The Interactive Fly
Genes involved in tissue and organ development
Dualistic thinking presents a pitfall in any attempt to explain a real world vastly more complex than an either/or perspective. These difficulties are exemplified when formulating models to describe the basis of polarity in Drosophila leg morphogenesis.
A polar coordinate model is appealing because of the circular symmetry of the leg. In this hypothesis cells receive positional information from the disc center. The presumed coordinates are given by distance from the center (radial coordinate) and circumferential location (angular coordinate). Such a model could work in the real world if decapentaplegic or wingless transcription were limited to a quadrant of the disc. The diffusion of their protein products in this case would set the angular coordinates necessary for cell fate specification. The advantage of this system is that it fits in nicely with the radial symmetry of the leg disc.
A Cartesian model is equally appealing. A wingless-decapentaplegic zone could function as an X axis to specify one coordinate of cellular position while another unknown gene could perform a similar function for a Y axis. In this model positional information is determined by the lateral diffusion of multiple morphogens. The advantage inherent here is that it fits nicely with understanding of determination of positional information during segmentation, when gradients of multiple morphogens like Bicoid and Decapentaplegic establish positional identity along anterior-posterior and dorsal-ventral axes respectively.
Reality encroches upon these arguments when observations are made of the effects of mutation and of the expression patterns of genes involved in leg morphogenesis. The leg has a clear anterior-posterior boundary, suggesting a Cartesian model, yet expression patterns of Distal-less and aristaless show radial symmetry, suggesting a polar coordinate model. In addition, the sectored expression of wingless is also compatable with a polar model.
Current thinking inclines toward the Boundary model, a combination of the Cartesian and polar models. The Boundary model assumes that three or more compartments will be specified. This is thought to be the case in leg polarity, and known to be true for segmentation. These compartments would cooperate to cause the production of a specific morphogen at their point of intersection (the center of the disc). The conical concentration gradient formed by the diffusion of this morphogen would then specify a radial coordinate for all cells in the disc.
In the Boundary model, cell positional identity in the leg disc is defined by both Cartesian and polar coordinates. In Drosophila leg morphogenesis both models have to be called upon to explain all the facts.
Held, L. I. (1995). Axes, boundaries and coordinates: the ABCs of fly leg development. Bioessays 17: 721-732
Genes involved in organ development
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