Fine structure and behaviour of a pericentric inversion in the sand rat, Psammomys obesus

Abstract
In pachytene spermatocytes of the sand rat, Psammomys obesus, a long autosomal bivalent was observed, which was asynaptic for a large interstitial segment of its length in early pachytene. This bivalent also exhibited unaligned kinetochores. In late pachytene spermatocytes all autosomal bivalents were fully synapsed, but one of the shortest bivalents now possessed unaligned kinetochores. Evidence is presented in support of the proposition that the asynaptic interstitial region observed in early pachytene is due to the bivalent being heterozygous for a pericentric inversion. Using the maximum extent of homologous pairing, the break points were mapped at 26% from one end and 20% from the other. The unaligned kinetochores support the proposal that the aberration is an inversion and measurements of their positions confirm the estimated break points. In one cell a bivalent with interstitial (but no terminal) synapsis also confirms the inversion hypothesis. It is proposed that the bivalent is so small that topological considerations prevent the formation of the expected inversion loop. Evidence is also presented that complete synapsis of the bivalent during late pachytene can be attributed to ‘synaptic adjustment’, characterized by non-homologous synapsis (heterosynapsis). The position of the aberrant bivalent in relation to the sex chromosomes also changes during pachytene. When the bivalent is incompletely synapsed it generally associates by its ends with the ends of the sex chromosomes, but when it is non-homologously synapsed it is not associated with them.