Kinesin-2 family motors in the unusual photoreceptor cilium.

This review focuses on recent advances in the understanding of kinesin-2 family motors in vertebrate photoreceptor development. Zebrafish photoreceptors develop by the 3rd day of embryogenesis, making it possible to study mutant phenotypes without the use of conditional alleles. Recent work using a zebrafish kif3b mutant allele validates the concept that the heterotrimeric kinesin II motor is generally required for ciliogenesis. In zebrafish photoreceptors, however, loss of kif3b function delays but does not block cilium formation. This is thought to occur because both kif3b or kif3c can dimerize with kif3a and function redundantly. The second ciliary kinesin thought to function in photoreceptor cells is kif17. Prior work has shown that either morpholino knockdown of this gene or the overexpression of its dominant negative form can reduce or delay photoreceptor cilium development without any evident impact on ciliogenesis in general. This has led to the idea that kif17 may play an important role only in some specialized cilium types, such the one in photoreceptor cells. In a recently identified kif17 mutant, however, photoreceptor outer segments are formed by 5 dpf and an obvious delay of outer segment formation is seen only at the earliest stage analyzed (3 dpf). This work suggests that kif17 plays a significant role mainly at an early stage of photoreceptor development. Taken together, these studies lead to an intriguing concept that as they differentiate photoreceptors alter their kinesin repertoire.