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folliculogenesis
Wednesday 10 November 2010
follicle development
Definition: In biology, folliculogenesis is the maturation of the ovarian follicle, a densely-packed shell of somatic cells that contains an immature oocyte.
Folliculogenesis describes the progression of a number of small primordial follicles into large preovulatory follicles that enter the menstrual cycle.
Contrary to male spermatogenesis, which can last indefinitely, folliculogenesis ends when the remaining follicles in the ovaries are incapable of responding to the hormonal cues that previously recruited some follicles to mature. This depletion in follicle supply signals the beginning of the menopause.
The primary role of the follicle is oocyte support.
Primordial follicles are indiscernible to the naked eye. However, these eventually develop into primary, secondary and tertiary vesicular follicles. Tertiary vesicular follicles (also called "mature vesicular follicles" or "ripe vesicular follicles") are sometimes called Graafian follicles (after Regnier de Graaf).
In humans, oocytes are established in the ovary before birth and may lie dormant awaiting initiation for up to 50 years.
From birth, the ovaries of the human female contain a number of immature, primordial follicles. These follicles each contain a similarly immature primary oocyte.
After puberty and commencing with the first menstruation, a clutch of follicles begins folliculogenesis, entering a growth pattern that will end in death or in ovulation (the process where the oocyte leaves the follicle). After rupturing, the follicle is turned into a corpus luteum.
During post-pubescent follicular development, and over the course of roughly a year, primordial follicles that have begun development undergo a series of critical changes in character, both histologically and hormonally.
Two-thirds of the way through this process, the follicles have transitioned to tertiary follicle (antral, follicles). At this stage in development, they become dependent on hormones emanating from the host body, causing a substantial increase in their growth rate.
With a little more than ten days until the end of the period of follicular development, most of the original group of follicles have died (a process known as atresia). The remaining cohort of follicles enter the menstrual cycle, competing with each other until only one follicle is left.
This remaining follicle, the late tertiary follicle (pre-ovulatory follicle), ruptures and discharges the oocyte (that has since grown into a secondary oocyte), ending folliculogenesis.
Folliculogenesis lasts for approximately 375 days. It coincides with thirteen menstrual cycles. The process begins continuously, meaning that at any time the ovary contains follicles in all stages of development, and ends when a mature oocyte departs from the preovulatory follicle in a process called ovulation.
The growing follicle passes through the following distinct stages that are defined by certain structural characteristics (unfamiliar terms will be defined in their respective sections):
In a larger perspective, the whole folliculogenesis, from primordial to preovulatory follicle, belongs to the stage of ootidogenesis of oogenesis.
Up until the preovulatory stage, the follicle contains a primary oocyte that is arrested in prophase of meiosis I. During the late preovulatory stage, the oocyte continues meiosis and becomes a secondary oocyte, arrested in metaphase II.
Development of oocytes in ovarian follicles - Oogenesis
In a larger perspective, the whole folliculogenesis from "primordial follicle" to "preovulatory follicle" is located in the stage of meiosis I of ootidogenesis in oogenesis.
The embryonic development doesn’t differ from the male one, but follows the common path before gametogenesis. Once gametogonia enter the gonadal ridge, however, they attempt to associate with these somatic cells. Development proceeds and the gametogonia turns into oogonia, which become fully surrounded by a layer of cells (pre-granulosa cells).
The Oogonia multiply by dividing mitotically; this proliferation ends when the oogonia enter meiosis. The amount of time that oogonia multiply by mitosis is not species specific. In the human fetus, cells undergoing mitosis are seen until the second and third trimester of pregnancy.
After beginning the meiotic process, the oogonia (now called primary oocytes) can no longer replicate. Therefore the total number of gametes is established at this time. Once the primary oocytes stop dividing the cells enter a prolonged ‘resting phase’. This ‘resting phase’ or dictyate stage can last anywhere up to fifty years in the human.
For several primary oocytes that undergoes meiosis, only one functional oocyte is produced. The other two or three cells produced are called polar bodies. Polar bodies have no function and eventually deteriorate.
The primary oocyte turns into a secondary oocyte in mature ovarian follicles. Unlike the sperm, the egg is arrested in the secondary stage of meiosis (Meiosis 2) until fertilization.
Upon fertilization by sperm, the secondary oocyte continues the second part of meiosis and becomes a zygote.
See also
ovarian follicle