iEEG implantation, behavioral task and computational modeling.

(A) Anatomical location of intracerebral electrodes across the 17 epileptic patients. Anterior insula (aINS, n=75), dorsolateral prefrontal cortex (dlPFC, n=70), lateral orbitofrontal cortex (lOFC, n=59), ventromedial prefrontal cortex (vmPFC, n=44), (B) Number of pairwise connectivity links (i.e. within patients) within and across regions, (C) Example of a typical trial in the reward (top) and punishment (bottom) conditions. Participants had to select one abstract visual cue among the two presented on each side of a central visual fixation cross and subsequently observed the outcome. Duration is given in milliseconds, (D) Number of trials where participants received outcomes +1€ (142±44, mean±std) vs. 0€ (93±33) in the rewarding condition (blue) and outcomes 0€ (141±42) vs. -1€ (93±27) in the punishment condition (red), (E) Across participants trial-wise reward PE (RPE - blue) and punishment PE (PPE - red), ± 95% confidence interval.

Local probabilistic encoding of reward and punishment prediction error signals.

(A) Time-courses of mutual information (MI in bits) estimated between the gamma power and the reward (blue) and punishment (red) PE signals. The solid line and the shaded area represent the mean and SEM of the across-contacts MI. Significant clusters of MI at the group level are plotted with horizontal bold lines (p<0.05, cluster-based correction, non-parametric randomization across epochs), (B) Instantaneous proportions of task-irrelevant (gray) and task-relevant bipolar derivations presenting a significant relation with either the RPE (blue), the PPE (red) or with both RPE and PPE (purple). Data are aligned to the outcome presentation (vertical line at 0 seconds).

Encoding of PE signals emerges from redundancy-dominated subsystems.

Dynamic interaction information (II in bits) within- (A) and between-regions (B) about the RPE (IIRPE) and PPE (IIPPE) are plotted in blue and red. Significant clusters of IIRPE and IIPPE are displayed with horizontal bold blue and red lines (p<0.05, cluster-based correction, non-parametric randomization across epochs). Significant differences between IIRPE and IIPPE are displayed in green. Shaded areas represent the SEM. The vertical gray line at 0 seconds represents the outcome presentation.

Contextual modulation of information transfer.

Time courses of transfer entropy (TE, in bits) from the aINS to the dlPFC (aINS→dlPFC) and from the vmPFC to the lOFC (vmPFC→lOFC), estimated during the rewarding condition (TERew in blue) and punishing condition (TEPun in red). Significant differences (p<0.05, cluster-based correction, non-parametric randomization across epochs) of TE between conditions are displayed with horizontal bold lines (blue for TERew > TEPun and red for TEPun > TERew). Shaded areas represent the SEM. The vertical gray line at 0 seconds represents the outcome presentation.

Synergistic interactions about the full PE signals between recordings of the dlPFC and vmPFC.

(A) Dynamic interaction information (II in bits) between the dlPFC and vmPFC about the full prediction error (IIdlPFC-vmPFC). Hot and cold colors indicate synergy- and redundancy-dominated II about the full PE. Significant clusters of II are displayed with a horizontal bold green line (p<0.05, cluster-based correction, non-parametric randomization across epochs). Shaded areas represent the SEM. The vertical gray line at 0 seconds represents the outcome presentation. (B) Dynamic IIdlPFC-vmPFC binned according to the local specificity PPE-RPE (IIPPE-RPE in pink) or mixed (IIMixed in purple) (C) Distributions of the mean of the IIPPE-RPE and IIMixed for each pair of recordings (IIPPE-RPE: one sample t-test against 0; dof=34; P fdr-corrected=0.015*; T=2.86; CI(95%)=[6.5e-5, 3.9e-4]; IIMixed: dof=33; P fdr-corrected=0.015*; T=2.84; CI(95%)=[5.4e-5, 3.3e-4]).

Summary of findings.

The four nodes represent the investigated regions, namely the anterior insula (aINS), the dorsolateral and ventromedial parts of the prefrontal cortex (dlPFC and vmPFC) and the lateral orbitofrontal cortex (lOFC). The outer disc represents the local probabilistic encoding i.e. the different proportions of contacts over time having a significant relationship between the gamma power and PE signals. In blue, is the proportion of contacts with a significant relation with the PE across rewarding trials (RPE-specific). Conversely, in red for punishment trials (PPE-specific). In purple, the proportion of contacts with a significant relationship with both the RPE and PPE. In gray, is the remaining proportion of non-significant contacts. Regarding interactions, we found that information transfer between aINS and dlPFC carried redundant information about PPE only and information transfer between vmPFC and lOFC about RPE only. This information transfer occurred with a leading role of the aINS in the punishment context and the vmPFC in the rewarding context. Finally, we found synergistic interactions between the dlPFC and the vmPFC about the full PE, without splitting into rewarding and punishing conditions.

Results of the one sample t-test performed against 0

Local encoding of prediction error signals within the gamma band.

(A) Distribution of information in the aINS, dlPFC, lOFC and vmPFC about the R/PPE in the frequency domain. The solid line and the shaded area respectively represent the mean and SEM of the across-contacts MI. Horizontal thick lines represent significant clusters of information (p<0.05, cluster-based correction, non-parametric randomization across epochs), (B) Density of information in the [50, 100]Hz, [100, 150]Hz and [150, 200]Hz bins.

Optimal delay interval for maximizing information transfer.

Modulation of transfer entropy (TE in bits) as a function of the delay between source and target areas. In blue, the TE computed across rewarding trials and in red, the TE computed across punishing trials. Shaded areas surrounding the time courses represent the 95% confidence interval estimated using a bootstrapping strategy.

Contextual modulation of the information transfer.

Time courses of transfer entropy (TE, in bits) estimated during the rewarding condition (TERew in blue) and punishing condition (TEPun in red). Significant differences (p<0.05, cluster-based correction, non-parametric randomization across epochs) of TE between conditions are displayed with horizontal bold lines (blue for TERew > TEPun and red for TEPun > TERew). Shaded areas represent the SEM. The vertical grey line at 0 seconds represents the outcome presentation.

Cortico-cortical interactions about the full PE signals.

Dynamic interaction information (II in bits) between-regions about the full prediction error (IIPE). Hot and cold colors indicate synergy- and redundancy-dominated interactions about the full PE. Significant clusters of IIPE are displayed with a horizontal bold green line (p<0.05, cluster-based correction, non-parametric randomization across epochs). Shaded areas represent the SEM. The vertical gray line at 0 seconds represents the outcome presentation.

Interaction information binned according to the local specificity.

(A) Dynamic interaction information (II in bits) between the dlPFC and vmPFC (IIdlPFC-vmPFC) binned according to the local specificity toward the RPE and PPE. Shaded areas represent the SEM. The vertical gray line at 0 seconds represents the outcome presentation. (B) Mean II between time points from 250 to 600ms after outcome presentation per category of local specificity. Each individual point represents one pairs of recordings from the dlPFC and vmPFC.