Organization of the dopaminergic innervation of forebrain areas relevant to learning: A combined immunohistochemical/retrograde tracing study in the domestic chick

The mediorostral neostriatum/hyperstriatum ventrale (MNH) and neostriatum dorsocaudale (Ndc) of the domestic chick are crucially involved in auditory filial imprinting, whereas the lobus parolfactorius (LPO) seems to be involved in the emotional modulation of behavior. Because there is evidence that MNH and Ndc are akin to higher association areas in mammals, the present study evaluates the dopaminergic and thalamic input to these areas, as well as to the avian caudate/putamen homologue LPO, by using retrograde pathway tracing, together with dopamine (DA) and tyrosine hydroxylase (TH) immunohistochemistry. By combining DA immunohistochemistry with retrograde fluorescent tracing, we demonstrated that dopaminergic afferents to the MNH and Ndc arise mainly from the area ventralis, whereas the main dopaminergic input to the LPO arises from the substantia nigra. The main thalamic input to the MNH and LPO arises from the dorsal thalamic nuclei, n. dorsomedialis anterior and n. dorsolateralis anterior, whereas the thalamic input to the Ndc arises from the n. dorsolateralis posterior and n. subrotundus. Furthermore, there are reciprocal intratelencephalic connections between distinct parts of the neostriatum caudale and the mediorostral neostriatum. DA-immunoreactive (ir) fibers are present at moderate densities in the MNH and Ndc and at high densities in the LPO. At the ultrastructural level, DA- and TH-ir axon terminals in the MNH and Ndc form predominantly symmetric synaptic contacts with dendritic shafts, which are often situated in close vicinity to unstained terminals. These results indicate that the general organization of dopaminergic afferents to the chick telecephalon is similar to that of the mesotelencephalic dopaminergic subsystems in mammals such as the mesostriatal and mesolimbocortical DA system.