The folding and unfolding of eukaryotic chromatin

In vivo, chromatin exists as fibres with differing degrees of compaction. We argue here that the packing density of the chromatin fibre is an important parameter, such that fibres with six nucleosomes/11 nm are enriched in ‘euchromatin’ while more highly compacted forms with higher packing densities correspond to some heterochromatic regions. The fibre forms differ in the extent of nucleosome stacking—in the ‘30 nm’ fibre stacking is suboptimal while in ‘heterochromatic’ fibres optimal stacking allows a greater compaction. One factor affecting the choice of different endpoints in fibre formation depends on the homogeneity and optimisation of linker length within a nucleosomal array. The ‘30 nm’ fibre can accommodate some variation in linker length while formation of the more compact forms requires that linker lengths be homogeneous and optimal. In vivo, chromatin remodelling machines and histone tail modifications would mediate and regulate this optimisation.