Localized requirement for torso-like expression in follicle cells for development of terminal anlagen of the Drosophila embryo.

The torso (tor) gene, one of six identified maternal genes essential for the development of the anterior and posterior terminal structures in the Drosophila embryo, is likely to function as a transmembrane receptor tyrosine kinase. Although tor protein is uniformly distributed in the membrane of the egg cell and syncytial embryo, genetic and molecular data suggest that tor is locally activated at the ends of the embryo by a ligand present in the perivitelline space. Local activation of tor could be achieved if the ligand were expressed by a subpopulation of the follicle cells that surround the developing oocyte. Here we describe torso-like (tsl), the sixth member of the terminal gene class, and show that it is unique among these genes in that its expression is required in the somatic follicle cells rather than in the germ line. Moreover, mosaic analysis demonstrates that tsl expression is necessary only in subpopulations of follicle cells located at the poles of the oocyte. Thus, the spatially regulated expression of tsl in the follicle cell layer may generate a localized signal that is transduced by tor, ultimately resulting in the formation of the terminal structures of the embryo.

Segmental polarity and identity in the abdomen of Drosophila is controlled by the relative position of gap gene expression.

The establishment of the segmental pattern in the Drosophila embryo is directed by three sets of maternal genes: the anterior, the terminal and the posterior group of genes. Embryos derived from females mutant for one of the posterior group genes lack abdominal segmentation. This phenotype can be rescued by transplantation of posterior pole plasm into the abdominal region of mutant embryos. We transplanted posterior pole plasma into the middle of embryos mutant either for the posterior, the anterior and posterior, or all three maternal systems and monitored the segmentation pattern as well as the expression of the zygotic gap gene Krüppel in control and injected embryos. We conclude that polarity and identity of the abdominal segments do not depend on the relative concentration of posterior activity but rather on the position of gap gene expression. By changing the pattern of gap gene expression, the orientation of the abdomen can be reversed. These experiments suggest that maternal gene products act in a strictly hierarchical manner. The function of the maternal gene products becomes dispensable once the position of the zygotically expressed gap genes is determined. Subsequently the gap genes will control the pattern of the pair-rule and segment polarity genes.

Determination of anteroposterior polarity in Drosophila.

The principles of pattern formation in embryogenesis can be studied in Drosophila by means of a powerful combination of genetic and transplantation experiments. The segmented pattern of the Drosophila embryo is organized by two activities localized at the anterior and posterior egg poles. Both activities exert inducing and polarizing effects on the pattern when transplanted to other egg regions. A small set of maternal genes have been identified that are required for these activities. Mutants in these genes lack either the anterior or posterior part of the segmented pattern. The unsegmented terminal embryonic regions require a third class of genes and form independently of the anterior and posterior centers.