Zygotene-Pachytene Substaging and Synaptonemal Complex Karyotyping of Boar Spermatocytes

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Abstract
Synaptonemal complex analysis, by electron microscopy of spread spermatocytes after phosphotungstic acid staining, made possible description of the chromosomal synaptic patterns and the synaptonemal complex karyotype of the pig (Sus scrofa domestica). The autosomal synaptic pattern in conjunction with the sex chromosome morphology and pairing behaviour can serve as a reference for the meiotic cell progression from the zygotene stage to the pachytene. The autosomes started terminal synapsis at early zygotene (Z1) and at mid-zygotene (Z2) some of the small bivalents were completely paired. The extension of pairing between the X and the Y chromosome, and the differentiation of their axes, disclosed seven types of sex bivalent (Types 0-VI). The Type 0 sex bivalent occurred in late zygotene (Z3), at which the X and Y axes began terminal synapsis by their short arms. Each one of the pachytene substages, early, mid-, and late, included two sex bivalent types. By early pachytene (P1-P2) the entire Y chromosome was paired with the X axis. Subsequently, progressive desynapsis and differentiation of the sex chromosome axes defined the mid- (P3-P4) and late pachytene substages (P5-P6). At mid-pachytene, the unpaired XY segments were associated end-to-end and showed differing degrees of complexity (thickening, splitting, despiralization and strandedness). These axial complexities were replaced at late pachytene by fine excrescences along the sex chromosome axes, which still formed a short SC stretch. Additionally, an attempt to construct an SC karyotype for the pig is presented. There was general agreement between the autosomal SC karyotype and the autosomal somatic karyotype when comparisons were made, considering the relative lengths and arm ratios of individual chromosome pairs.