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GroEL1: A Dedicated Chaperone Involved in Mycolic Acid Biosynthesis during Biofilm Formation in Mycobacteria Anil Ojha,1 Mridula Anand,1 Apoorva Bhatt,2 Laurent Kremer,3 William R. Jacobs, Jr., 2 and Graham F. Hatfull1,* 1 Pittsburgh Bacteriophage Institute, Department of Biological Sciences, University of Pittsburgh, Pittsburgh, PA 15260, USA 2 Howard Hughes Medical Institute, Albert Einstein College of Medicine, 1300 Morris Park Avenue, Bronx, NY 10461, USA 3 ´ ´ ` Laboratoire de Dynamique Moleculaire des Interactions Membranaires, CNRS UMR 5539, Universite de Montpellier II, Place Eugene Bataillon, 34095 Montpellier Cedex 05, France *Contact: gfh@ DOI 10.1016/j.cell.2005.09.012 SUMMARY (Kolter and Losick, 1998; Miller and Bassler, 2001; Fux et al., 2005; Reguera and Kolter, 2005), but the role of bio- Mycobacteria are unusual in encoding films in the pathogenesis of Mycobacterium tuberculosis two GroEL paralogs, GroEL1 and GroEL2. and Mycobacterium leprae (the causative agents of tubercu- GroEL2 is essential—presumably providing losis and leprosy, respectively) remains unclear. Several non- tuberculous mycobacteria—including the saprophytic My- the housekeeping chaperone functions— cobacterium smegmatis and the opportunistic pathogens while groEL1 is nonessential, contains the Mycobacterium avium and Mycobacterium chelonae— attB site for phage Bxb1 integration, and