“…Histone H3 variant H3.3 correlates with active gene expression (Stroud et al, 2012;Wollmann et al, 2012Wollmann et al, , 2017Shu et al, 2014;Duc et al, 2017), its incorporation results in more unstable nucleosomes (Jin et al, 2009;Thakar et al, 2009) and a transcription-permissive chromatin state, whereas H3.1 is enriched at transcriptionally inactive chromatin (Stroud et al, 2012;Wollmann et al, 2012). Another feature of heterochromatin is the presence of linker histones H1 that bind to DNA both at the entry and exit site of the nucleosome particle and which are enriched at CCs (She et al, 2013;Fyodorov et al, 2018). Although the characteristics of heterochromatin in leaf tissues with conspicuous CCs were well defined in previous reports, few studies have yet addressed the dynamic nature of CC structures and the implicated changes in chromatin features (Tessadori et al, 2007a,b;van Zanten et al, 2012;She et al, 2013).…”