Right BA 10 lesions impair performance on real-world planning but are not sensitive to problem novelty or tower tasks.

The prefrontal cortex (PFC) is associated with many cognitive functions, including planning. In the neuropsychology literature planning is reduced to "look ahead" ability and most extensively studied with the "tower" tasks. The most influential theoretical explanation is that planning is required in the absence of a routine solution and PFC patients have difficulty coping with novelty. There is an alternate view of planning that emphasizes the distinction between real world tasks and laboratory tower tasks. This account focuses on the structure of problem spaces and why patients with lesions to right PFC have difficulty navigating ill-structured problem spaces. To further explore these issues we administered two real world travel planning tasks to 56 Vietnam War veterans with penetrating brain lesions and 14 matched normal controls. One planning task involved familiar knowledge while the other involved knowledge unfamiliar to our participants. Participants also completed the D-KEFS tower task. A subset of 18 patients-with lesions to right anterior prefrontal cortex (BA 10)-were impaired in the travel planning task compared to normal controls. The task familiarity/novelty dimension affected performance across participant groups (familiar-task scores were higher than unfamiliar-task scores), but it did not differentially affect any group. An examination of cognitive strategies utilized by participants revealed that the impaired patient group had difficulty maintaining a sufficient level of abstraction and engaged the task at a much more concrete level than other participants. Interestingly, patients impaired in the real-world planning tasks were not impaired in the tower tasks. We conclude that patients with lesions to right BA 10 have difficulty in real-world planning tasks that can be attributed to difficulties in engaging problems at the appropriate level of abstraction.

Perceived danger associated with a property modulates cross category generalization.

Identifying projectable predicates is a key issue in understanding inductive inference. It is proposed that looking into the evolutionary psychology literature for adaptive properties may be one useful approach. One hypothesis that emerges from this literature is that properties that signal danger or harm should be more salient than properties that do not. Two studies are carried out to test this hypothesis. In study 1 participants were presented with a scenario involving the discovery of novel animals, for which there was incomplete information. Three types of properties (a harmful property, a neutral property, a beneficial property) were associated with animals in one (base) category and participants were asked to indicate strength of generalization of the property to a target within the category, and to a target across to another category. In the second experiment, the procedure was repeated, but in addition, subjects were also explicitly asked to indicate whether the base and target belonged to the same or different categories. Study 1 showed that the harmful property was more projectable compared to the beneficial and neutral properties. Study 2 reconfirmed this and further showed that it also promoted excessive generalization across categories. The results suggest that examination of adaptations identified by evolutionary psychologists may be a useful source of insight in the study of inductive inference. Supplementary Information: The online version contains supplementary material available at 10.1007/s11571-022-09793-3.

Hemispheric asymmetry in the prefrontal cortex for complex cognition.

With the exception of language, hemispheric asymmetry has not historically been an important issue in the frontal lobe literature. Data generated over the past 20 years is forcing a reconsideration of this position. There is now considerable evidence to suggest that the left prefrontal cortex is an inference engine that automatically makes simple conceptual, logical, and causal connections to fill in missing information and eliminate uncertainty or indeterminacy. This is a fine-tuning of the "left hemisphere interpreter" account from the callosotomy patient literature. What is new is an understanding of the important contributions of the right prefrontal cortex to formal logical inference, conflict detection, and indeterminacy tolerance and maintenance. This chapter articulates these claims and reviews the data on which they are based. The chapter concludes by speculating that the inference capabilities of the left prefrontal cortex are built into the very fabric of language and can be accounted for by the left hemisphere dominance for language. The roles of the right PFC require multiple mechanisms for explanation. Its role in formal inference may be a function of its visual-spatial processing capabilities. Its role in conflict detection may be explained as a system for checking for consistency between existing beliefs and new information coming into the system and inferences drawn from beliefs and/or new information. There are at least three possible mechanisms to account for its role in indeterminacy tolerance. First, it could contain a representational system with properties very different from those of language, and an accompanying inference engine. Second, it could just contain this different representational system, and the information is at some point passed back to the left prefrontal cortex for inference. Third, the role of the right prefrontal cortex may be largely preventative. That is, it doesn't provide alternative representational and inference capabilities but simply prevents the left prefrontal cortex from settling on initial, local inferences. The current data do not allow differentiating between these possibilities. Successful real-world functioning requires the participation of both hemispheres.

