Superficial cleavage

• In Drosophila, 256 nuclei are produced by a series of eight nuclear

divisions averaging 8 minutes each. The nuclei then migrate to the
periphery of the egg, where the mitoses continue, albeit at a
progressively slower rate.
• During the ninth division cycle, about five nuclei reach the surface of
the posterior pole of the embryo .These nuclei become enclosed by cell
membranes and generate the pole cells that give rise to the gametes of
the adult.
• Most of the other nuclei arrive at the periphery of the embryo at cycle
10 and then undergo four more divisions at progressively slower rates.
• During these stages of nuclear division, the embryo is called a
syncytial blastoderm.
• Although the nuclei divide within a common cytoplasm, this does not
mean that the cytoplasm is itself uniform; each nucleus within the
• syncytial blastoderm is contained within its own little territory of
cytoskeletal proteins. The nuclei and their associated cytoplasmic
islands are called energids.
superficial cleavage
 Gastrulation
• The prospective mesoderm about 1000 cells constituting the ventral
midline of the embryo folds inward to produce the ventral furrow.
• This furrow eventually pinches off from the surface to become a ventral
tube within the embryo. It then flattens to form a layer of mesodermal
tissue beneath the ventral ectoderm.
• The prospective endoderm invaginates as two pockets at the anterior
and posterior ends of the ventral furrow. The pole cells are internalized
along with the endoderm. At this time, the embryo bends to form the
cephalic furrow.
• The ectodermal cells on the surface and the mesoderm undergo
convergence and extension, migrating toward the ventral midline to
form the germ band, a collection of cells along the ventral midline that
includes all the cells that will form the trunk of the embryo. The germ
band extends posteriorly and, perhaps because of the egg case, wraps
around the top (dorsal) surface of the embryo.
• Gastrulation
• The body segments begin to
appear, dividing the ectoderm
and mesoderm. The germ band
then retracts, placing the
presumptive posterior segments
into the posterior tip of the
embryo
• The general body plan of
Drosophila is the same in the
embryo, the larva, and the adult,
each of which has a distinct head
end and a distinct tail end,
between which are repeating
segmental units. Three of these
segments form the thorax, while
another eight segments form the
abdomen. Each segment of the
adult fly has its own identity.
• How does this pattern arise?
 Anterior –Posterior axis
• The maternal effect genes expressed in the
mother's ovaries produce messenger RNAs that
are placed in different regions of the egg.
• Two of these proteins, Bicoid and Hunchback,
regulate th production of anterior structures,
while another pair of maternally specified
proteins, Nanos and Caudal, regulates the
formation of the posterior parts of the embryo.
• The zygotic genes regulated by these maternal
factors are expressed in certain broad (about
three segments wide), partially overlapping
domains. These genes are called gap genes
(because mutations in them cause gaps in the
segmentation pattern), and they are among the
first genes transcribed in the embryo.
• Differing concentrations of the gap gene proteins
cause the transcription of pair-rule genes, which
divide the embryo into periodic units. The
transcription of the different pair-rule genes results
in a striped pattern of seven vertical bands
perpendicular to the anteriorposterior axis. T
• he pair-rule gene proteins activate the transcription
of the segment polarity genes, whose mRNA and
protein products divide the embryo into 14
segment-wide units, establishing the periodicity of
the embryo. At the same time, the protein products
of the gap, pairrule, and segment polarity genes
interact to regulate another class of genes, the
homeotic selector genes, whose transcription
determines the developmental fate of each
segment.
An anterior-to-posterior gradient of Bicoid protein
An anterior-to-posterior gradient of Hunchback protein
A posterior-to-anterior gradient of Nanos protein
A posterior-to-anterior gradient of Caudal protein
bicoid and hunchback mRNAs, whose protein products
are critical for head and thorax formation
 Gap genes
• High levels of Hunchback protein
induce the expression of giant, while
the Krüppel transcript appears over the
region where Hunchback begins to
decline.
• It is thought that a gradient of the
Caudal protein, highest at the posterior
pole, is responsible for activating the
abdominal gap genes knirps and giant.
• Krüppel gene expression is negatively
regulated on its anterior boundary by
the Hunchback and Giant proteins and
on its posterior boundary by the
Knirps and Tailless proteins
Pair rule genes
• Three genes are known to be the primary
pair-rule genes. These genes hairy, even
skipped, and runt are essential for the
formation of the periodic pattern, and they
are directly controlled by the gap gene
proteins.
• The second even skipped stripe is
repressed by both Giant and Krüppel
proteins and is activated by Hunchback
protein and low concentrations of Bicoid.
• Similarly, even-skipped stripe 5 is
regulated negatively by Krüppel protein
(on its anterior border) and by Giant
protein (on its posterior border).
 Segment polarity genes
• Interactions take place between the cells.
• reinforce the para-segmental periodicity
established by the earlier transcription factors.
• Second, through this cell-to-cell signalling, cell
fates are established within each para-segment.
• Mutations in these genes lead to defects in
segmentation and in gene expression pattern
across each parasegment.
 Segment polarity genes
• The engrailed gene is activated when cells have high levels of the
Even-skipped, Fushi tarazu, or Paired transcription factors; it is
repressed in those cells that receive high levels of Odd skipped,
Runt, or Sloppy-paired proteins.
• These stripes of engrailed transcription mark the anterior boundary
of each parasegment and the posterior border of each segment.
• The wingless gene is activated in those bands of cells that receive
little or no Even-skipped or Fushi tarazu proteins, but which do
contain the Sloppy-paired protein. This causes wingless to be
transcribed solely in the row of cells directly anterior to the cells
where engrailed is transcribed.
• This pattern is maintained as Engrailed-synthesizing cells secrete
the Hedgehog protein, which maintains the expression of the
wingless gene in the neighbouring cells, while the
Wingless-secreting cells maintain the expression of the engrailed
and hedgehog genes in their neighbours in turn.
The Homeotic Selector Genes
 Homeotic Antennapedia gene is
mutants expressed in the
head

When the ultrabithorax gene is

deleted