Chapter: Medical Physiology: Reproductive and Hormonal Functions of the Male (and Function of the Pineal Gland)

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Semen

“Capacitation” of the Spermatozoa—Making It Possible for Them to Penetrate the Ovum, Acrosome Enzymes, the “Acrosome Reaction,” and Penetration of the Ovum.

Semen

Semen, which is ejaculated during the male sexual act, is composed of the fluid and sperm from the vas deferens (about 10 per cent of the total), fluid from the seminal vesicles (almost 60 per cent), fluid from the prostate gland (about 30 per cent), and small amounts from the mucous glands, especially the bulbourethral glands. Thus, the bulk of the semen is seminal vesicle fluid, which is the last to be ejaculated and serves to wash the sperm through the ejaculatory duct and urethra.

The average pH of the combined semen is about 7.5, the alkaline prostatic fluid having more than neutral-ized the mild acidity of the other portions of the semen. The prostatic fluid gives the semen a milky appearance, and fluid from the seminal vesicles and mucous glands gives the semen a mucoid consistency. Also, a clotting enzyme from the prostatic fluid causes the fibrinogen of the seminal vesicle fluid to form a weak fibrin coagulum that holds the semen in the deeper regions of the vagina where the uterine cervix lies. The coagulum then dissolves during the next 15 to 30 minutes because of lysis by fibrinolysin formed from the prostatic profibrinolysin. In the early minutes after ejaculation, the sperm remain relatively immo-bile, possibly because of the viscosity of the coagulum. As the coagulum dissolves, the sperm simultaneously become highly motile.

Although sperm can live for many weeks in the male genital ducts, once they are ejaculated in the semen, their maximal life span is only 24 to 48 hours at body temperature. At lowered temperatures, however, semen can be stored for several weeks, and when frozen at temperatures below -100°C, sperm have been preserved for years.

“Capacitation” of the Spermatozoa—Making It Possible for Them to Penetrate the Ovum

Although spermatozoa are said to be “mature” when they leave the epididymis, their activity is held in check by multiple inhibitory factors secreted by the genital duct epithelia. Therefore, when they are first expelled in the semen, they are unable to perform their duties in fertilizing the ovum. However, on coming in contact with the fluids of the female genital tract, multiple changes occur that activate the sperm for the final processes of fertilization. These collective changes are called capacitation of the spermatozoa. This normally requires from 1 to 10 hours. Some changes that are believed to occur are the following:

1. The uterine and fallopian tube fluids wash away the various inhibitory factors that suppress sperm activity in the male genital ducts.

 

2. While the spermatozoa remain in the fluid of the male genital ducts, they are continually exposed to many floating vesicles from the seminiferous tubules containing large amounts of cholesterol.This cholesterol is continually added to the cellular membrane covering the sperm acrosome, toughening this membrane and preventing release of its enzymes. After ejaculation, the sperm deposited in the vagina swim away from the cholesterol vesicles upward into the uterine cavity, and they gradually lose much of their other excess cholesterol over the next few hours. In so doing, the membrane at the head of the sperm (the acrosome) becomes much weaker.

 

3.               The membrane of the sperm also becomes much more permeable to calcium ions, so that calcium now enters the sperm in abundance and changes the activity of the flagellum, giving it a powerful whiplash motion in contrast to its previously weak undulating motion. In addition, the calcium ions cause changes in the cellular membrane that covers the leading edge of the acrosome, making it possible for the acrosome to release its enzymes rapidly and easily as the sperm penetrates the granulosa cell mass surrounding the ovum, and even more so as it attempts to penetrate the zona pellucida of the ovum itself.

 

Thus, multiple changes occur during the process of capacitation. Without these, the sperm cannot make its way to the interior of the ovum to cause fertilization.

Acrosome Enzymes, the “Acrosome Reaction,” and Penetration of the Ovum

Stored in the acrosome of the sperm are large quanti-ties of hyaluronidase and proteolytic enzymes. Hyaluronidase depolymerizes the hyaluronic acid polymers in the intercellular cement that hold the ovarian granulosa cells together. The proteolytic enzymes digest proteins in the structural elements of tissue cells that still adhere to the ovum.

When the ovum is expelled from the ovarian fol-licle into the fallopian tube, it still carries with it mul-tiple layers of granulosa cells. Before a sperm can fertilize the ovum, it must dissolute these granulosa cell layers, and then it must penetrate though the thick covering of the ovum itself, the zona pellucida. To achieve this, the stored enzymes in the acrosome begin to be released. It is believed that the hyaluronidase among these enzymes is especially important in open-ing pathways between the granulosa cells so that the sperm can reach the ovum.

When the sperm reaches the zona pellucida of the ovum, the anterior membrane of the sperm itself binds specifically with receptor proteins in the zona pellu-cida. Then, rapidly, the entire acrosome dissolves, and all the acrosomal enzymes are released. Within minutes, these enzymes open a penetrating pathway for passage of the sperm head through the zona pel-lucida to the inside of the ovum. Within another 30 minutes, the cell membranes of the sperm head and of the oocyte fuse with each other to form a single cell. At the same time, the genetic material of the sperm and the oocyte combine to form a completely new cell genome, containing equal numbers of chromosomes and genes from mother and father. This is the process of fertilization; then the embryo begins to develop.

Why Does Only One Sperm Enter the Oocyte? With as manysperm as there are, why does only one enter the oocyte? The reason is not entirely known, but within a few minutes after the first sperm penetrates the zona pellucida of the ovum, calcium ions diffuse inward through the oocyte membrane and cause multiple cortical granules to be released by exocytosis from the oocyte into the perivitelline space. These granules contain substances that permeate all portions of the zona pellucida and prevent binding of additional sperm, and they even cause any sperm that have already begun to bind to fall off. Thus, almost never does more than one sperm enter the oocyte during fertilization.


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