Evolutionary changes in the olfactory projection neuron pathways of eumalacostracan crustaceans

Output from the olfactory lobe (primary olfactory center) of eumalacostracan crustaceans is transmitted to the medulla terminalis (MT) and hemiellipsoid body (HB) in the lateral protocerebrum (higher order center) by a large population of projection neurons. In eureptantian crustaceans (lobsters, crayfish, and crabs), these projection neurons also form the output pathway from an additional neuropil, the accessory lobe (higher order center), which appears to have arisen de novo in these animals. In a previous study of lobsters and crayfish we showed that whereas projection neurons innervating the olfactory lobe project primarily to the MT, those innervating the accessory lobe project exclusively to the HB (Sullivan and Beltz [ 2001a] J. Comp. Neurol. 441:9-22). In the present study, we used focal dye injections to examine the olfactory projection neuron pathways of representatives of four eumalacostracan taxa (Stomatopoda, Dendrobranchiata, Caridea, and Stenopodidea) that diverged from the eureptantian line prior to the appearance of the accessory lobe. These experiments were undertaken both to examine the evolution of the olfactory pathway in the Eumalacostraca and to provide insights into the changes in this pathway that accompanied the appearance of the accessory lobe. The innervation patterns of the olfactory projection neurons of the species examined were found to differ markedly, varying from that observed in the most basal taxon examined (Stomatopoda), in which the neurons primarily project to the MT, to those observed in the two highest taxa examined (Caridea and Stenopodidea), in which they primarily target the HB. These results suggest that substantial changes in the relative importance of the MT and HB within the olfactory pathway have occurred during the evolution of the Eumalacostraca.