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Histone deposition proteins: links between the DNA replication machinery and epigenetic gene silencing.

organized into a nucleoprotein complex called chromatin. Chromatin is essential for compacting genomic DNA and plays a primary role in governing the expression status of genes. Chromatin is a diverse and dynamic polymer whose composition is modulated to build specialized chromosome structures, such as those found at centromeres and telomeres (for review, see Grewal and Moazed 2003). The fundamental repeating unit of chromatin is the nucleosome, containing an octamer of histone proteins, two each of H2A, H2B, H3, and H4, around which 146 bp of DNA wraps 1.7 times. This review will focus on how nucleosomes are assembled by histone chaperones that deposit histones during DNA replication or RNA transcription. We will begin by describing the biophysical properties governing nucleosome formation and the mechanisms of de novo nucleosome formation at sites of DNA synthesis. Then, we will discuss the functional overlap between histone chaperones involved in DNA replication-coupled and replication-independent nucleosome assembly. Finally, we will review recent data demonstrating the role of histone chaperones in the tight temporal coupling of histone deposition and DNA replication.