Five chromatosome structures used as reference in this study: PDBs 4QLC, 5NL0, and 5WCU (rows 1–3) exhibit an on-dyad binding mode of the linker histone.

The reference structure of the off-dyad binding mode (row 5) was obtained via HADDOCK docking using a nucleosome from 1ZBB and PDB 1HST as the histone. [10, 50]. PDB 1ZBB is used as a reference for a an array of multiple nucleosomes. The last row represents the present study. We use PDB 1GHC as the linker histone (also see Fig 1 for a sequence alignment of the linker histones).