Interacting functions of snail, twist and huckebein during the early development of germ layers in Drosophila.

Two zygotic genes, snail (sna) and twist (twi), are required for mesoderm development, which begins with the formation of the ventral furrow. Both twi and sna are expressed ventrally in the blastoderm, encode transcription factors and promote the invagination of the ventral furrow by activating or repressing appropriate target genes. However, sna and twi alone do not define the position of the ventral furrow, since they are also expressed in ventral cells that do not invaginate. We show that huckebein (hkb) sets the anterior and the posterior borders of the ventral furrow, but acts by different modes of regulation. In the posterior part of the blastoderm, hkb represses the expression of sna in the endodermal primordium (which we suggest to be adjacent to the mesodermal primordium). In the anterior part, hkb antagonizes the activation of target genes by twi and sna. Here, bicoid permits the co-expression of hkb, sna and twi, which are all required for the development of the anterior digestive tract. We suggest that mesodermal fate is determined where sna and twi but not hkb are expressed. Anteriorly hkb together with sna determines endodermal fate, and hkb together with sna and twi are required for foregut development.