Growing evidence suggests that the prefrontal cortex (PFC) is organized hierarchically, with more anterior regions having increasingly abstract representations. How does this organization support hierarchical cognitive control and the rapid discovery of abstract action rules? We present computational models at different levels of description. A neural circuit model simulates interacting corticostriatal circuits organized hierarchically. In each circuit the basal ganglia (BG) gate frontal actions, with some striatal units gating the inputs to PFC, and others gating the outputs to influence response selection. Learning at all of these levels is accomplished via dopaminergic reward prediction error signals in each corticostriatal circuit. This functionality allows the system to exhibit conditional if-then hypothesis testing and to learn rapidly in environments with hierarchical structure. We also develop a hybrid Bayesian-RL mixture of experts (MoE) model, which can estimate the most likely hypothesis state of individual participants based on their observed sequence of choices and rewards. This model yields accurate probabilistic estimates about which hypotheses are attended by manipulating attentional states in the generative neural model and recovering them with the MoE model. This two-pronged modeling approach leads to multiple quantitative predictions that are tested with fMRI in the companion paper.

Functional neuroimaging and lesion studies have frequently reported thalamic and putamen activation during reading and speech production. However, it is currently unknown how activity in these structures interacts with that in other reading and speech production areas. This study investigates how reading aloud modulates the neuronal interactions between visual recognition and articulatory areas, when both the putamen and thalamus are explicitly included. Using dynamic causal modeling in skilled readers who were reading regularly spelled English words, we compared 27 possible pathways that might connect the ventral anterior occipito-temporal sulcus (aOT) to articulatory areas in the precentral cortex (PrC). We focused on whether the neuronal interactions within these pathways were increased by reading relative to picture naming and other visual and articulatory control conditions. The results provide strong evidence that reading boosts the aOT--PrC pathway via the putamen but not the thalamus. However, the putamen pathway was not exclusive because there was also evidence for another reading pathway that did not involve either the putamen or the thalamus. We conclude that the putamen plays a special role in reading but this is likely to vary with individual reading preferences and strategies.

■ Appetitive goal-directed behavior can be associated with a cue-triggered expectancy that it will lead to a particular reward, a process thought to depend on the OFC and basolateral amygdala complex. We developed a biologically informed neural network model of this system to investigate the separable and complementary roles of these areas as the main components of a flexible expectancy system. These areas of interest are part of a neural network with additional subcortical areas, including the central nucleus of amygdala, ventral (limbic) and dorsomedial (associative) striatum. Our simulations are consistent with the view that the amygdala maintains Pavlovian associations through incremental updating of synaptic strength and that the OFC supports flexibility by maintaining an activation-based working memory of the recent reward history. Our model provides a mechanistic explanation for electrophysiological evidence that cue-related firing in OFC neurons is nonselectively early after a contingency change and why this nonselective firing is critical for promoting plasticity in the amygdala. This ambiguous activation results from the simultaneous maintenance of recent outcomes and obsolete Pavlovian contingencies in working memory. Furthermore, at the beginning of reversal, the OFC is critical for supporting responses that are no longer inappropriate. This result is inconsistent with an exclusive inhibitory account of OFC function. ■

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Due to basal ganglia dysfunction, bimanual motor performance in Parkinson patients reportedly relies on compensatory brain acti-vation in premotor–parietal–cerebellar circuitries. A subgroup of Par-kinson’s disease (PD) patients with freezing of gait (FOG) may exhibit greater bimanual impairments up to the point that motor blocks occur. This study investigated the neural mechanisms of upper limb motor blocks and explored their relation with FOG. Brain activation was measured using functional magnetic resonance imaging during bilateral finger movements in 16 PD with FOG, 16 without FOG (PD+ FOG and PD − FOG), and 16 controls. During suc-cessful movement, PD+ FOG showed decreased activation in right dorsolateral prefrontal cortex (PFC), left dorsal premotor cortex (PMd), as well as left M1 and bilaterally increased activation in dorsal putamen, pallidum, as well as subthalamic nucleus compared with PD − FOG and controls. On the contrary, upper limb motor blocks were associated with increased activation in right M1, PMd, supplementary motor area, and left PFC compared with successful movement, whereas bilateral pallidum and putamen activity was de-creased. Complex striatofrontal activation changes may be involved in the difficulties of PD+ FOG to perform bimanual movements, or sequential movements in general. These novel results suggest that, whatever the exact underlying cause, PD+ FOG seem to have reached a saturation point of normal neural compensation and respond belatedly to actual movement breakdown.

Winning against an opponent in a competitive video game can be ex-pected to be more rewarding than losing, especially when the opponent is a fellow human player rather than a computer. We show that winning versus losing in a first-person video game activates the brain’s reward circuit and the ventromedial prefrontal cortex (vmPFC) differ-ently depending on the type of the opponent. Participants played a com-petitive tank shooter game against alleged human and computer opponents while their brain activity was measured with functional magnetic resonance imaging. Brain responses to wins and losses were contrasted by fitting an event-related model to the hemodynamic data. Stronger activation to winning was observed in ventral and dorsal stria-tum as well as in vmPFC. Activation in ventral striatum was associated with participants ’ self-ratings of pleasure. During winning, ventral stria-tum showed stronger functional coupling with right insula, and weaker coupling with dorsal striatum, sensorimotor pre- and postcentral gyri,

Although characteristic motor symptoms of Parkinson’s disease such as bradykinesia typically improve under dopaminergic medica-tion, deficits in higher motor control are less responsive. We here investigated the dopaminergic modulation of network dynamics underlying basic motor performance, i.e. finger tapping, and higher motor control, i.e. internally and externally cued movement preparation and selection. Twelve patients, assessed ON and OFF medication, and 12 age-matched healthy subjects underwent func-tional magnetic resonance imaging. Dynamic causal modelling was used to assess effective connectivity in a motor network comprising cortical and subcortical regions. In particular, we investigated whether impairments in basic and higher motor control, and the effects induced by dopaminergic treatment are due to connectivity changes in (i) the mesial premotor loop comprising the supplementarymotor area; (ii) the lateral premotor loop comprising lateral premotor cortex; and (iii) cortico-subcortical interactions. At the behavioural level, we observed a marked slowing of movement preparation and selection when patients were internally as opposed to externally cued. Preserved performance during external cueing was associated with enhanced connectivity between prefrontal cortex and lateral pre-motor cortex OFF medication, compatible with a context-dependent compensatory role of the lateral premotor loop in the hypodo-paminergic state. Dopaminergic medication significantly improved finger tapping speed in patients, which correlated with a drug-induced coupling increase of prefrontal cortex with the supplementary motor area, i.e. the mesial premotor loop. In addition, only in the finger tapping condition, patients ON medication showed enhanced excitatory influences exerted by cortical premotor regions and the thalamus upon the putamen. In conclusion, the amelioration of bradykinesia by dopaminergic medication seems to be driven by

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Parcellation of the cingulate cortex at rest and during tasks: