Atresia (regression) of follicles is rare until
they reach 1 mm
The types of follicular waves that develop in mares are
major waves (characterized by dominant and subordinate
follicles) and minor waves (largest follicle does not attain
the diameter of a dominant follicle).
In some breeds (e.g. Quarter-Horses, Ponies), usually only
one major wave develops in late diestrus and culminates in the
estrous ovulation. In other breeds (e.g. Thoroughbreds), a
secondary major wave frequently develops in early diestrus,
and the dominant follicle may be anovulatory (more common),
or occasionally ovulatory (diestrous ovulation).

Minor follicular waves have been demonstrated in mares in

variable patterns. In all horse breeds, however, a major
ovulatory wave begins at mid-cycle and one follicle
(occasionally two, rarely three) becomes dominant and
ovulates.
• The ovulatory waves, as well as major
anovulatory waves and minor waves,
originate from the stimulation of an FSH
surge, which reaches a peak when the
largest follicle is about 13 mm.

• The physiologically selected dominant

follicle grows to a large diameter (≥ 28
mm) and then either regresses
(anovulatory major wave) or ovulates
(ovulatory wave).
• Maximum diameter of the dominant follicle is
smaller for major anovulatory waves than for
ovulatory waves (e.g., means of 37 and 46
mm

• A paradox in mares is the occurrence of

ovulation from the dominant follicle of what
was expected to be an anovulatory wave.
These ovulations have been called
secondary or diestrus ovulations and occur
during high progesterone concentrations.
 Follicular dynamics in sheep
• In sheep follicles ≥5mm in diameter would
exhibit a wave-like pattern while follicles
smaller than 4mm would grow randomly. Both
patterns of growth are classified as primary
and secondary follicular waves, respectively.

• Two to 3 follicular waves per cycle are

common in sheep, and 3 or 4 follicular waves
per cycle are most common in goats
• During estrous cycles, the dominant and
largest subordinate follicles reach maximum
diameters of 5 to 7 mm and 3 to 5 mm
respectively in sheep. In most cases the
ovulatory follicles develop from the cohort of
follicles from the last follicular wave. However,
the ovulatory follicles can also derive from the
penultimate follicular wave with the result
that follicles can ovulate both from the last
and the second last follicular waves of the
cycle.
 Follicular dynamics in goats

• In goats on any one day of the estrous cycle

there are 5 to 10 follicles > 3 mm in diameter
in the ovaries and follicles ovulate at or
between 6 and 9 mm in diameter. As in sheep
double ovulations can derive from follicles of
the last and the second last wave.
 THANKS
Kindly share the video and
subscribe to my channel G N
Purohit if you like them