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Mendoza, Princess Grace N.

Fertilization in Musmusculus
Fertilization is the union of the male and females gametes. The mouse has been
frequently used to study mammalian fertilization due to (1) thetremendous wealth
of knowledge available about mouse developmental and reproductive biology, (2)
the relatively low cost and ease with which mice can be obtained, housed, and
handled, (3) the well-established protocols available for reliably obtaining and
culturing relatively large numbers of mouse gametes, (4) the well-established
protocols available for carrying out fertilization with mice, both in vivo and in vitro,
and (5) the firm belief that fertilization in mice is an appropriate model for
understanding many aspects of human fertilization. These factors made the mouse
as a major contributor to the understanding of the mammalian fertilization.
There are three mouse egg components directly involved in the mouse fertilization
process; (1) the zona pellucida (ZP); (2) the plasma membrane; and (3) the cortical
granules (CG). The mouse egg zona pellucida is about 7 µm thick and contains 3-4
ng of protein. It is composed of three relatively acidic glycoproteins, called ZP1,
ZP2, and ZP3. All three glycoproteins are synthesized and secreted by oocytes
during their 2 to 3-week growth phase, the period when the ZP first appears. The ZP
increases in thickness as the oocyte increases in diameter. The ZP consists of long
filaments that are composed of ZP2 and ZP3 (located every 15 nm or so) and are
cross-linked by ZP1 to form a very porous, three-dimensional matrix. The ZP
glycoproteins interact with one another via noncovalent bonds.The mouse egg
plasma membrane is extensive since it is associated with thousands of long (0. 31 µm) microvilli. Only the surface ofthe egg where emission of the first polar body
occurred is free of microvilli. During oocyte growth the number and length of
microvilli increase markedly as the oocyte increases in diameter. Furthermore,
fusion between gametes nearly always involves egg microvillar membranes,
probably because they have a low radius of curvature (as compared with
amicrovillar plasma membrane) that permits maximum apposition of sperm and
egg. In general, the egg plasma membrane does not appear to confer species
specificity on gamete fusion, and there does not appear to be a specific site on the
egg surface where gamete fusion must occur. The mouse egg contains about 4500
cortical granules (-30 CG per 100 µm plasma membrane) rangingfrom 200 nm to
600 nm in diameter and located within about 2 µm of the plasma membrane. The
CG are heterogeneous in appearance and, in some cases, appear to be attached to
the cytoplasmic face of the egg plasma membrane. Like the ZP, CG first appear in
the oocyte during its growth phase, as a product ofthe Golgi, and increase in
number as the oocyte increases in diameter.
There are several steps that take place in mouse fertilization: (1) attachment of the
sperm to the unfertilized egg extracellular coat or zona pellucida, (2) binding of the
sperm to the zona pellucida via plasma membrane overlying the anterior region of
the sperm head, (3) bound sperm undergo acrosome reaction (AR) which is a form
of signal-transduced exocytosis, (4) acrosome-reacted sperm penetrate the zona
pellucida, reach the perivitelline space between the ZP and egg plasma membrane,
and then fuse with the egg plasma membrane to form a zygote. Fertilization by a

single sperm induces eggs to undergo the cortical reaction (CR), which activates the
zona reaction (ZR). This involves the fusion of CG membrane with egg plasma
membrane and the hardening of the zona pellucida, respectively.