Histology by Charlotte L. Ownby, PhD.  Links to beginning of Histology web site.

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  Return to Oogenesis  Link to Ovaries Link to Oviducts (uterine tubes) Link to Interaction during estrous cycle Link to Vagina Link to Review Materials Proceed to Follicle Development

Oogenesis: Development of Follicles in the Ovary

Oogenesis occurs in specialized structures in the cortex of the ovary called ovarian follicles.  
Click to see larger images with labels Ovarian follicles are composed of: 
  1. the developing gamete
  2. associated follicular cells

The two images at the left show the typical appearance of follicles in the cortex of the ovary.

Click the images to see them larger and labeled.

There are three stages in the development of follicles: 
  1. pre-ovulation
  2. ovulation
  3. post-ovulation.

On this page we will consider pre-ovulation only.

The pars distalis of the pituitary gland (hypophysis) controls the process of follicular development and maturation by the secretion of: 

  1. gonadotropic hormones FSH (follicle stimulating hormone) 
  2. LH (luteinizing hormone)
  3. prolactin (in some species).

The hormonal interactions will be explained in the discussion of the ovarian cycle.

Pre-ovulation: Development of Follicles in the Ovary.   
Click each image below to view the histological image.

Notice the flattened follicluar cells surrounding the oocyte.   The primordial (quiescent) follicle consists of a primary oocyte and a single layer of flattened follicular cells.  As the follicle develops, alterations occur in the primary oocyte and the surrounding follicular cells.  The primary oocyte produces yolk granules and the follicular cells change from flattened to cuboidal or columnar. 
Notice the single layer of cuboidal cells The primary follicle consists of a primary oocyte with a single layer of cuboidal/columnar follicular cells.  As development proceeds, the number of follicular cells increases by mitosis forming several layers around the primary oocyte. As these cells enlarge they release steroid hormones called estrogens of which estradiol is the dominant one prior to ovulation.  During each cycle, a few primary follicles will continue to develop into secondary follicles.
Notice more than one layer of granulosa cells. The secondary follicle consists of several layers of cuboidal/columnar follicular cells, now collectively called the membrana granulosa which begin to secrete follicular fluid.   A thick, amorphous layer, the zona pellucida, forms between the primary oocyte and the membrana granulosa. Previously undifferentiated stromal cells now develop into two distinct layers around the developing follicle: the theca interna and the theca externa (see below). Cells in the theca interna are large, rounded and epithelial-like; cells in the theca externa are smaller, fibroblasts. Both layers of theca cells are separated from the membrana granulosa cells of the follicle by a basement membrane. As the follicular fluid secreted by the membrana graulosa cells accumulates, small pockets of fluid between granulosa cells begin to appear. Usually in human females only one secondary follicle will continue to develop.
The secondary-vesicular follicle (below) is characterized by the presence of pockets of follicular fluid within the membrana granulosa. As the follicle continues to develop, the separate pockets fuse to form one large pocket of fluid called the follicular antrum.   During this development of the follicular antrum, the oocyte is still a primary oocyte, arrested in prophase of Meiosis I. It is still surrounded by granulosa cells which are contiguous with the membrana granulosa present around the periphery of the growing follicle. 
Notice the pockets of follicular fluid within the membrana granulosa Two regions of cells can be identified in the layer of granulosa cells surrounding the oocyte: 
  1. the corona radiata contains granulosa cells which remain attached to the oocyte after ovulation and are in close contact with the oocyte through cytoplasmic processes which pass through the zona pellucida and contact microvilli of the oocyte; 
  2. the cumulus oophorus contains granulosa cells which surround the oocyte and are continuous with the displaced cells of the membrana granulosa but remains in the ovary after ovulation. 

The other granulosa cells form a layer around the periphery of the follicle and are separated from the theca interna cells by a distinct basement membrane.

Notice the antrum of follicular fluid   

The mature follicle (left), sometimes called the pre-ovulatory follicle, has all of the components of the secondary-vesicular follicle but is much larger and contains one single large antrum of follicular fluid. These follicles are very large and usually extend from the deepest parts of the cortex and protrude from the surface of the ovary. In some species just before ovulation, the primary oocyte in the mature follicle completes meiosis I producing a secondary oocyte and a polar body. 

Click here to see follicle development

Click here

Click here to see follicle development in the ovary

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Pre-ovulation: Development of Theca Cells in the Ovary.   

As the oocyte and follicular (granulosa) cells are growing and developing in the ovary, the stromal cells differentiate and develop into the theca interna and theca externa cells.  As a follicle goes from a primary to a secondary follicle, the stromal cells immediately surrounding the follicle differentiate into the theca folliculi.  The cells closest to the follicle become the theca interna cells,  round, foamy cells that secrete androgens, including testosterone.  These two “male” hormones are converted by the granulosa cells to estrogens. The stomal cells farther away from the developing follicle become the theca externa cells, fibroblast-like cells arranged around the follicle outside the theca interna cells.

 Click the images on the right to see them enlarged and labeled. 

Click to see larger images with labels

 Quick Checks:
1.  Development of follicles in the ovary involves which of the following:
an increase in the number of oocytes
a decrease in the number of oocytes and follicular cells
an increase in the number of follicular and granulosa cells

2. Which of the following statements is True:
Estrogen is made and secreted by primordial follicles.
Estrogen is made and secreted by theca externa cells.
Estrogen is secreted by the primary oocyte.
Estrogen is secreted by granulosa cells.

 3.  The development of follicles in the ovary is under the control of which gland and which hormones:
pituitary gland; FSH and LH
hypothalamus; oxytocin and vasopressin
pineal gland; prolactin

Return to Oogenesis      Proceed to Post-Ovulation

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This page was last updated 10/19/07 09:53:56 AM