Fertilization

After ovulation, the unfertilized egg is arrested in prophase of meiosis II and

contains one polar body left over from meiosis I. Fertilization is a process of several
events and typically takes place in the ampullated portion of the uterine tube:
Capacitation
Changes take place in the glycoprotein coat of sperm as they travel up the
female reproductive tract. Capacitation is associated with removal of adherent
seminal plasma proteins, reorganization of plasma membrane lipids and proteins.
These changes are absolutely essential for fertilization. Sperm that have undergone
capacitation are said to become hyperactivated, and among other things, display
hyperactivated motility. Most importantly however, capacitation appears to
destabilize the sperm's membrane to prepare it for the acrosome reaction. Thus, to
perform successful in vitro fertilization you must add some tissue extracted from the
female reproductive tract in addition to the sperm and egg extracted from the
parents.
Approximation
Only a tiny fraction of sperm actually reaches the ampulla of the uterine tube
to be near the egg.
Penetration of Corona Radiata
The sperm uses both chemical and physical means to penetrate the eggs
corona radiata:

The action of membrane-bound enzyme hyaluronidase on its coat, and

Swimming motion of its flagellum.

Penetration of Zona Pellucida
The acellular zona pellucida that surrounds the egg at ovulation and remains
in place until implantation has two major functions in the fertilization process:
- The zona pellucida contains ligands for sperm
- The zona pellucida undergoes the zona reaction in which the zona
becomes impervious to other sperm once the fertilizing sperm penetrates,
and thus it provides a bar to polyploidy.
Once inside the corona radiata, the sperm binds to the species-specific ZP3
receptor on the eggs glycoprotein coat. Binding to the zona pellucida is required to
permit a component of the zona to induce the acrosomal reaction. This triggers the
acrosomal reaction, or the release of enzymes stored in the sperms acrosome (e.g.
acrosin - a trypsin-like proteinase). These enzymes help the sperm drill through
the zona pellucida along with movement of the head and tail of the sperm.
Once the sperm has penetrated the outer layers it fuses with the plasma
membrane of the egg and releases its contents inside. The head and the tail of the

sperm degrade, so that all mitochondria in the embryo (and all mitochondrial DNA)
come from the mother.

Cortical Reaction
Entry of a sperm into the egg triggers changes that prevent polyspermy
(fertilization of an egg by more than one sperm). These changes are known as the
cortical reaction. In cortical reaction, release of materials from the cortical granules,
lysosome-like organelles that are found just below the egg surface contains
materials including various hydrolytic enzymes. Changes brought about by these
enzymes lead to the hardening of the extracellular layer by cross-linking of
structural proteins, and inactivation of ligands for sperm receptors.
Cortical Reaction
Phase
Fast block

Slow block

Description
Electrical depolarization of the eggs surface
works for a short time to repel other sperm
electrostatically with release of calcium ion
from Calmodulin.
A wave of Ca++ ions released from the point of
sperm entry spreads through the egg. This
causes cortical granules in the egg to release
their contents.
Proteases in the cortical
granules reach the outside of the egg and
harden the zona to prevent sperm penetration.
Other enzymes in the granules break down the
ZP3 receptors in the zona pellucida and also
further harden the coat.

Fusion of Pronuclei
DNA in the male pronucleus is packed very tightly with protamines to make it
compact enough to fit inside a sperm. These protamines are replaced by histones
inside the egg, unpacking the DNA. The chromatin material of the sperm head
decondenses, and the male pronucleus is formed. The male and the female
pronuclei migrate toward each other, and as they move into close proximity the
limiting membranes break down, and a spindle is formed on which the
chromosomes become arranged. Thus, the stage is set for the first cell division. The
fertilized egg is now known as the zygote (together).
It is vital to not only understand how normal fertilization takes place but also
in the occurrence of abnormal events that can interfere with pregnancy. It is
worthwhile, therefore, to consider the failures that occur in association with in vivo
fertilization. By using sensitive pregnancy tests, it has been suggested that the total
rate of pregnancy loss after implantation is approximately 30%. When the loss of

fertilized oocytes before implantation is included, approximately 46% of all

pregnancies end before the pregnancy is clinically perceived.