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J. Biol. Chem., Vol. 279, Issue 23, 24757-24764, June 4, 2004

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The Trypanosoma brucei Cyclin, CYC2, Is Required for Cell Cycle Progression through G₁ Phase and for Maintenance of Procyclic Form Cell Morphology^*

Tansy C. Hammarton, Markus Engstler, and Jeremy C. Mottram¶

From the Wellcome Centre for Molecular Parasitology, the Anderson College, University of Glasgow, 56 Dumbarton Road, Glasgow G11 6NU, Scotland, United Kingdom, and Ludwig-Maximilians-Universität, Department Biologie I, Genetik, Maria-Ward-Strasse 1a, München 80638, Germany

CYC2 is an essential PHO80-like cyclin that forms a complex with the cdc2-related kinase CRK3 in Trypanosoma brucei. In both procyclic and bloodstream form T. brucei, knock-down of CYC2 by RNA interference (RNAi) led to an accumulation of cells in G₁ phase. Additionally, in procyclic cells, but not in bloodstream form cells, CYC2 RNAi induced a specific cell elongation at the posterior end. The G₁ block, as well as the posterior end elongation in the procyclic form, was irreversible once established. Staining for tyrosinated -tubulin and morphometric analyses showed that the posterior end elongation occurred through active microtubule extension, with no repositioning of the kinetoplast. Hence, these cells can be classified as exhibiting the "nozzle" phenotype as has been described for cells that ectopically express TbZFP2, a zinc finger protein that is involved in the differentiation of the bloodstream form to procyclic form. Within the tsetse fly, procyclic trypanosomes differentiate to elongated mesocyclic cells. However, although mesocyclic trypanosomes isolated from tsetse flies also show active microtubule extension at the posterior end, the kinetoplast is coincidentally repositioned such that it always lies approximately midway between the nucleus and posterior end of the cell. Thus, in the procyclic form CYC2 has dual functionality and is required for both cell cycle progression through G₁ and for the maintenance of correct cell morphology, whereas in the bloodstream form only a role for CYC2 in G₁ progression is evident.

Received for publication, February 5, 2004 , and in revised form, March 16, 2004.

^* This work was supported by the Medical Research Council and the British Council. The costs of publication of this article were defrayed in part by the payment of page charges. This article must therefore be hereby marked "advertisement" in accordance with 18 U.S.C. Section 1734 solely to indicate this fact.

^¶ To whom correspondence should be addressed. Tel.: 44-141-330-3745; Fax: 44-141-330-5422; E-mail: j.mottram{at}udcf.gla.ac.uk.

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