Early Development in Sea Urchin
Early Development in Sea Urchin
SEA URCHINS
Sea Urchin Cleavage
• Cleavage in Sea Urchins is holoblastic
radial.
• the first and second cleavages are both
meridional and are perpendicular to each
other
• The third cleavage is equatorial,
perpendicular to the first two cleavage
planes, and separates the animal and
vegetal hemispheres from each other.
• The fourth cleavage, however, is very
different from the first three. The four
cells of the animal tier divide
meridionally into eight blastomeres, each
with the same volume.
• These eight cells are called mesomeres.
The vegetal tier, however, undergoes an
unequal equatorial cleavage. to produce
four large cells—the macromeres—and
four smaller micromeres at the vegetal
pole.
Cleavage to Blastula
• At later blastula, in which the cells form a hollow
sphere surrounding central cavity, or blastocoel is
called mature blastula . From here on, the pattern of
divisions becomes less regular.
• After entering into mature blastula the cell cleavage
become invariant.
• By this time, the fates of the cells have become
specified and each cell becomes ciliated on the region
of the cell membrane farthest from the blastocoel.
• This ciliated blastula begins to rotate within the
fertilization envelope. Soon afterward, differences are
seen in the cells.
• The cells at the vegetal pole of the blastula begin to
thicken, forming a vegetal plate.
• The cells of the animal hemisphere synthesize and
secrete a hatching enzyme that digests the
fertilization envelope. The embryo is now a free-
swimming hatched blastula.
Fate maps and the determination of sea
urchin blastomeres
• A fate map of the 60-cell sea urchin embryo is shown
in Figure. The animal half of the embryo consistently
gives rise to the ectoderm—the larval skin and its
neurons.
• The veg1 layer produces cells that can enter into
either the ectodermal or the endodermal organs.
• The veg2 layer gives rise to cells that can populate
three different structures—the endoderm, the coelom
(internal mesodermal body wall), and the non-
skeletogenic mesenchyme (sometimes called
secondary mesenchyme), which generates pigment
cells, immunocytes, and muscle cells.
• The first tier of micromeres (the large micromeres)
produces the skeletogenic mesenchyme (also called
primary mesenchyme), which forms the larval
skeleton.
• The second-tier micromeres (i.e., the small
micromeres) play no role in embryonic development
but appear to contribute cells to the larval coelom
Fate maps and the determination of sea
urchin blastomeres
• Although the early blastomeres have
consistent fates in the larva, most of these fates
are achieved by conditional specification.
• The only cells whose fates are determined
autonomously are the skeletogenic
micromeres.
• The micromeres appear to produce a signal
that tells the cells adjacent to them to become
endoderm and induces them to invaginate into
the embryo.
Axis specification Dorsal
Hbox-12 P38-MAPK
Nodal
Nodal
• In the sea urchin blastula, the cell fates line up along the animal-vegetal SoxB1 P38-MAPK
axis established in the egg cytoplasm prior to fertilization.
• The animal-vegetal axis also appears to structure the future anterior-
posterior axis, with the vegetal region sequestering those maternal Ventral
components necessary for posterior development.
Sea Urchin Gastrulation
• The late sea urchin blastula consists of a single layer of about 1000 cells that form a hollow ball,
somewhat flattened at the vegetal end. The blastomeres, derived from different regions of the zygote,
have different sizes and properties.
• The fates of the various regions of the blastula as it develops through gastrulation to the pluteus
larva stage characteristic of sea urchins. The fate of each cell layer can be seen through its movements
during gastrulation.
Ingression of primary mesenchyme
Function of primary mesenchyme cells
• During gastrulation the cells at vegetal side become thickened
and flat at the center of this flat vegetal plate, a cluster of small
cells begins to change. These cells begin extending and
contracting long, thin (30 × 5 μm) processes
called filopodia from their inner surfaces.
• The cells then dissociate from the epithelial monolayer and
ingress into the blastocoel. These cells, derived from the
micromeres, are called the primary mesenchyme. They will
form the larval skeleton, so they are sometimes called
the skeletogenic mesenchyme.
• Originally, all the cells of the blastula are connected on their
outer surface to the hyaline layer and on their inner surface
to a basal lamina secreted by the cells.
Stages of Gastrulation
First stage of archenteron invagination
• As the ring of primary mesenchyme cells leaves the vegetal region of the blastocoel, important changes are
occurring in the cells that remain at the vegetal plate. These cells remain bound to one another and to the
hyaline layer of the egg, and they move to fill the gaps caused by the ingression of the primary mesenchyme.
• Moreover, the vegetal plate bends inward and invaginates about one-fourth to one-half the way into the
blastocoel and then take a pause.
• The invaginated region is called the archenteron (primitive gut), and the opening of the archenteron at the
vegetal region is called the blastopore.
Second and third stages of archenteron invagination
• The invagination of the vegetal cells occurs in three discrete stages. After a brief pause,
the second phase of archenteron formation begins.
• During this time, the archenteron extends dramatically, sometimes tripling its length.
In this process of extension, the wide, short gut rudiment is transformed into a long, thin
tube.