Left Amygdala and Putamen Activation Modulate Emotion Driven Decisions in the Iterated Prisoner's Dilemma Game.

Although economic decision-making is commonly characterized as a purely rational phenomenon, it is clear that real-world decision-making is influenced by emotions. Yet, relatively little is known about the neural correlates of this process. To explore this issue, 20 participants underwent fMRI scanning while engaged in the Prisoner's Dilemma game under partner-directed sympathy, anger and neutral emotion conditions. Participants were most and least likely to cooperate after sympathy and anger induction, respectively, with the neutral condition eliciting intermediate cooperation rates. Moreover, the sympathy condition elicited quicker responses for cooperation than defection choices, whereas this pattern was reversed in the anger and neutral conditions. Left amygdala activation showed a positive correlation with cooperation rates and self-reports of partner directed sympathy in the sympathy condition. In the anger condition, left putamen activation was positively correlated with cooperation rates and negatively correlated with self-reports of partner directed anger strength. These findings indicate that while the left amygdala activation may be indicative of emotion enhancement and increase of cooperative behavior, the left putamen may help to suppress an emotion to overcome anger and engage in cooperation.

Patients with Lesions to Left Prefrontal Cortex (BA 9 and BA 10) Have Less Entrenched Beliefs and Are More Skeptical Reasoners.

The effect of prior beliefs on reasoning and decision-making is a robust, poorly understood phenomenon, exhibiting considerable individual variation. Neuroimaging studies widely show the involvement of the left pFC in reasoning involving beliefs. However, little patient data exist to speak to the necessity and role of the left pFC in belief-based inference. To address this shortcoming, we tested 102 patients with unilateral focal penetrating traumatic brain injuries and 49 matched controls. Participants provided plausibility ratings (plausible/implausible) to simple inductive arguments and (separately) strength of believability ratings of the conclusion to those same arguments. A voxel-based lesion symptom mapping analysis identified 10 patients, all with lesions to the left pFC (BA 9 and BA 10) as rating significantly fewer arguments with highly believable conclusions as "plausible," compared with all other patients. Subsequent analyses, incorporating the right hemisphere homologue of these patients (n = 12) and normal controls (n = 24), revealed patients with lesions to left pFC found fewer arguments plausible in the high believable than either of these groups, and there was no difference in the behavioral scores of the right pFC patients and normal controls. Further analysis, utilizing the belief ratings as the dependent measure, revealed a Group × Belief Rating interaction, with left pFC patients having less intense beliefs about the conclusions of moderately believable and highly believable arguments. We interpreted these results to indicate that lesions to left pFC (BA 9, BA 10) increase incredulity and make these patients more skeptical reasoners. The former can partially, but not fully, explain the latter. The other relevant factor may be that unilateral left pFC lesions disrupt hemispheric equilibrium and allow for an increased inhibitory role of the right pFC. We speculate that individual differences in belief bias in reasoning in the normal population may be a function of individual differences in the left and right pFC interactional dynamics.

Developmental grey matter changes in superior parietal cortex accompany improved transitive reasoning.

The neural basis of developmental changes in transitive reasoning in parietal regions was examined, using voxel-based morphometry. Young adolescents and adults performed a transitive reasoning task, subsequent to undergoing anatomical magnetic resonance imaging (MRI) brain scans. Behaviorally, adults reasoned more accurately than did the young adolescents. Neural results showed (i) less grey matter density in superior parietal cortex in the adults than in the young adolescents, possibly due to a developmental period of synaptic pruning; (ii) improved performance in the reasoning task was negatively correlated with grey matter density in superior parietal cortex in the adolescents, but not in the adult group; and (iii) the latter results were driven by the more difficult trials, requiring greater spatial manipulation. Taken together, the results support the idea that during development, regions in superior parietal cortex are fine-tuned, to support more robust spatial manipulation, resulting in greater accuracy and efficiency in transitive reasoning.

Differential roles of polar orbital prefrontal cortex and parietal lobes in logical reasoning with neutral and negative emotional content.

