The mechanisms that determine the shape and organization of cells remain largely unknown. Green algae such as Chlamydomonas provide excellent model systems for studying cell geometry owing to their highly reproducible cell organization. Structural and genetic studies suggest that asymmetry of the centriole (basal body) plays a critical determining role in organizing the internal organization of algal cells, through the attachment of microtubule rootlets and other large fiber systems to specific sets of microtubule triplets on the centriole. Thus to understand cell organization, it will be critical to understand how the different triplets of the centriole come to have distinct molecular identities.
"Centrioles are barrel-shaped structures that, in human cells, are made up of triplet microtubules. The older of the two centrioles, often referred to as the 'mature' centriole, carries distal and subdistal appendages , and the two centrioles can be distinguished by staining for marker proteins    . In most species, centriole biogenesis involves the early formation of a so-called cartwheel at the base of the nascent centriole . "
[Show abstract][Hide abstract] ABSTRACT: Centrioles function in the assembly of centrosomes and cilia. Structural and numerical centrosome aberrations have long been implicated in cancer, and more recent genetic evidence directly links centrosomal proteins to the etiology of ciliopathies, dwarfism and microcephaly. To better understand these disease connections, it will be important to elucidate the biogenesis of centrioles as well as the controls that govern centriole duplication during the cell cycle. Moreover, it remains to be fully understood how these organelles organize a variety of dynamic microtubule-based structures in response to different physiological conditions. In proliferating cells, centrosomes are crucial for the assembly of microtubule arrays, including mitotic spindles, whereas in quiescent cells centrioles function as basal bodies in the formation of ciliary axonemes. In this short review, we briefly introduce the key gene products required for centriole duplication. Then we discuss recent findings on the centriole duplication factor STIL that point to centrosome amplification as a potential root cause for primary microcephaly in humans. We also present recent data on the role of a disease-related centriole-associated protein complex, Cep164-TTBK2, in ciliogenesis.
[Show abstract][Hide abstract] ABSTRACT: Directed fluid flow, which is achieved by the coordinated beating of motile cilia, is required for processes as diverse as cellular swimming, developmental patterning, and mucus clearance. Cilia are nucleated, anchored and aligned at the plasma membrane by basal bodies, cylindrical microtubule based structures with nine-fold radial symmetry. In the unicellular ciliate, Tetrahymena thermophila, two centrin family members associated with the basal body are important for both basal body organization and stabilization. We have identified a family of thirteen proteins in Tetrahymena containing centrin-binding repeats, related to those identified in the Saccharomyces cerevisiae Sfi1 protein, that we have named Sfr1-13 for Sfi1-repeat. Nine of the Sfr proteins localize in unique polarized patterns surrounding the basal body, suggesting non-identical roles in basal body organization and association with basal body accessory structures. Furthermore, the Sfr proteins are found in distinct basal body populations in Tetrahymena cells, indicating they are responsive to particular developmental programs. A complete genetic deletion of one of the family members, Sfr13, causes unstable basal bodies and defects in daughter basal body separation from the mother, phenotypes also observed with centrin disruption. It is likely the other Sfr family members are involved in distinct centrin functions, providing specificity to the tasks centrins perform at basal bodies.
[Show abstract][Hide abstract] ABSTRACT: Like many algae, Chlamydomonas reinhardtii is phototactic, using two anterior flagella to swim toward light optimal for photosynthesis. The flagella are responsive to signals initiated at the photosensory eyespot, which comprises photoreceptors in the plasma membrane and layers of pigment granules in the chloroplast. Phototaxis depends on placement of the eyespot at a specific asymmetric location relative to the flagella, basal bodies, and bundles of two or four highly acetylated microtubules, termed rootlets, which extend from the basal bodies toward the posterior of the cell. Previous work has shown that the eyespot is disassembled prior to cell division, and new eyespots are assembled in daughter cells adjacent to the nascent four-membered rootlet associated with the daughter basal body (D4), but the chronology of these assembly events has not been determined. Here we use immunofluorescence microscopy to follow assembly and acetylation of the D4 rootlet, localization of individual eyespot components in the plasma membrane or chloroplast envelope, and flagellar emergence during and immediately following cell division. We find that the D4 rootlet is assembled before the initiation of eyespot assembly, which occurs within the same time frame as rootlet acetylation and flagellar outgrowth. Photoreceptors in the plasma membrane are correctly localized in eyespot mutant cells lacking pigment granule layers, and chloroplast components of the eyespot assemble in mutant cells in which photoreceptor localization is retarded. The data are suggestive that plasma membrane and chloroplast components of the eyespot are independently responsive to a cytoskeletal positioning cue.
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