Orcokinin in the central complex of the locust Schistocerca gregaria: Identification of immunostained neurons and colocalization with other neuroactive substances.

The central complex is a group of highly interconnected neuropils in the insect brain. It is involved in the control of spatial orientation, based on external compass cues and various internal needs. The functional and neurochemical organization of the central complex has been studied in detail in the desert locust Schistocerca gregaria. In addition to classical neurotransmitters, immunocytochemistry has provided evidence for a major contribution of neuropeptides to neural signaling within the central complex. To complement these data, we have identified all orcokinin-immunoreactive neurons in the locust central complex and associated brain areas. About 50 bilateral pairs of neurons innervating all substructures of the central complex exhibit orcokinin immunoreactivity. Among these were about 20 columnar neurons, 33 bilateral pairs of tangential neurons of the central body, and seven pairs of tangential neurons of the protocerebral bridge. In silico transcript analysis suggests the presence of eight different orcokinin-A type peptides in the desert locust. Double label experiments showed that all orcokinin-immunostained tangential neurons of the lateral accessory lobe cluster were also immunoreactive for GABA and the GABA-synthesizing enzyme glutamic acid decarboxylase. Two types of tangential neurons of the upper division of the central body were, furthermore, also labeled with an antiserum against Dip-allatostatin I. No colocalization was found with serotonin immunostaining. The data provide additional insights into the neurochemical organization of the locust central complex and suggest that orcokinin-peptides of the orcokinin-A gene act as neuroactive substances at all stages of signal processing in this brain area.

Auditory cortex reflects goal-directed movement but is not necessary for behavioral adaptation in sound-cued reward tracking.

Mounting evidence suggests that the role of sensory cortices in perceptual decision making goes beyond the mere representation of the discriminative stimuli and additionally involves the representation of nonsensory variables such as reward expectation. However, the relevance of these representations for behavior is not clear. To address this issue, we trained rats to discriminate sounds in a single-interval forced-choice task and then confronted the animals with unsignaled blockwise changes of reward probabilities. We found that unequal reward probabilities for the two choice options led to substantial shifts in response bias without concomitant reduction in stimulus discrimination. Although decisional biases were on average less extreme than required to maximize overall reinforcement, a model-based analysis revealed that rats managed to harvest >97% of rewards. Neurons in auditory cortex recorded during task performance weakly differentiated the discriminative stimuli but more strongly the subsequent goal-directed movement. Although 10-20% of units exhibited significantly different firing rates between task epochs with different response biases, control experiments showed this to result from inflated false positive rates due to unspecific temporal correlations of spiking activity rather than changing reinforcement contingencies. Transient pharmacological inactivation of auditory cortex reduced sound discriminability without affecting other measures of performance, whereas inactivation of medial prefrontal cortex affected both discriminability and bias. Together, these results suggest that auditory cortex activity only weakly reflects decisional variables during flexible updating of stimulus-response-outcome contingencies and does not play a crucial role in sound-cued adaptive behavior, beyond the representation of the discriminative stimuli.NEW & NOTEWORTHY Recent evidence suggests that sensory cortex represents nonsensory variables such as reward expectation, but the relevance of these representations for behavior is not well understood. We show that rat auditory cortex (AC) is modulated during movement and reward anticipation in a sound-cued reward tracking task, whereas AC inactivation only impaired discrimination without affecting reward tracking, consistent with a predominantly sensory role of AC.