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Fertilization
The process by which the sperm initiates and participates, with the egg, in the
development of the embryo
What are the factors that affect the sperm’s potential for
successfully fertilizing the egg?
1. Capacitation - occurs in the female’s vagina. Vaginal secretions cause a molecular
change in the sperm plasmalemma (removal of decapacitating factor - semen proteins,
results in increased membrane fluidity,). Takes 4-5 hr in humans, 1 hr in mice, 6 hr in
rabbits.
http://www.erin.utoronto.ca/~w3bio380/l
ecture/Lect07/L7.htm
Effects of Capacitation on
Galactosyltransferase is a protein in the Sperm
sperm plasmalemma that MAY act as the
receptor that binds to the zona pellucida • Increased rate of metabolism
and initiates the acrosome reaction.
• Flagellum beats more rapidly; Result:
Sperm are more motile
• Changes in sperm plasmalemma proteins
allow sperm-egg binding and occurrence of
the acrosome reaction
• Pro-Acrosin (inactive) is converted to
acrosin (active)
• Sperm become capable of chemotaxis
http://www.erin.utoronto.ca/~w3bio380/l
ecture/Lect07/L7.htm
Factors that affect the sperm’s potential for successfully
fertilizing the egg? (cont.)
2. Dilution
Marine invertebrates - Free spawning of sperm into surrounding ocean water.
Mammals - Dilution: 40 - 400 million spermatozoa in vagina, only a few hundred to a
thousand reach the upper oviduct.
http://www.erin.utoronto.ca/~w3bio3
80/lecture/Lect07/L7.htm
Factors that affect the sperm’s potential for successfully
fertilizing the egg? (cont.)
4. Structures surrounding the egg - barriers
• Follicle cells
• Zona pellucida (vitelline membrane in non-
mammals)
• Oolemma (plasmalemma of egg)
http://www.talbotcentral.ucr.edu/mammalianfert.htm
http://arbl.cvmbs.colostate.edu/hbooks/pathphys/reprod/fert/gxport.html
6
Acrosome Reaction
How does the sperm get through the barriers surrounding
the egg?
1. Used to be thought that the acrosome enzymes did it all.
2. More recent data
http://arbl.cvmbs.colostate.edu/hbooks/pathphys/reprod/fert/gxport.html
Getting through the zona pellucida
What happens when the sperm gets to the zona pellucida?
1. Attachment - loose association
2. Binding - strong attachment
3. Acrosome reaction - release of enzymes
4. Penetration of the zona pellucida by the sperm
http://www.talbotcentral.ucr.edu/mammalianfert.htm
How is this accomplished?
Wassermann and co-workers (1980, 1985, 1987, 1988)
Found that zona pellucida is composed of 3 glycoproteins
ZP1, ZP2, ZP3
Repeating subunits of ZP2 and ZP3 form filaments that are bound
together by ZP1
1. Wassermann et al. found that when the sperm
binds to ZP3 it causes a change in Ca+2 and Na+
flux across the sperm plasmalemma that results
in the acrosome reaction. (ZP1 and ZP2 will not
cause this to happen).
2. The sperm actually binds to an o-linked
oligosaccharide that is part of ZP3 - (a
carbohydrate component of the glycoprotein)
3. The identity of the receptor in the sperm
plasmalemma that binds to this oligosaccharide
is, as yet, uncertain. It has been suggested that
a part of PH-20 is the receptor; however, more
recent work indicates this may not be the case.
When the acrosome reaction occurs, a number
of proteolytic enzymes are exposed or released.
One or more of these enzymes is responsible for
digesting the hole through the zona pellucida
through which the sperm enters the perivitelline
space.
Following fusion of the sperm with the egg,
http://www.molbiolcell.org/cgi/content/full/9/7/1609/F6
In mammals:
1. The male nucleus enters the egg cytoplasm
and becomes the male pronucleus.
2. As a result of the sperm fusing with the egg
plasmalemma, the oocyte nucleus, which is at
metaphase of the second meiotic division,
completes that division giving rise to another
polar body.
3. Following the second meiotic division, what is
now the nucleus of the ovum becomes the
female pronucleus. Male and female pronuclei in a
fertilized hampster egg
4. The haploid male and female pronuclei move
toward one and other, meet, and fuse to form
the diploid nucleus of the zygote.
5. The zygote will now proceed to undergo cleavage.
http://www.talbotcentral.ucr.edu/mammalianfert.htm Cleavage
http://arbl.cvmbs.colostate.edu/hbooks/pathphys/reprod/fert/cleavage.html
Blocks to polyspermy
1. Dilution
2. Fertilizin like secretions cause agglutination of sperm in some
invertebrate species (Does not occur in mammals)
3. “Hardening” of vitelline membrane or zona pellucida that results from
cortical granule rupture - - slow block to polyspermy, takes 1 - 5 min
http://www.molbiolcell.org/cgi/content/full/9/7/1609
http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=dbio.figgrp.1397
Blocks to polyspermy
4. Depolarization of the oolemma - membrane potential changes from
negative to positive, sperm cannot fuse with positively charged oolemma -
fast block to polyspermy, takes 1 - 2 seconds. Caused by influx of sodium
ions.
Thus, only
1) dilution of the sperm and
2) hardening of the zona pellucida due to the
contents of the cortical granules
act to prevent polyspermy in most mammals,
including humans.
Egg Activation
A series of morphological, physiological and molecular changes that
occur in the egg in response to fusion of the sperm with the egg.
What does egg activation accomplish?
1. Frees egg from constraints that kept it from developing prior to
fertilization.
2. Prepares the egg for development.
http://www.molbiolcell.org/cgi/content/full/9/7/1609
“The fertilization transient can be seen at the
very start of the animation. Later there is a
Sea Urchin rise in calcium starting at the point of sperm
entry which correlates with the start of the
movement of the female pronucleus to the
center of the egg and then a further general
rise in calcium which correlates with the
fusion of the two pronuclei.”
http://petrus.ncl.ac.uk/urchins/fertilizationb.html
The seven events that characterize egg activation following
fusion of the sperm with the egg:
3. In many species, an influx of Na+ (sodium) into the egg cytoplasm that
causes a change in membrane potential - fast block to polyspermy.
4. In many species a reorganization of the egg cytoplasm.
5. In most cases, completion of meiosis by the egg.
6. An efflux of H+ (hydrogen) ions causing an increase in cytoplasmic pH -
this activates previously inhibited synthetic pathways.
7. Increase in metabolism - zygote gears up for development.