To answer the question of how brain pathology affects reasoning about negative emotional content, we administered a disjunctive logical reasoning task involving arguments with neutral content (e.g. Either there are tigers or women in NYC, but not both; There are no tigers in NYC; There are women in NYC) and emotionally laden content (e.g. Either there are pedophiles or politicians in Texas, but not both; There are politicians in Texas; There are no pedophiles in Texas) to 92 neurological patients with focal lesions to various parts of the brain. A Voxel Lesion Symptom Mapping (VLSM) analysis identified 16 patients, all with lesions to the orbital polar prefrontal cortex (BA 10 & 11), as being selectively impaired in the emotional reasoning condition. Another 17 patients, all with lesions to the parietal cortex, were identified as being impaired in the neutral content condition. The reasoning scores of these two patient groups, along with 23 matched normal controls, underwent additional analysis to explore the effect of belief bias. This analysis revealed that the differences identified above were largely driven by trials where there was an incongruency between the believability of the conclusion and the validity of the argument (i.e. valid argument/false conclusion or invalid argument/true conclusion). Patients with lesions to polar orbital prefrontal cortex underperformed in incongruent emotional content trials and over performed in incongruent neutral content trials (compared to both normal controls and patients with parietal lobe lesions). Patients with lesions to parietal lobes underperformed normal controls (at a trend level) in neutral trials where there was a congruency between the believability of the conclusion and the validity of the argument (i.e. valid argument/true conclusion or invalid argument/false conclusion). We conclude that lesions to the polar orbital prefrontal cortex (i) prevent these patients from enjoying any emotionally induced cognitive boost, and (ii) block the belief bias processing route in the neutral condition. Lesions to parietal lobes result in a generalized impairment in logical reasoning with neutral content.

Lesions to polar/orbital prefrontal cortex selectively impair reasoning about emotional material.

While it is widely accepted that lesions to orbital prefrontal cortex lead to emotion related disruptions and poor decision-making, there is very little patient data on this issue involving actual logical reasoning tasks. We tested patients with circumscribed, focal lesions largely confined to polar/orbital prefrontal cortex (BA 10 & 11) (N=17) on logical reasoning tasks involving neutral and emotional content, and compared their performance to that of an age and education-matched normal control group (N=22) and a posterior lesion control group (N=24). Our results revealed a significant group by content interaction driven by a selective impairment in the polar/orbital prefrontal cortex group compared to healthy normal controls and to the parietal patient group, in the emotional content reasoning trials. Subsequent analyses of congruent and incongruent reasoning trials indicated that this impairment was driven by the poor performance of patients with polar/orbital lesions in the incongruent trials. We conclude that the polar/orbital prefrontal cortex plays a critical role in filtering emotionally charged content from the material before it is passed on to the reasoning system in lateral/dorsal regions of prefrontal cortex. Where unfiltered content is passed to the reasoning engine, either as a result of pathology (as in the case of our patients) or as a result of individual differences, reasoning performance suffers.

Indeterminacy tolerance as a basis of hemispheric asymmetry within prefrontal cortex.

There is an important hemispheric distinction in the functional organization of prefrontal cortex (PFC) that has not been fully recognized and explored. Research with split-brain patients provides considerable evidence for a left hemisphere (LH) "interpreter" that abhors indeterminacy and automatically draws inferences to complete patterns (real or imaginary). It is suggested that this "interpreter" function may be a byproduct of the linguistic capabilities of the LH. This same literature initially limited the role of the right hemisphere (RH) to little more than visual organization. Recent reviews have garnered evidence for several different roles for the right PFC in reasoning, problem solving, and decision-making. We here focus on the beneficial but neglected role of indeterminacy in real-world problem solving and argue that the right PFC complements the left PFC "interpreter" by maintaining, and even enhancing indeterminacy. Successful real-world functioning is a delicate balancing act between these two systems.

Syllogisms delivered in an angry voice lead to improved performance and engagement of a different neural system compared to neutral voice.

