Neural Representations of Hierarchical Rule Sets: The Human Control System Represents Rules Irrespective of the Hierarchical Level to Which They Belong

Humans use rules to organize their actions to achieve specific goals. Although simple rules that link a sensory stimulus to one response may suffice in some situations, often, the application of multiple, hierarchically organized rules is required. Recent theories suggest that progressively higher level rules are encoded along an anterior-to-posterior gradient within PFC. Although some evidence supports the existence of such a functional gradient, other studies argue for a lesser degree of specialization within PFC. We used fMRI to investigate whether rules at different hierarchical levels are represented at distinct locations in the brain or encoded by a single system. Thirty-seven male and female participants represented and applied hierarchical rule sets containing one lower-level stimulus–response rule and one higher-level selection rule. We used multivariate pattern analysis to investigate directly the representation of rules at each hierarchical level in absence of information about rules from other levels or other task-related information, thus providing a clear identification of low- and high-level rule representations. We could decode low- and high-level rules from local patterns of brain activity within a wide frontoparietal network. However, no significant difference existed between regions encoding representations of rules from both levels except for precentral gyrus, which represented only low-level rule information. Our findings show that the brain represents conditional rules regardless of their level in the explored hierarchy, so the human control system did not organize task representation according to this dimension. Our paradigm represents a promising approach to identifying critical principles that shape this control system.

SIGNIFICANCE STATEMENT Several recent studies investigating the organization of the human control system propose that rules at different control levels are organized along an anterior-to-posterior gradient within PFC. In this study, we used multivariate pattern analysis to explore independently the representation of formally identical conditional rules belonging to different levels of a cognitive hierarchy and provide for the first time a clear identification of low- and high-level rule representations. We found no major spatial differences between regions encoding rules from different hierarchical levels. This suggests that the human brain does not use levels in the investigated hierarchy as a topographical organization principle to represent rules controlling our behavior. Our paradigm represents a promising approach to identifying which principles are critical.