Tyrosine hydroxylase immunostaining in the central complex of dicondylian insects

Dopamine acts as a neurohormone and neurotransmitter in the insect nervous system and controls a variety of physiological processes. Dopaminergic neurons also innervate the central complex (CX), a multisensory center of the insect brain involved in sky compass navigation, goal-directed locomotion and sleep control. To infer a possible influence of evolutionary history and lifestyle on the neurochemical architecture of the CX, we have studied the distribution of neurons immunoreactive to tyrosine hydroxylase (TH), the rate-limiting enzyme in dopamine biosynthesis. Analysis of representatives from 12 insect orders ranging from firebrats to flies revealed high conservation of immunolabeled neurons. One type of TH-immunoreactive neuron was found in all species studied. The neurons have somata in the pars intercerebralis, arborizations in the lateral accessory lobes, and axonal ramifications in the central body and noduli. In all pterygote species, a second type of tangential neuron of the upper division of the central body was TH-immunoreactive. The neurons have cell bodies near the calyces and arborizations in the superior protocerebrum. Both types of neuron showed species-specific variations in cell number and in the innervated areas outside and inside the CX. Additional neurons were found in only two taxa: one type of columnar neuron showed TH immunostaining in the water strider Gerris lacustris, but not in other Heteroptera, and a tritocerebral neuron innervating the protocerebral bridge was immunolabeled in Diptera. The data show largely taxon-specific variations of a common ground pattern of putatively dopaminergic neurons that may be commonly involved in state-dependent modulation of CX function.