Linking cell-fate specification to planar polarity: determination of the R3/R4 photoreceptors is a prerequisite for the interpretation of the Frizzled mediated polarity signal

The adult eye of Drosophila is a highly ordered structure composed of about 800 ommatidia, each displaying precise polarity. The planar polarity is reflected in the mirror-symmetric arrangement of ommatidia relative to the dorso-ventral midline, the equator. This arrangement is generated when ommatidia rotate towards the equator and the photoreceptor R3 displaces R4 creating different chiral forms in each half. Analysis of ommatidia mosaic for the tissue polarity gene frizzled (fz) has shown that the presence of a single Fz+ photoreceptor cell within the R3/ R4 pair is critical for the direction of rotation and chirality. By analysing clones mutant for seven-up (svp), in which R3/R4 precursors reside in their normal positions and become photoreceptor neurones but fail to adopt the normal R3/R4 fate, we find that the R3/R4 photoreceptor subtype specification is a prerequisite for planar polarisation in the eye. Moreover, in mosaic R3/R4 pairs we find that the svp- cell always adopts the R4 position. This bias is reminiscent of what happens in fz mosaic R3/R4 pairs, where the fz- cell also almost always adopts the R4 position. In addition, we find that in genotypes where too many cells adopt the R3/R4 fate, ommatidial polarity is also disturbed. Taken together, these data imply that correct specification of a single R3 cell per ommatidium is essential for the normal interpretation of the Fz-mediated polarity signal.