At birth, the human ovary contains approximately 2 million ovarian follicles, which consist of germ cells, or oocytes, surrounded by a cluster of endocrine cells that provide an isolated and protected environment for the oocyte. In the first stage of follicle development, occurring mostly before puberty, and taking anywhere from 13 to 50 years, the cells surrounding the oocyte (the granulosa cells) divide and form several layers around the oocyte, forming the so-called secondary follicle. Subsequent to puberty, these secondary follicles form a reserve pool from which follicles are recruited to begin the second stage of development. In this second stage, follicles increase to a final size of up to 20 mm before they rupture and release the oocyte to be fertilized. The release of the oocyte is called ovulation. Although many follicles begin this second developmental stage, few reach full maturity and ovulate, as the rest atrophy and die. In fact, the number of oocytes reaching full maturity is carefully controlled, so that litter sizes are generally restricted to within a relatively narrow range, and different species have different typical litter sizes. For example, to quote some interesting, if not particularly useful, facts from Asdell (1946), both the dugong and llama have a typical litter size of 1, the crestless Himalayan porcupine typically gives birth to two offspring, while the dingo produces, on average, 3. Different breeds of pigs have litter sizes ranging from 6 to 11.
There must therefore be a complex process that, despite the continuous recruitment of secondary follicles into the second developmental stage, allows precise regulation of the number remaining at ovulation. Further, the temporal periodicity of ovulation is tightly controlled, with ovulation occurring at regularly spaced time intervals.
In addition to questions related to the nature of the control of ovulation, there is the question of efficiency. It appears inefficient to regulate the final number of mature follicles by initiating the growth of many and killing off most of them. One might speculate that it would be more reasonable to initiate growth in only the required number and ensure that they all progress through to ovulation.
Normal ovulation involves growth in both ovaries. However, since removal of one ovary does not change the total number of eggs released during ovulation, the control mechanism is not a local one, but a global one, known to operate through the circulatory system. Maturation of follicles is stimulated by gonadotropin, which is released from the pituitary gland. Gonadotropin consists of two different hormones called follicle-stimulating hormone (FSH) and luteinizing hormone (LH). However, follicles themselves secrete estradiol, which stimulates the production of gonadotropin, forming a feedback control loop for the control of follicle maturation (Fig. 19.1).
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