Karmina Santos September 19, 2016

4BIO3 Embryology Lec

Fertilization and Cleavage of C. elegans

The process of fertilization in C. elegans occurs within the spermatheca of the

adult hermaphrodite: Contractions of the oviduct force an oocyte into the spermatheca,
where the leading end of the oocyte appears to engulf a single sperm. Supernumerary
sperm carried into the uterus on the oocyte abruptly migrate back through the
constriction to the spermatheca so that every sperm can fertilize an oocyte. When
males mate with hermaphrodites they deposit their sperm in the region of the vulva and
these sperm migrate past the zygotes in the uterus to the spermatheca. When the male
sperm arrive at the spermatheca they preferentially fertilize the subsequent oocytes
even though hermaphrodite sperm are still present. The sperm are ameboid cells with a
specialized pseudopodial region that is extended from the cell while the sperm is
migrating. The sperm contain specialized membraneous vesicles that can fuse with the
plasma membrane. The newly fertilized egg exits its prophase arrest state and
completes meioses I and II, extruding two polar bodies at the future anterior end of the
embryo. Concurrently, a hard and impermeable eggshell forms around the embryo.

First, sperm entry appears to activate the oocyte to begin embryogenesis.

Unfertilized oocytes do not complete meiosis or become surrounded by an eggshell;
they undergo rounds of DNA replication, but in the absence of functional microtubule-
organizing centers (MTOCs), they do not undergo mitosis or cytokinesis.

Second, centrosomes are paternally inherited in C. elegans. The poles of the

oocyte meiotic spindle lack centrioles and are apparently incapable of functioning as
MTOCs for mitosis. The centriole-containing centrosome present in the sperm
duplicates after fertilization, and the resulting MTOCs nucleate the microtubules that
mediate pronuclear migration and compose the mitotic spindle
 Third, the site of sperm entry specifies the future posterior end of the embryo.
The sperm normally enters the end of the oocyte that first penetrates the spermatheca,
and this end becomes posterior. The oocyte nucleus normally resides at the opposite
end, and its polar body products usually mark the future anterior of the embryo The
unfertilized C. elegans oocyte has no predetermined axis. The sperm is thought to
specify the initial asymmetries in the embryo by directing cytoplasmic rearrangements
that cause determinants to become asymmetrically localized

Cleavage

The C. elegans demonstrates rotational holoblastic cleavage. It also exhibits a

series of asymmetrical and asynchronous divisions meaning its process of cell division
does not necessarily happen at the same time and may not be of equal shape. There is
the presence of P granules at the posterior end of the cell. The cells which have the P
granules are collectively called the P cells. The founder cells are called AB, MS, E, C,
D and P4.

In the one-cell stage the decision as to which end will become the anterior and
which the posterior will depend on the position of the sperm pronucleus. Then, the
Centriole with the sperm pronucleus initiates cytoplasmic movements that push the
male pronucleus to the posterior pole. A second anterior-posterior asymmetry is then
followed shortly after fertilization and it is the migration of the P-granules. The P-
granules are ribonucleoprotein complexes that probably function in specifying the germ
cells.

The two-cell stage then occurs after the pronuclei have fused. The zygote (P0)
divides into the Anterior AB cell and Posterior P1 cell. The P-granules are concentrated
at the P1 cell. In the four cell stage on the other hand, the AB cell then further divides
into the anterior (ABa) and posterior (ABp) cells. The ABa gives rise to neurons,
hypodermis and anterior pharynx cells. The ABp gives rise to neurons and hypodermal
cells only. P1 undergoes an asymmetric division that yields the EMS cell and P2 cell.
The ABp cell slides over the EMS cell which is result of division of P1 cell. ABp
then becomes the dorsal part. EMS would become the ventral part.
 In the 8 cell stage, the ABa cell further divides into ABal and ABar cells which
represent the left and right side respectively. The ABp cell divides into ABpr and ABpl
cells. EMS divides asymmetrically to produce E and MS. E is the precursor of all
intestinal tissue. MS is the major muscular precursor. P2 divides asymmetrically to
produce C and P3. C is a muscle and hypodermis precursor.

References:

Riddle, D. L., Blumenthal, T., & Meyer, B. J. (1997). C. elegans II (2nd ed.). Cold Spring
Harbor, New York: Cold Spring Harbor Laboratory Press.

Ward, S., Korman, G., & Carrel, J. (n.d.). Fertilization in C. elegans. Worm Breeders'
Gazette, 2(2).

http://www.wormbook.org/chapters/www_asymcelldiv/asymcelldiv.html

http://www.wormatlas.org/ver1/handbook/anatomyintro/anatomyintro.htm