Functional diversity of ISWI complexes

The yeast SWI/SNF ATP-dependent chromatin remodeling complex was first identified and characterized over 10 years ago (F. Winston and M. Carlson. 1992. Trends Genet. 8: 387-391.) Since then, the number of distinct ATP-dependent chromatin remodeling complexes and the variety of roles they play in nuclear processes have become dizzying (J.A. Martens and F. Winston. 2003. Curr. Opin. Genet. Dev. 13: 136-142; A. Vacquero et al. 2003. Sci. Aging Knowledge Environ. 2003: RE4) - and that does not even include the companion suite of histone modifying enzymes, which exhibit a comparable diversity in both number of complexes and variety of functions (M.J. Carrozza et al. 2003. Trends Genet. 19: 321-329; W. Fischle et al. 2003. Curr. Opin. Cell Biol. 15: 172-183; M. Iizuka and M.M. Smith. 2003. Curr. Opin. Genet. Dev. 13: 1529-1539). This vast complexity is hardly surprising, given that all nuclear processes that involve DNA - transcription, replication, repair, recombination, sister chromatid cohesion, etc. - must all occur in the context of chromatin. The SWI/SNF-related ATP-dependent remodelers are divided into a number of subfamilies, all related by the SWI2/SNF2 ATPase at their catalytic core. In nearly every species where researchers have looked for them, one or more members of each subfamily have been identified. Even the budding yeast, with its comparatively small genome, contains eight different chromatin remodelers in five different subfamilies. This review will focus on just one subfamily, the Imitation Switch (ISWI) family, which is proving to be one of the most diverse groups of chromatin remodelers in both form and function.