Distribution of centrifugal neurons targeting the soma clusters of the olfactory midbrain among decapod crustaceans.

To determine the distribution of two systems of centrifugal neurons innervating the soma clusters of the olfactory midbrain across decapod crustaceans, brains of the following nine species comprising most infraorders were immunostained with antibodies against dopamine and the neuropeptides substance P and FMRFamide: Macrobrachium rosenbergii, Homarus americanus, Cherax destructor, Orconectes limosus, Procambarus clarkii, Astacus leptodactylus, Carcinus maenas, Eriocheir sinensis and Pagurus bernhardus. One system consisting of several neurons with dopamine-like immunoreactivity that originate in the eyestalk ganglia was present in the four crayfish but not in any other species. These neurons project mainly into the lateral soma clusters (cluster 10) comprising the somata of ascending olfactory projection neurons and innervate very sparsely the medial soma clusters (clusters 9 and 11) containing the somata of local interneurons. In the innervation pattern of the lateral cluster, the dopamine-immunoreactive neurons showed large species-specific differences. The other system comprises a pair of giant neurons with substance P-like immunoreactivity. These neurons have somata in the median protocerebrum of the central brain and major projections into the lateral clusters and the core of the olfactory lobes, the neuropils that are the first synaptic relay in the central olfactory pathway of decapods; minor arborizations are present in the medial clusters. The system of substance P-immunoreactive giant neurons was present and of great morphological similarity in all studied species. Only in one species, the shrimp Macrobrachium rosenbergii, evidence for co-localization of FMRFamide-like with substance P-like immunoreactivity in these neurons was obtained. These and previously collected data indicate that the centrifugal neurons with dopamine-like immunoreactivity may be associated with the presence of an accessory lobe, a second-order neuropil that receives input from the olfactory lobe and only occurs in spiny lobsters, clawed lobsters and crayfish. The pair of centrifugal giant neurons with substance P-like immunoreactivity, on the other hand, appears to be a constitutive component of the decapod crustacean brain that most likely is functionally associated with the olfactory lobe. Both systems apparently exert modulatory functions on olfactory information processing by preferentially targeting the somata of the projection neurons. Thus, in the olfactory projection neurons, the somata seem to be more directly involved in information processing than in most other neurons of the arthropod CNS.