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Here we focus on the XY 1 Y 2  condition in male common shrew    Sorex   araneus    Linnaeus, 1758, applying electron microscopy and immunocytochemistry for a comprehensive analysis of structure, synapsis and behaviour of the sex trivalent in pachytene spermatocytes. The pachytene sex trivalent consists of three distinct parts: short and long synaptic  SC  fragments (between the X and Y 1  and between the X and Y 2 , respectively) and a long asynaptic region of the X in-between. Chromatin inactivation was revealed in the XY 1  synaptic region, the asynaptic region of the X and a very small asynaptic part of the Y 2 . This inactive part of the sex trivalent, that we named the ‘head’, forms a typical sex body and is located at the periphery of the meiotic nucleus at mid pachytene. The second part or ‘tail’, a long region of synapsis between the X and Y 2  chromosomes, is directed from the periphery into the nucleus. Based on the distribution patterns of four proteins involved in chromatin inactivation, we propose a model of meiotic silencing in shrew sex chromosomes. Thus, we conclude that pachytene sex chromosomes are structurally and functionally two different chromatin domains with specific nuclear topology: the peripheral inactivated ‘true’ sex chromosome regions (part of the X and the Y 1 ) and more centrally located transcriptionally active autosomal segments (part of the X and the Y 2 ).

Dual mechanism of chromatin remodeling in the common shrew sex trivalent (XY 1 Y 2 )