Developmental Cell Biology 220
Lecture Outline for
February 12, 1999
Rearrangement of Egg Cytoplasm
& Cleavage
Rearrangement of egg cytoplasm
-
Morphogenetic determinants are separated into different areas of the newl
fertilized egg
-
Tunicate egg (Styela partita) - see Figure 4.32
-
After fertilization yellow lipid-containing cytoplasm migrates ventrally.
-
A yellow crescent is formed in the area which will later become larval
muscle cells.
-
These movements depend on Ca+2 released after fertilization
and construction of microtubules.
-
Frog eggs
-
After fertlization the surface layers of the animal hemisphere rotate 30o
toward the point of sperm entry.
-
The newly exposed surface area will later invaginate during gastrulation.
-
Polymerization of microtubules is involved.
Cleavage
-
The site of the first cleavage furrow is determined by the point of sperm
entry and the subsequent cytoplasmic movements of the newly fertilized
egg.
-
In general, the embryonic volume does not change from that of the newly
fertilized egg until after the first12 cell divisions (producing ~ 4000
cells).
-
Patterns of cleavage (see Table 5.1)
-
Are determined by the amount and distribution of yolk protein
-
Cytoplasmic factors influence the angle and timing of the formation of
the mitotic spindle.
-
Based on whether the daughter cells are completely separated or not.
-
holoblastic - formation of complete and separate cells
-
meroblastic - incomplete separation of cells
-
Based on position/amount of yolk
-
Isolecithal
-
Mesolecithal
-
Telolecithal
-
Centrolecithal
-
Based on the position of the mitotic spindle and the daughter cells
-
Radial
-
Spiral
-
Bilateral
-
Rotational
-
Discoidal
-
Superficial
-
Sea urchin eggs (isolecithal) have radial, holoblastic cleavage.
(Fig 5.3)
-
Formation of mesomeres, macromeres, micromeres in the 16 cell embryo.
-
More layers are formed in the next two rounds of cell division.
-
The blastula starts with 128 cells and tight junctions between the cells
-
blastocoel formation involves accumulation of proteins within the blastocoel,
followed by accumulation of water (by osmosis)
-
blastocoel formation also involves adhesive connections to the hyaline
layer.
-
Most amphibian eggs (mesolecithal) are examples of radial, holoblastic
cleavage.
-
Because of the concentration of yolk in the vegetal hemisphere the formation
of the cleavage furrow in the yolk-containing ventral portion is slower
and lags behind that in the animal hemisphere.
-
Macromeres (large yolk-containing cells) and micromeres (animal pole cells)
form.
-
E-cadherin is important in adhesion between blastula cells.
-
Experiment: Injecting antisense oligonucleotides complementary
to E-cadherin mRNA prevents Xenopus embryos from forming a blastocoel.
-
Eggs of the snail (Lymnaea peregra) are examples of spiral, holoblastic
cleavage.
-
Cleavage occurs at oblique angles with daughter cells either moving to
the right or to the left.
-
In snails with a shell coil that opened to the right, after the second
cell division (Figure 5.14) the micromeres moved to the right; in snails
whose coil opened to the left, after the second cell division the micromeres
moved to the left.
-
The direction of daughter cell movement is controlled by the D gene of
the mother (cytoplasmic determinants).
-
DD or Dd mother, all offspring had coils which opened on the right.
-
dd mother, all offspring had coils which opened on the left.
-
Eggs of the tunicate (Styela partita) have bilateral holoblastic
cleavage; Fig 5.17
-
During cleavage, cytoplasm that will give rise to different parts of the
embryo is segregated into different blastomeres.
-
At the 8 cell stage:
-
Clear cytoplasm = ectoderm
-
Yellow cytoplasm = mesoderm
-
Dark gray cytoplasm = endoderm
-
Light gray cytoplasm = neural tube and notochord.