In both vertebrates and invertebrates, odorant molecules reach the dendrite
s of olfactory receptor cells through an aqueous medium, which reflects the
evolutionary origin of these systems in a marine environment. Important re
cent advances, however, have demonstrated striking interphyletic difference
s between the structure of vertebrate and invertebrate olfactory receptor p
roteins, as well as the organization of the genes encoding them. While thes
e disparities support independent origins for odor-processing systems in cr
aniates and protostomes (and even between the nasal and vomeronasal systems
of craniates), olfactory neuropils share close neuroanatomical and physiol
ogical characters. Whereas there is a case to be made for homology among me
mbers of the two great protostome clades (the ecdysozoans and lophotrochozo
ans), the position of the craniates remains ambiguous.