Conditional Visuomotor Learning Task SAMPLE DELAY CHOICE 1st Reversal 2nd Reversal A A A B B B etc…. 0 500 1500 Trial Time (ms) Asaad, W.F., Rainer, G. and Miller, E.K. (1998) Neuron, 21:1399-1407. Object and Direction selective PF neuron (nonlinear interaction) SAMPLE DELAY Spikes per second 50 A - go left A - go right B - go left B - go right 40 50 30 40 20 30 0 200 400 600 800 1000 Time (ms) 1200 1400 Movement Direction Left Right Stimulus Left Mapping Right Left Right Prefrontal cortex : May mediate learning of arbitrary associations. Many PF neurons coded both an object and a currently-associated directional response. During learning, information about the cue object and the action it instructed gradually merged together in PF activity. This may reflect the role of the PF cortex in acquiring and representing behavior-guiding rules, a function crucial for intelligent, adaptive behavior. Asaad, W.F., Rainer, G. and Miller, E.K. (1998) Neuron, 21:1399-1407. We trained monkeys to switch between two abstract rules: “match” and “nonmatch” Match (bar release) Match Rule (DMS) Sample Delay Test Nonmatch rule (DNMS) Four samples were used. New objects every day. Nonmatch (bar release) Wallis, J.D., Anderson, K.C., and Miller, E.K. (2000) Soc. Neurosci. Abstr. Condition Sample Rule Example Trial Cue Lever release + reward Match + low tone + no reward Nonmatch + high tone Lever release A Prefrontal Neuron Tuned to the Match Rule 2D Graph 1 Firing rate (Hz) 20 Sample Delay MATCH - reward MATCH - low tone 15 10 NONMATCH - no reward NONMATCH - high tone 5 500 1000 1500 Time (milliseconds) 2000 A Prefrontal Neuron Tuned to the Nonmatch Rule 2D Graph 1 20 Firing rate (Hz) Sample Delay NONMATCH - no reward NONMATCH - high tone 15 10 MATCH - reward MATCH - low tone 5 0 500 1000 1500 Time (milliseconds) 2000 The prefrontal cortex: Selectively represents goal-relevant information (focal attention, recall). Synthesizes information from diverse sources to serve a common behavioral goal (sensory inputs, stored knowledge). Is plastic: it neural activity changes to meet behavioral demands. Knits together arbitrary associations between diverse, but behaviorally-related information. Conveys information about the behavioral context in which the animals are engaged and the rules used to guide behavior. This may reflect the role of the PF cortex in acquiring and representing the formal demands of behavior, rules or models of tasks that provide a foundation for complex, intelligent behavior. (Cohen and Servan-Schreiber, 1992; Passingham, 1993; Grafman, 1994;Wise et al., 1996; Dehaene et al., 1998; Miller, 1999, Miller, 2000, Miller and Cohen, 2001). The PF cortex and cognitive control Phone rings Answer Don’t answer Active Inactive The PF cortex and cognitive control Phone rings Answer Don’t answer Active Inactive The PF cortex and cognitive control At home Guest Phone rings Answer Don’t answer Active Inactive The PF cortex and cognitive control PF cortex At home Guest Phone rings Answer Don’t answer Active Inactive The PF cortex and cognitive control PF cortex At home Reward signals (VTA neurons?) Guest Phone rings Answer Don’t answer Active Inactive The PF cortex and cognitive control PF cortex At home Guest Phone rings Answer Don’t answer Active Inactive The PF cortex and cognitive control PF cortex At home Reward signals (VTA neurons?) Guest Phone rings Answer Don’t answer Active Inactive The PF cortex and cognitive control PF cortex At home Guest Phone rings Answer Don’t answer Active Inactive The PF cortex and cognitive control PF cortex At home Guest Phone rings Answer Don’t answer Active Inactive The PF cortex and cognitive control PF cortex At home Guest Phone rings Answer Don’t answer Active Inactive The PF cortex and cognitive control PF cortex At home Guest Phone rings Answer Don’t answer Active Inactive The PF cortex and cognitive control PF cortex At home Guest Phone rings Answer Don’t answer Active Inactive The PF cortex and cognitive control PF cortex At home Guest Phone rings Answer Don’t answer Active Inactive The prefrontal cortex may be like a switch operator in a system of railroad tracks: Its integrative anatomy allows it to rapidly acquire a “map” that specifies which pattern of “tracks” (neural pathways) are needed to solve a given task. PF cortex The prefrontal cortex may be like a switch operator in a system of railroad tracks: Its integrative anatomy allows it to rapidly acquire a “map” that specifies which pattern of “tracks” (neural pathways) are needed to solve a given task. PF cortex The PF cortex actively maintains this pattern during task performance, allowing feedback signals to bias the flow of activity in other brain areas along those tracks. The prefrontal cortex may be like a switch operator in a system of railroad tracks: Its integrative anatomy allows it to rapidly acquire a “map” that specifies which pattern of “tracks” (neural pathways) are needed to solve a given task. PF cortex The PF cortex actively maintains this pattern during task performance, allowing feedback signals to bias the flow of activity in other brain areas along those tracks. Attention, retrieval, representation of rules and goals, response selection, inhibitory control, etc. can be explained by PF bias signals acting on different brain structures.