The cilium is an elongated and continuous structure that spans two major subcellular domains. The cytoplasmic domain contains a short centriole, which serves to nucleate the main projection of the cilium. This projection, known as the axoneme, remains separated from the cytoplasm by a specialized gatekeeping complex within a ciliary subdomain called the transition zone. In this way, the axoneme is compartmentalized. Intriguingly, however, this general principle of cilium biology is altered in the sperm cells of many animals, which instead contain a cytoplasmic axoneme domain. Here, we discuss the hypothesis that the formation of specialized sperm giant centrioles and cytoplasmic cilia is mediated by the migration of the transition zone from its typical location as part of a structure known as the annulus and examine the intrinsic properties of the transition zone that may facilitate its migratory behavior.
