Structural models of site-specific recombinases from the lambda integrase f
amily of enzymes have in the last four years provided an important new pers
pective on the three-dimensional nature of the recombination pathway. Membe
rs of this family, which include the bacteriophage pi Cre recombinase, bact
eriophage lambda integrase, the yeast Flp recombinase, and the bacterial Xe
rCD recombinases, exchange strands between DNA substrates in a stepwise pro
cess. One pair of strands is exchanged to form a Holliday junction intermed
iate, and the second pair of strands is exchanged during resolution of the
junction to products. Crystal structures of reaction intermediates in the C
re-loxP site-specific recombination system, together with recent biochemica
l studies in the field, support a "strand swapping" model for recombination
that does not require branch migration of the Holliday junction intermedia
te in order to test homology between recombining sites.