The product topology of Xer-mediated site-specific recombination at pl
asmid sites has been determined. The product of deletion at pSC101 psi
is a right-handed antiparallel 4-noded catenane. The ColE1 cer deleti
on product has an identical topology, except that only one pair of str
ands is exchanged. These specific product topologies imply that the pr
oductive synaptic complex and the strand exchange mechanism have fixed
topologies. Further analysis suggests that synapsis traps exactly thr
ee negative supercoils between recombining sites, and that strand exch
ange introduces a further negative topological node in the deletion re
action. We present a model in which the requirement for a specific syn
aptic stucture, with two recombination sites interwrapped around the a
ccessory proteins ArgR and PepA, ensures that recombination only occur
s efficiently between directly repeated sites on the same DNA molecule
.