The synaptonemal complex, recombination nodules and chiasmata in human spermatocytes

Reconstruction from serial sections of 120 human spermatocyte nuclei, ranging in stage from leptotene to anaphase I, has led to the following conclusions. The reconstructed nuclei form a continuous sequence of developmental stages. With few exceptions, all of the bivalents are identifiable by length, centromere index and morphological markers. Crossing over is accompanied by a transformation of the recombination nodule (RN) into a fusiform bar lying across the synaptonemal complex (SC). The evolution of the crossover distribution at late pachytene is regulated at two levels both involving the SC. Each bivalent displays a pattern of SC regions - domains - with high affinity for RNs in the middle and low affinity at the ends. The occurrence of crossing over in a domain reduces the probability of attachment of new RNs and causes RNs not yet involved in crossing over to be released from that domain. Positive interference between crossovers is a consequence of these phenomena. At diplotene the degradation of the SC is initiated in regions with low affinity for RNs and small segments of the SC are retained at the crossover sites until late diakinesis. The site of crossing over is only identifiable by an RN for a certain period of time during pachytene. The distribution of total crossovers in a nucleus is similar but not identical to the distribution of chiasmata as seen in the light microscope. The number of crossovers has been estimated to be 70 and exceeds the number of chiasmata determined by light microscopy by a factor of 1.4.