The goal of our research is to understand the molecular mechanism that controls head development. We are studying head formation in a simple model organism, the fruitfly Drosophila melanogaster. Many of the genes that control the development of the brain, sensory organs, and other head structures have been conserved throughout animal evolution. Using the powerful approaches possible in Drosophila research, we will determine how the head is divided into segments, and how the fates of these segments are specified. In the proposed experiments, we will investigate the role of the orthodenticle (otd) gene in head and brain development. otd-related genes play important roles in brain and sensory organ development in vertebrates. We will determine the molecular hierarchy in which otd participates and compare it to the hierarchy of genes that patterns the trunk region of the embryo. The molecular mechanism that patterns the head is not well understood. This study will contribute to elucidating this mechanism and determining how it differs from that which controls trunk development. Our results will provide a foundation for analyzing head/brain formation in higher animals. Ultimately, such studies will be critical for understanding the genetic defects that perturb normal pathways of development.