Synaptic initiation and extension as prerequisites for crossing over

The effect on crossover frequency in maize of three hour heat treatment was studied when treatment was applied at zygotene (substantially later than the major DNA synthetic period) and at pachytene. Crossover frequency assay was based upon bridge and fragment frequency at anaphase I in heterozygotes for a short paracentric inversion. Effect of treatment was studied in three distinguishable synaptic classes: (1) overall crossover frequency within the inversion, (2) double crossover frequency where two separate events of pairing initiation are required (coincident crossovers within and proximal to the inversion) and (3) double crossover frequency within the inversion, where spreading of synapsis over a short distance from a single event of pairing initiation can provide the requisite pairing. Evidence is reported: (1) that overall crossover frequency within the inversion was very significantly increased by treatment at zygotene but not detectably affected by treatment at pachytene; (2) that double crossover frequency within the inversion was very significantly increased by treatment at pachytene and may have been somewhat increased by treatment at zygotene. Results are consistent with the model that most crossover sites may be established at, or approximately at, events of synaptic initiation but that establishment of infrequent second crossover sites near those formed first can follow or accompany the spreading of synapsis to adjoining regions.