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Originally published In Press as doi:10.1074/jbc.M208652200 on November 9, 2002

J. Biol. Chem., Vol. 278, Issue 4, 2604-2613, January 24, 2003
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Functional Analysis of a Divergent System II Protein, Ccs1, Involved in c-Type Cytochrome Biogenesis*,

Beth Welty Dreyfuss, Patrice P. Hamel, Stacie S. Nakamoto, and Sabeeha MerchantDagger

From the Department of Chemistry and Biochemistry, UCLA, Los Angeles, California 90095-1569

The Ccs1 gene, encoding a highly divergent novel component of a system II type c-type cytochrome biogenesis pathway, is encoded by the previously defined CCS1 locus in Chlamydomonas reinhardtii. phoA and lacZalpha bacterial topological reporters were used to deduce a topological model of the Synechocystis sp. 6803 Ccs1 homologue, CcsB. CcsB, and therefore by analogy Ccs1, possesses a large soluble lumenal domain at its C terminus that is tethered in the thylakoid membrane by three closely spaced transmembrane domains in the N-terminal portion of the protein. Molecular analysis of ccs1 alleles reveals that the entire C-terminal soluble domain is essential for Ccs1 function and that a stromal loop appears to be important in vivo, at least for maintenance of Ccs1. Site-directed mutational analysis reveals that a single histidine (His274) within the last transmembrane domain, preceding the large lumenal domain, is required for c-type cytochrome assembly, whereas an invariant cysteine residue (Cys199) is shown to be non-essential. Ccs1 is proposed to interact with other Ccs components based on its reduced accumulation in ccs2, ccs3, ccs4, and ccsA strains.

* This work was supported by National Institutes of Health Grant GM48350, NRSA Grant GM17483 from the National Institutes of Health (to B. W. D.), American Heart Association Post-doctoral Fellowship 0120100Y (to P. H.), and a United States Public Health Service NRSA Award GM07185 from the National Institutes of Health (to S. S. N.).The costs of publication of this article were defrayed in part by the payment of page charges. The article must therefore be hereby marked "advertisement" in accordance with 18 U.S.C. Section 1734 solely to indicate this fact.

The on-line version of this article (available at http://www.jbc.org) contains Experimental Procedures, Figs. 1 and 2, and Table I.

Dagger To whom correspondence should be addressed: Dept. of Chemistry and Biochemistry, UCLA, Box 951569, Los Angeles, CA 90095-1569. Tel.: 310-825-8300; Fax: 310-206-1035; E-mail: merchant@chem.ucla.edu.

Copyright © 2003 by The American Society for Biochemistry and Molecular Biology, Inc.
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