Chua, Mark Ericson T.

4B3
C. elegans Fertilization and Cleavage
Fertilization
Fertilization involves the union of a sperm and an egg to form a diploid
zygote. In C. elegans, fertilization may occur in two ways. One way is for the
hermaphrodite to undergo self-fertilization. As oocytes are ovulated to the
uterus, they pass through the spermatheca where sperm is stored. The
passing oocyte is fertilized in the spermatheca by the stored sperm. The
second way fertilization occurs is by copulation of a male with a
hermaphrodite. Sperm is released from the cloaca of the male into the
vulva of the hermaphrodite. The cloaca of the male C. elegans is connected
to a modified rectum called a proctodeum. The proctodeum is connected to
both the intestine and the gonads. The proctodeum has two prong-like
structures called spicules, which is used to locate the vulval slit of the
female and to hold the vulva open during sperm transfer as well. Fertilized
eggs for both methods of fertilization undergo a series of cleavage before it
is eventually laid through the vulva at the 40-cell stage.
Primary oocytes are fertilized before the first meiotic division is
completed and maturation of the oocyte reinitiates in the presence of major
sperm protein (MSP). In the resulting zygote, the centriole and
microtubule-organizing center (MTOCs) come from the sperm while
the cytoplasm comes from the egg. The site of sperm entry on the egg
determines the future posterior end of the embryo. This is visible as a
smooth posterior cortical region while the rest of the egg becomes ruffled.
Cleavage
C. elegans displays rotational holoblastic cleavage. In addition,
cleavage of the zygote is asymmetrical and asynchronous, which means
that cleavage occurs at different times and results in cells of different sizes.
In C. elegans, complexes of ribonucleoprotein that probably function in
specifying the germ cell layer called P-granules can be found. P-granules
tend to concentrate on a single cell during each cleavage, which results into
a P-cell that contains P-granules, and a cell that contains little to no Pgranules called founder cells. Founder cells are named using capital letters
such as AB, MS, E, C and D, and they give rise to somatic cells instead of
germ cells.
One-Cell Stage
During the newly formed one-cell stage, the anterior and posterior
ends are determined by the position of the male pronucleus. The centriole
with the sperm pronucleus initiates cytoplasmic movements that push the
male pronucleus to the posterior of the cell. The male and female
pronuclei eventually fuse to form a zygote (P0). After fusion of the pronuclei,
the zygote formed can undergo cleavage resulting in two cells. The larger,

anterior cell is called the AB cell while the smaller and more posterior cell is
called the P1 cell. P-granules are highly concentrated in the P1 cell.

2-Cell Stage
Now at the 2-cell stage, the AB and P1 cells will divide and will result in
4 cells. The AB cell will divide into the more anterior ABa cell and the
more posterior ABp cell. The ABa will give rise to the future cells of the
neurons, hypodermis and anterior pharynx cells while ABp will give rise to
future neurons and hypodermal cells only. The P1 cell will divide into the
EMS cell and the P2 cell. Similar to the P1 cell, the P-granules are now
concentrated at the P2 cells. The ABp cell becomes the dorsal part of
the cell while the EMS becomes the ventral part.
4-Cell Stage
The 4 cells of the embryo will further divide. The ABa cell will divide
into the ABal and the ABar cells, which are the anterior left and right
cells, respectively. Similarly, the ABp cell will divide into the ABpl and
ABpr. The ABpl is the left cell at the dorsal region while ABpr is the right one.
The EMS cell divides into the E and MS cells. The E cell is the precursor
for all intestinal tissue of C. elegans, while the MS cell will give rise to the
muscles. P2 will divide into the C cell and the P3 cell. The P3 cell
contains the P3 granules. The C cell is the precursor cell for muscle cells and
as well as hypodermal cells. This division results in a total of 8 cells.
8-Cell Stage
In the 8-cell stage, the P3 cell will further undergo further
cleavage to give rise to the D cell and the P4 cell. The D cell is a
muscle precursor while the P4 will eventually give rise to the germ line. The
other cells will continue to divide into to give rise to their previously
mentioned organs.
Reference:
Altun,

Z.F.

and

Hall,

D.H.

2009.

Introduction.

In

WormAtlas.

doi:10.3908/wormatlas.1.1
Gilbert, S. (2000). Developmental Biology (6th ed.). Sunderland: Sinauer Associates
Gnczy, P. & Rose, L.S. (2005). Assymetric cell division and axis formation in the
embryo. Wormbook.
doi/10.1895/wormbook.1.30.1

Lints, R. and Hall, D.H. 2009. Male neuronal support cells, spicules. In

WormAtlas.

doi:10.3908/wormatlas.2.11
Riddle, D. L., Blumenthal, T., & Meyer, B. J. (1997). C. elegans II (2nd ed.). Cold
Spring Harbor, New York: Cold Spring Harbor Laboratory Press
Slack, J.M.W. (2013). Essential developmental biology (3rd ed.). Oxford: Blackwell
Publishing Ltd.
Ward, S., Korman, G., & Carrel, J. (n.d.). Fertilization in C. elegans. Worm Breeders'
Gazette, 2(2)