The anterior and posterior signaling systems
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