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Characterizing the roles of the central apparatus C1a projection complexes in motility regulation and axoneme assembly

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Author	Goduti, Daniel J.
Title	Characterizing the roles of the central apparatus C1a projection complexes in motility regulation and axoneme assembly
Date	2012
Abstract	The eukaryotic cilium (also called flagellum) is a complex, microtubule-based organelle that is found on many different cell types. It produces movement by the precise activation of dyneins that are located along the length and the circumference of the structure. The dynein activity is regulated in part by the central pair and radial spoke structures of the axoneme, which are required for wild-type motility. For example, loss of the C1a projection, which assembles onto the central apparatus, results in twitchy flagellar movements and paralysis (Dutcher et al., 1984). This central pair projection comprises several proteins, including the calcium-interacting protein calmodulin, suggesting that C1a is required for calcium-dependent motility regulation (Wargo et al., 2005). To further characterize the role of the projection components in flagellar motility, we generated Chlamydomonas mutants that assemble partial central pair projections onto the axoneme. These mutants were analyzed biochemically, structurally, and functionally to determine the impact of the missing components. From these results, we conclude that the C1a projection has a role in regulating both inner and outer dynein arm activity to generate wild-type motility. Specifically, we have determined that the amino terminus of PF6, a protein component of the C1a complex, is important for the stable assembly of the C1a-34, C1a-32, and C1a-18 proteins, and that the loss of these components results in a reduced flagellar beat frequency and reduced swimming velocity of the cell. The flagellar waveform is not affected by this loss. Additionally, the carboxyl terminus of PF6 is required for motility. Attempts to reduce expression of C1a-86 by an artificial microRNA technique were successful for knockdown, but also generated additional mutations in genes necessary for central pair assembly, resulting in paralysis and preventing further analysis of any motility phenotypes. Overall, these conclusions suggest that each of the C1a components, including the PF6 protein, contribute to a unique feature of flagellar motility regulation. Future experiments will focus on generating and analyzing additional mutants lacking subsets of the C1a projection components as well as determining direct, indirect, or functional interactions between C1a and other axonemal components that are important for motility.
Description	xii, 132 leaves : ill. (some col.) ; 28 cm.
Format	application/pdf
Advisor	Smith, Elizabeth F.; Compton, Duane A.; Higgs, Henry N.; Sale, Winfield S.
College	Dartmouth College -- Department of Biological Sciences.
Issuing Body	Dartmouth Dissertations are made available through the Dartmouth Digital Library.
Degree	Thesis (Ph.D.)--Dartmouth College.
Copyright	All rights reserved. No part of this book may be reproduced in any form by any electronic or mechanical means without permission in writing from the author.
DOI	http://dx.doi.org/10.1349/ddlp.429

Title	Page 1
Format	application/pdf
Issuing Body	Dartmouth Dissertations are made available through the Dartmouth Digital Library.
Degree	Thesis (Ph.D.)--Dartmouth College.
Full Text	Characterizing the roles of the central apparatus C1a projection complexes in motility regulation and axoneme assembly A Thesis Submitted to the Faculty in partial fulfillment of the requirements for the degree of Doctor of Philosophy in the Department of Biological Sciences by Daniel J. Goduti DARTMOUTH COLLEGE Hanover, New Hampshire May 4, 2012 Examining Committee: (chair) Elizabeth F. Smith, Ph.D. Duane A. Compton, Ph.D. Henry N. Higgs, Ph.D. Winfield S. Sale, Ph.D. Brian W. Pogue, Ph.D. Dean of Graduate Studies

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