The nucleosome: A little variation goes a long way

Emily Bernstein, Sandra B. Hake

Research output: Contribution to journalArticlepeer-review

99 Scopus citations


Changes in the overall structure of chromatin are essential for the proper regulation of cellular processes, including gene activation and silencing, DNA repair, chromosome segregation during mitosis and meiosis, X chromosome inactivation in female mammals, and chromatin compaction during apoptosis. Such alterations of the chromatin template occur through at least 3 interrelated mechanisms: post-translational modifications of histones, ATP-dependent chromatin remodeling, and the incorporation (or replacement) of specialized histone variants into chromatin. Of these mechanisms, the exchange of variants into and out of chromatin is the least well understood. However, the exchange of conventional histones for variant histones has distinct and profound consequences within the cell. This review focuses on the growing number of mammalian histone variants, their particular biological functions and unique features, and how they may affect the structure of the nucleosome. We propose that a given nucleosome might not consist of heterotypic variants, but rather, that only specific histone variants come together to form a homotypic nucleosome, a hypothesis that we refer to as the nucleosome code. Such nucleosomes might in turn participate in marking specific chromatin domains that may contribute to epigenetic inheritance.

Original languageEnglish
Pages (from-to)505-517
Number of pages13
JournalBiochemistry and Cell Biology
Issue number4
StatePublished - Aug 2006
Externally publishedYes


  • Heterotypic variant nucleosome
  • Histone variants
  • Homotypic variant nucleosome
  • Nucleosome code


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