The ultrastructure of theChlamydomonas reinhardtiibasal apparatus: identification of an early marker of radial asymmetry inherent in the basal body

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Abstract
The biflagellate unicellular green alga Chlamydomonas reinhardtii is a classic model organism for the analysis of flagella and their organizers, the basal bodies. In this cell, the two flagella-bearing basal bodies, along with two probasal bodies and an array of fibers and microtubules, form a complex organelle called the basal apparatus. The ultrastructure of the basal apparatus was analysed in detail by serial thin-section electron microscopy of isolated cytoskeletons and several newly discovered features are described, including a marker for the rotational asymmetry inherent in the basal bodies and probasal bodies. In addition, the complex three-dimensional basal apparatus ultrastructure is resolved and illustrated, including the attachment sites of all basal apparatus elements to specific microtubular triplets of the basal bodies and probasal bodies. These data will facilitate both the localization of novel basal apparatus proteins and the analysis of mutants and RNA interference cells with only subtle defects in basal apparatus ultrastructure. The early harbinger of radial asymmetry described here could play a crucial role during basal body maturation by orienting the asymmetric attachment of the various associated fibers and therefore might define the orientation of the basal bodies and, ultimately, the direction of flagellar beating.