▪ Abstract The prefrontal cortex has long been suspected to play an important role in cognitive control, in the ability to orchestrate thought and action in accordance with internal goals. Its neural basis, however, has remained a mystery. Here, we propose that cognitive control stems from the active maintenance of patterns of activity in the prefrontal cortex that represent goals and the means to achieve them. They provide bias signals to other brain structures whose net effect is to guide the flow of activity along neural pathways that establish the proper mappings between inputs, internal states, and outputs needed to perform a given task. We review neurophysiological, neurobiological, neuroimaging, and computational studies that support this theory and discuss its implications as well as further issues to be addressed

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An integrative theory of prefrontal cortex function

Probability Distributions.- Linear Models for Regression.- Linear Models for Classification.- Neural Networks.- Kernel Methods.- Sparse Kernel Machines.- Graphical Models.- Mixture Models and EM.- Approximate Inference.- Sampling Methods.- Continuous Latent Variables.- Sequential Data.- Combining Models.

Pattern Recognition and Machine Learning

Prelude. Cycle 1. Introduction. Cycle 2. Structure defines function. Cycle 3. Diversity of cortical functions is provided by inhibition. Cycle 4. Windows on the brain. Cycle 5. A system of rhythms: from simple to complex dynamics. Cycle 6. Synchronization by oscillation. Cycle 7. The brain's default state: self-organized oscillations in rest and sleep. Cycle 8. Perturbation of the default patterns by experience. Cycle 9. The gamma buzz: gluing by oscillations in the waking brain. Cycle 10. Perceptions and actions are brain state-dependent. Cycle 11. Oscillations in the "other cortex:" navigation in real and memory space. Cycle 12. Coupling of systems by oscillations. Cycle 13. The tough problem. References.

Rhythms of the brain

The authors present a unified account of 2 neural systems concerned with the development and expression of adaptive behaviors: a mesencephalic dopamine system for reinforcement learning and a “generic” error-processing system associated with the anterior cingulate cortex The existence of the error-processing system has been inferred from the error-related negativity (ERN), a component of the event-related brain potential elicited when human participants commit errors in reaction-time tasks The authors propose that the ERN is generated when a negative reinforcement learning signal is conveyed to the anterior cingulate cortex via the mesencephalic dopamine system and that this signal is used by the anterior cingulate cortex to modify performance on the task at hand They provide support for this proposal using both computational modeling and psychophysiological experimentation

/pdf/the-neural-basis-of-human-error-processing-reinforcement-246y7vy4ka.pdf

The neural basis of human error processing: Reinforcement learning, dopamine, and the error-related negativity.

Assessments of anterior cingulate cortex in experimental animals and humans have led to unifying theories of its structural organization and contributions to mammalian behaviour. The anterior cingulate cortex forms a large region around the rostrum of the corpus callosum that is termed the anterior executive region. This region has numerous projections into motor systems, however, since these projections originate from different parts of anterior cingulate cortex and because functional studies have shown that it does not have a uniform contribution to brain functions, the anterior executive region is further subdivided into ‘affect’ and ‘cognition’ components. The affect division includes areas 25, 33 and rostral area 24, and has extensive connections with the amygdala and periaqueductal grey, and parts of it project to autonomic brainstem motor nuclei. In addition to regulating autonomic and endocrine functions, it is involved in conditioned emotional learning, vocalizations associated with expressing internal states, assessments of motivational content and assigning emotional valence to internal and external stimuli, and maternal—infant interactions. The cognition division includes caudal areas 24' and 32', the cingulate motor areas in the cingulate sulcus and nociceptive cortex. The cingulate motor areas project to the spinal cord and red nucleus and have premotor functions, while the nociceptive area is engaged in both response selection and cognitively demanding information processing. The cingulate epilepsy syndrome provides important support of experimental animal and human functional imaging studies for the role of anterior cingulate cortex in movement, affect and social behaviours. Excessive cingulate activity in cases with seizures confirmed in anterior cingulate cortex with subdural electrode recordings, can impair consciousness, alter affective state and expression, and influence skeletomotor and autonomic activity. Interictally, patients with anterior cingulate cortex epilepsy often display psychopathic or sociopathic behaviours. In other clinical examples of elevated anterior cingulate cortex activity it may contribute to tics, obsessive—compulsive behaviours, and aberrent social behaviour. Conversely, reduced cingulate activity following infarcts or surgery can contribute to behavioural disorders including akinetic mutism, diminished self-awareness and depression, motor neglect and impaired motor initiation, reduced responses to pain, and aberrent social behaviour. The role of anterior cingulate cortex in pain responsiveness is suggested by cingulumotomy results and functional imaging studies during noxious somatic stimulation. The affect division of anterior cingulate cortex modulates autonomic activity and internal emotional responses, while the cognition division is engaged in response selection associated with skeletomotor activity and responses to noxious stimuli. Overall, anterior cingulate cortex appears to play a crucial role in initiation, motivation, and goal-directed behaviours. The anterior cingulate cortex is part of a larger matrix of structures that are engaged in similar functions. These structures form the rostral limbic system and include the amygdala, periaqueductal grey, ventral striatum, orbitofrontal and anterior insular cortices. The system formed by these interconnected areas assesses the motivational content of internal and external stimuli and regulates context-dependent behaviours.

