Initial asymmetries
in the egg along the anterior-posterior axis are set up by bicoid
and nanos mRNA localization at the anterior and posterior pole of
the embryo respectively. Their translated gene products diffuse through
the embryo, forming two opposing long-range protein gradients.
bicoid
and nanos regulate the translation of two other maternal mRNAs present
in the embryo: hunchback and caudal. hunchback and
caudal mRNA are uniformly distributed throughout the embryo. nanos,
the posterior determinant, represses hunchback translation, leading
to a Hunchback protein gradient complementary to that of Nanos. Similarly,
translational repression of caudal through bicoid leads to
a posterior Caudal gradient.
Although Bicoid acts as a translational repressor, its main function is
the transcriptional regulation of downstream zygotic target genes. bicoid
cooperates with maternal hunchback in the regulation of zygotic genes
expressed in the anterior region of the embryo: orthodenticle, buttonhead,
empty spiracles, zygotic hunchback and the anterior stripe
of giant expression. Together with caudal, it regulates genes
in the posterior region, like knirps and the posterior stripe of
giant expression. In contrast to the dual function of bicoid
as a regulator at the DNA and RNA level, the sole role of nanos is
to repress maternal hunchback translation. How the target genes of
Bicoid, Hunchback and Caudal pattern the embryo, eventually leading to its
subdivision in 14 segments, is extensively discussed under the topic segmentation.
The
fact that bicoid acts very early in development in a large syncytial
cell is the prerequisite for the rare circumstance that it is a transcription
factor AND a morphogen. As soon as cells are present, morphogens would have
to move between them.
The different responsiveness of target genes to bicoid concentration translates
the gradient into a spatial pattern. A gene X, responding to a certain concentration
of bicoid will be active only in a defined region of the embryo. A gene
Y may respond to a different concentration of bicoid. Therefore, the clue
for the spatial organization along the anterior-posterior axis lies in the
cis-regulatory regions of the target genes. In addition to Bicoid, factors
like Hunchback and Caudal, cooperate in conveying positional information.

Media list
Maternal interactions
Concentration gradient
Genes discussed
Gene
|
Gene product
- Domains
|
Function
|
Links
|
bicoid
(bcd)
|
transcription factor - homeodomain
|
maternally provided morphogen involved in anterior
patterning of the Drosophila embryo
|
|
buttonhead
(btd)
|
transcription
factor - zinc finger
|
transcriptonal
activator that regulates the segmentation of the head
|
|
caudal
(cad)
|
transcription
factor - homeodomain
|
plays a role in establishing the posterior domains
of the embryo
|
|
empty
spiracles (ems)
|
transcription
factor - homeodomain
|
required for head + brain development
|
|
hunchback
(hb)
|
transcription factor - zinc
finger
|
gap gene, later required for proper temporal generation
of NB sublineages
|
|
nanos
(nos)
|
translational
repressor - zinc finger
|
targets
Hunchback- and Bicoid-mRNAs to achieve posterior identity
|
|
orthodenticle
(otd)
|
transcription
factor - homeodomain - paired-like
|
acts in a combinatorial fashion with the cephalic gap
genes empty spiracles and buttonhead to
assign segmental identities in the head and brain
|
|
|