PMC:400732 / 17593-21146 JSONTXT

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A future extension of this study would be to test performance over a fixed number of trials and determine patterns of error rates during acquisition of the discrimination and reversal learning tasks. This manipulation would control the number of errors based on trials performed to determine if the apparent difference in absolute number of errors demonstrated by the mutant mice (the putative performance deficit) is simply a reflection of extra trials. Nonetheless, the inability of D2R-/- mice to disengage from previously established S+/S- contingencies (responding to a discriminative stimulus; Fig. 4) strongly argues that mesolimbic dopamine, and in particular dopamine D2R-mediated signaling, modulates the process of alerting the subject that familiar contingencies are now associated unexpected consequences.\nSchultz and colleagues have demonstrated that dopamine cells display consistent tonic firing patterns during maintenance of associative learning tasks [33]. However, phasic burst activity of dopaminergic cells occurs when discrepancies between predicted and actual reinforcement contingencies transpire [14]. This robust increase of dopaminergic cell activation in response to unpredicted outcomes has been referred to as an \"error\" signal [14]. To date, the molecular basis of this error signal has not been identified. The inability of the D2R-/- mice to desist responding to a previously reinforced stimulus suggests that the dopamine D2R might in fact be the focal point of this error-signaling cascade.\nElectrophysiological data further support the hypothesis that signaling mediated by dopamine D2Rs tunes D1R-mediated mesocorticolimbic output. Calabresi et al. [34] demonstrated that tetanic stimulation of dorsal striatum slices prepared from D2R-/- mice is associated with enhanced EPSP and as a result increased striatal synaptic efficacy. In contrast, stimulation of dorsal striatum slices from wild-type mice resulted in IPSP activity, long-term depression, and decreased neuronal activity of striatal efferents [34]. Carlsson and colleagues [35] have speculated that dopamine D2R stimulation in the striatum serves to \"brake\" or diminish excitatory corticostriatal signaling and plasticity. Indeed, perseverative behavior is associated with over activity of the dorsal striatum in rodents [36] and over activity of the caudate in patients with ADHD [37] and a strong inverse correlation of D2R binding with compulsive behavior has been reported [22]. Importantly, our data indicate that the poor performances displayed by the D2R-/- mice are manifestations of reversal learning deficits and not gross motivational or sensory impairments. We therefore argue that D2Rs participate in signaling or alerting the organism of learning contingency changes during reversal learning and sculpt ongoing goal-directed 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