Contributions of anterior cingulate cortex to behaviour.

Most natural actions are chosen voluntarily from many possible choices. An action is often chosen based on the reward that it is expected to produce. What kind of cellular activity in which area of the cerebral cortex is involved in selecting an action according to the expected reward value? Results of an analysis in monkeys of cellular activity during the performance of reward-based motor selection and the effects of chemical inactivation are presented. We suggest that cells in the rostral cingulate motor area, one of the higher order motor areas in the cortex, play a part in processing the reward information for motor selection.

https://science.sciencemag.org/content/sci/282/5392/1335.full.pdf

Role for Cingulate Motor Area Cells in Voluntary Movement Selection Based on Reward

To achieve a volitional goal, we need to execute multiple movements in a specific temporal order. After repetitive performance of a particular sequence of movements, we are able to memorize and execute the whole sequence without external guidance. Where and how in the brain do we store information necessary for the orderly performance of multiple movements? We have found a group of cells in the cerebral cortex of monkeys whose activity is exclusively related to a sequence of multiple movements performed in a particular order. Such cellular activity exists in the supplementary motor area, but not in the primary motor cortex. We propose that these cells contribute a signal about the order of forthcoming multiple movements, and are useful for planning and coding of several movements ahead.

Role for supplementary motor area cells in planning several movements ahead

To study how neurons in the medial motor areas participate in performing sequential multiple movements that are individually separated in time, we analyzed neuronal activity in the supplementary (SMA) and presupplementary (pre-SMA) motor areas. Monkeys were trained to perform three different movements separated by waiting times, in four or six different orders. Initially each series of movements was learned during five trials guided by visual signals that indicated the correct movements. The monkeys subsequently executed the three movements in the memorized order without the visual signals. Three types of neuronal activity were of particular interest; these appeared to be crucially involved in sequencing the multiple motor tasks in different orders. First, we found activity changes that were selective for a particular sequence of the three movements that the monkeys were prepared to perform. The sequence-selective activity ceased when the monkeys initiated the first movement. Second, we found interval-selective activity that appeared in the interval between one particular movement and the next. Third, we found neuronal activity representing the rank order of three movements arranged chronologically; that is, the activity differed selectively in the process of preparing the first, second, or third movements in individual trials. The interval-selective activity was more prevalent in the SMA, whereas the rank-order selective activity was more frequently recorded in the pre-SMA. These results suggest how neurons in the SMA and pre-SMA are involved in sequencing multiple movements over time.

Neuronal activity in the supplementary and presupplementary motor areas for temporal organization of multiple movements.

1. Single-unit activity in the cingulate cortex of the monkey was recorded during the performance of sensorially (visual, auditory, or tactile) triggered or self-paced forelimb key press movements. 2. Microelectrodes were inserted into the broad rostrocaudal expanse of the cingulate cortex, including the upper and lower banks of the cingulate sulcus and the hemispheric medial wall of the cingulate gyrus. 3. A total of 1,042 task-related neurons were examined, the majority of which were related to the execution of the key press movements. In greater than 60% of them, the movement-related activity preceded the activity in the distal flexor muscles. 4. The movement-related neurons were distributed, in two foci, in the posterior and anterior parts of the cingulate cortex, both including the upper and lower banks of the cingulate sulcus. The posterior focus was found to largely overlap the area projecting to the forelimb area of the primary motor cortex by the use of the horseradish peroxidase (HRP) method. 5. About 40% of the cingulate cortical neurons showed equimagnitude responses during the signal-triggered and self-paced movements. The neurons exhibiting a selective or differential response to the self-paced motor task were more frequently observed in the anterior than in the posterior cingulate cortex. 6. The long-lead type of changes in activity, ranging from 500 ms to 2 s, were observed mainly before the self-paced and, much less frequently, before the triggered movements. They were particularly abundant in the anterior cingulate cortex. 7. Only a few of the neurons showed activity time-locked to the onset of the sensory signals. 8. These observations indicate that the anterior and posterior parts of the cingulate cortex are distinct entities participating in the performance of limb movements, even if the movements are simple, such as those in this study.

Two movement-related foci in the primate cingulate cortex observed in signal-triggered and self-paced forelimb movements

Interval timing is an essential guiding force of behavior. Previous reports have implicated the prefrontal and parietal cortex as being involved in time perception and in temporal decision making. We found that neurons in the medial motor areas, in particular the presupplementary motor area, participate in interval timing in the range of seconds. Monkeys were trained to perform an interval-generation task that required them to determine waiting periods of three different durations. Neuronal activity contributed to the process of retrieving time instructions from visual cues, signaled the initiation of action in a time-selective manner, and developed activity to represent the passage of time. These results specify how medial motor areas take part in initiating actions on the basis of self-generated time estimates.

Keisetsu Shima

Papers

Role for Cingulate Motor Area Cells in Voluntary Movement Selection Based on Reward

Role for supplementary motor area cells in planning several movements ahead

Neuronal activity in the supplementary and presupplementary motor areas for temporal organization of multiple movements.

Two movement-related foci in the primate cingulate cortex observed in signal-triggered and self-paced forelimb movements

Interval time coding by neurons in the presupplementary and supplementary motor areas.