Disentangling the prefrontal network for rule selection by means of a non‐verbal variant of the Wisconsin Card Sorting Test

Abstract

This study disentangles the prefrontal network underlying executive functions involved in the Wisconsin Card Sorting Test (WCST). During the WCST, subjects have to perform two key processes: first, they have to derive the correct sorting rule for each trial by trial-and-error, and, second, they have to detect when this sorting rule is changed by the investigator. Both cognitive processes constitute key components of the executive system, which is subserved by the prefrontal cortex. For the current fMRI experiment, we developed a non-verbal variant of the WCST. Subjects were instructed either to respond according to a given sorting rule or to detect the correct sorting rule, like in the original version of the WCST. Data were obtained from 14 healthy male volunteers and analysed using SPM and a random effects model. All conditions activated a fronto-parietal network, which was generally more active when subjects had to search for the correct sorting rule than when the rule was announced beforehand. Significant differences between these two conditions were seen in the dorsolateral prefrontal cortex (PFC) and the parietal lobe. In addition, the data provided new evidence for the assumption of differentiated roles of the left and right prefrontal cortex. Although the right PFC showed a general involvement in response selection and the execution of goal directed responses, based on given rules, the left PFC was only activated when inductive reasoning and feedback integration was required.

Mechanisms of Rule Acquisition and Rule Following in Inductive Reasoning

“…This generalization is in line with previous research (Reverberi et al, 2005a,b;Specht et al, 2009), but differs in emphasis from conclusions from other studies, including those using neurophysiological recordings in nonhuman primates. Thus, although some studies (Wang et al, 2000;Bussey et al, 2001) have shown that lesioning ventral PFC after learning does not affect subse- quent rule use in monkeys, other studies, in which recording was from regions in the principal sulcus extending to DLPFC (BA 9/46), have supported the idea that this latter region controls the guidance of behavior according to well learned rules (White and Wise, 1999;Asaad et al, 2000;Mansouri et al, 2006).…”

“…Thus, some studies, including neurophysiological recordings in nonhuman primates (White and Wise, 1999;Goel and Dolan, 2000;Seger et al, 2000;Strange et al, 2001;Bunge, 2004) have attributed a role for the dorsolateral prefrontal cortex (DLPFC) (particularly the left DLPFC) in the application and maintenance of rules, rather than in their initial learning. Nonetheless, recent evidence suggests that DLPFC plays a role in hypothesis generation during rule learning independently of working memory (Boettiger and D'Esposito, 2005;Reverberi et al, 2005b;Specht et al, 2009).…”

Role for Presupplementary Motor Area in Inhibition of Cognitive Set Interference

“…It has been demonstrated that performance of the WCST is impaired following lesion to the pFC (Aron, Monsell, Sahakian, & Robbins, 2004;Stuss et al, 2000;Dias, Robbins, & Roberts, 1996Owen et al, 1993;Janowski, Shimamura, Kritchevski, & Squire, 1989;Nelson, 1976;Drewe, 1974;Passingham, 1972;Milner, 1963). Consistent with the neuropsychological studies, prominent activation as measured with neuroimaging has been detected in the pFC during performance of the WCST (Nyhus & Barcelo, 2009;Specht, Lie, Shah, & Fink, 2009;Konishi et al, 1998Konishi et al, , 2002Konishi et al, , 2008Hampshire & Owen, 2006;Lie, Specht, Marshall, & Fink, 2006;Monchi et al, 2004;Monchi, Petrides, Petre, Worsley, & Dagher, 2001;Rogers, Andrews, Grasby, Brooks, & Robbins, 2000;Nagahama et al, 1999) and the task switching paradigms (Wylie, Murray, Javitt, & Foxe, 2009;Crone, Donohue, Honomichl, Wendelken, & Bunge, 2006;Brass & von Cramon, 2004;Cools, Clark, & Robbins, 2004;Braver, Reynolds, & Donaldson, 2003;Dove, Pollmann, Schubert, Wiggins, & von Cramon, 2002;Rushworth, Passingham, & Nobre, 2002;Pollmann, Weidner, Muller, & von Cramon, 2000;Sohn, Ursu, Anderson, Stenger, & Carter, 2000). It is to be noted, however, that the brain activity measured at the time of the dimension/task changes may reflect both inhibition of PI and reconfiguration of a task new set (Monsell, 2003), and thus it is not clear whether the detected prefrontal activation at the time of the dimension/task changes is specific to inhibition...…”

“…Previous neuroimaging studies have examined set shifting/ task switching by contrasting dimension change minus dimension repeat/task switch minus task repeat (Nyhus & Barcelo, 2009;Specht et al, 2009;Wylie et al, 2009;Crone et al, 2006;Hampshire & Owen, 2006;Lie et al, 2006;Konishi et al, 2002Konishi et al, , 2005Brass & von Cramon, 2004;Cools et al, 2004;Monchi et al, 2001Monchi et al, , 2004Braver et al, 2003;Dove et al, 2002;Rushworth et al, 2002;Pollmann et al, 2000;Rogers et al, 2000;Sohn et al, 2000;Nagahama et al, 1999). Assuming that set shifting/task switching consists of inhibition of PI from a previous set and reconfiguration of a new set, the weak activation associated with inhibition, as opposed to reconfiguration, reported in the present study suggests that the major component of the lateral prefrontal activation during set shifting/task switching reported previously, including the inferior prefrontal activation, might be related to reconfiguration of a new task set.…”

Interindividual differences in cognitive flexibility: Influence of gray-matter volume, functional connectivity and trait impulsivity

“…Alternatively and especially in the context of cognitive control, the aINS might sometimes be mislabeled as inferior frontal cortex (Swick et al 2011). In addition, even though there are studies reporting the insula in executive functioning tasks (Lie et al 2006;Specht et al 2009), its specific role has in this context hardly been discussed. Nevertheless, there is growing evidence that the aINS plays a major role in cognitive and attentional control (Menon and Uddin 2010;Dosenbach et al 2006;Kurth et al 2010).…”