Gastrulation

"It is not birth,

marriage, or death,
but gastrulation,
which is truly the
most important time
in your life."
Lewis Wolpert (1986)
–series of highly
coordinated cell
movements to
reorganize the simple
blastula into a
complex embryo with
3-D body plan of 3
Physiological changes
during gastrulation
●
rearrangement of cells
by morphogenetic
movements
●
rhythm of cellular
divisions is slowed
●
oxidation is intensified
●
nuclei become more
active in controlling
activities of the cells
●
synthesis of proteins
not present in the egg
cytoplasm
What more do we need
to know?
●
types of cell movements
●
forces that direct cell
movement
●
fate maps and
gastrulation in model
Types of cell
movements
(individual or cell
sheet/layer)
1. Ingression,
egression
●
migration of individual
cells from the surface
layer into the interior or
out towards the surface
Sea urchin
2. Intercalation
●
movement of individual
cells into a cell layer or
in between two of its
neighbors
3. Condensation
●
dispersed cells
congregate to form a
cell sheet
Dispersal
●
cell sheet breaks up to
form dispersed
mesenchyme cells
4. Invagination
●
folding in of a sheet of
cells to form an
indentation
Evaginatio
● n
folding out of a sheet of
cells to form a protrusion
5. Involution
●
inward turning of an
expanding sheet of cells
– cells continue to
spread over the
internal surface of the
sheet and create a
6. Epiboly
●
movement of sheet of
cells that spread as a
unit to enclose the
deeper layers of the
embryo
7. Delamination
●
splitting of one layer of
cells into two parallel
layers
8. Convergent
extension
●
elongation of cell layer
in one dimension and
shortening in another
dimension
●
movement of cells
during gastrulation is
irreversible
= each part remains in
the position into
which it was brought
by the preceding
What drives
cell
movement?
Dictatorship or
democracy?
1. Differential cell
affinity
●
cells sort out according
to their cell type
– strength of adhesion
●
to attain the most
thermodynamically
stable pattern
Determinants of
strength of adhesion
☻ types of cell adhesion
molecules
☻ amount of cell
adhesion molecules
a. Cadherins
☻calcium-dependent
adhesion molecules
– establish and

maintain intercellular
connections
= cadherin-cadherin
Cadherins in
vertebrates
☻E-cadherin
– epithelial cadherin

(uvomorulin or L-CAM)
expressed in early
embryo but later
restricted to epithelia
☻P-cadherin
– placental cadherin

● expressed in the

trophoblast cells and

uterine wall cells
☻N-cadherin
– neural cadherin

● expressed in

mesodermal cells as
they lose their E-
cadherins
☻EP-cadherin
– maintains adhesion

between blastomeres
of Xenopus larva,
required for normal
cell movement
☻Protocadherins
– lack connections to

cytoskeleton through
catenins
– separates nochord

from mesoderm
b. Integrins
☻integral proteins that
bind cells to extracelluar
matrix
– enable the cell to

endure pulling forces

without being ripped
2. Cell contraction
●
apical constriction
– basolateral expansion
of cells -> change in
cell shape -> bending
of cell sheet
– actomyosin driven
●
basal protrusions
– filipodia or
lamellipodia formation
towards direction of
movement + traction
at cell base
– actin driven
3. Morphogenetic
gradients
●
chemotaxis of cells
which direct cell
movements