Despite the fact that most real-world reasoning occurs in some emotional context, very little is known about the underlying behavioral and neural implications of such context. To further understand the role of emotional context in logical reasoning we scanned 15 participants with fMRI while they engaged in logical reasoning about neutral syllogisms presented through the auditory channel in a sad, angry, or neutral tone of voice. Exposure to angry voice led to improved reasoning performance compared to exposure to sad and neutral voice. A likely explanation for this effect is that exposure to expressions of anger increases selective attention toward the relevant features of target stimuli, in this case the reasoning task. Supporting this interpretation, reasoning in the context of angry voice was accompanied by activation in the superior frontal gyrus-a region known to be associated with selective attention. Our findings contribute to a greater understanding of the neural processes that underlie reasoning in an emotional context by demonstrating that two emotional contexts, despite being of the same (negative) valence, have different effects on reasoning.

Different neural systems contribute to semantic bias and conflict detection in the inclusion fallacy task.

The inclusion fallacy is a phenomenon in which generalization from a specific premise category to a more general conclusion category is considered stronger than a generalization to a specific conclusion category nested within the more general set. Such inferences violate rational norms and are part of the reasoning fallacy literature that provides interesting tasks to explore cognitive and neural basis of reasoning. To explore the functional neuroanatomy of the inclusion fallacy, we used a 2 × 2 factorial design, with factors for quantification (explicit and implicit) and response (fallacious and non-fallacious). It was found that a left fronto-temporal system, along with a superior medial frontal system, was specifically activated in response to fallacious responses consistent with a semantic biasing of judgment explanation. A right fronto-parietal system was specifically recruited in response to detecting conflict associated with the heightened fallacy condition. These results are largely consistent with previous studies of reasoning fallacy and support a multiple systems model of reasoning.

Dissociable neural systems underwrite logical reasoning in the context of induced emotions with positive and negative valence.

How emotions influence syllogistic reasoning is not well understood. fMRI was employed to investigate the effects of induced positive or negative emotion on syllogistic reasoning. Specifically, on a trial-by-trial basis participants were exposed to a positive, negative, or neutral picture, immediately prior to engagement in a reasoning task. After viewing and rating the valence and intensity of each picture, participants indicated by keypress whether or not the conclusion of the syllogism followed logically from the premises. The content of all syllogisms was neutral, and the influence of belief-bias was controlled for in the study design. Emotion did not affect reasoning performance, although there was a trend in the expected direction based on accuracy rates for the positive (63%) and negative (64%) versus neutral (70%) condition. Nevertheless, exposure to positive and negative pictures led to dissociable patterns of neural activation during reasoning. Therefore, the neural basis of deductive reasoning differs as a function of the valence of the context.

Reason and less.

We consider ourselves to be rational beings. We feel that our choices, decisions, and actions are selected from a flexible array of possibilities, based upon reasons. When we vote for a political candidate, it is because they share our views on certain critical issues. When we hire an individual for a job, it is because they are the best qualified. However, if this is true, why does an analysis of the direction of shift in the timbre of the voice of political candidates during an exchange or debate, predict the winner of American presidential elections? Why is it that while only 3% of the American population consists of white men over 6'4″ tall, 30% of the CEOs of Fortune 500 companies are white men over 6'4″ tall? These are examples of "instinctual biases" affecting or modulating rational thought processes. I argue that existing theories of reasoning cannot substantively accommodate these ubiquitous, real-world phenomena. Failure to recognize and incorporate these types of phenomena into the study of human reasoning results in a distorted understanding of rationality. The goal of this article is to draw attention to these types of phenomena and propose an "adulterated rationality" account of reasoning as a first step in trying to explain them.

Creative brains: designing in the real world.

The process of designing artifacts is a creative activity. It is proposed that, at the cognitive level, one key to understanding design creativity is to understand the array of symbol systems designers utilize. These symbol systems range from being vague, imprecise, abstract, ambiguous, and indeterminate (like conceptual sketches), to being very precise, concrete, unambiguous, and determinate (like contract documents). The former types of symbol systems support associative processes that facilitate lateral (or divergent) transformations that broaden the problem space, while the latter types of symbol systems support inference processes facilitating vertical (or convergent) transformations that deepen of the problem space. The process of artifact design requires the judicious application of both lateral and vertical transformations. This leads to a dual mechanism model of design problem-solving comprising of an associative engine and an inference engine. It is further claimed that this dual mechanism model is supported by an interesting hemispheric dissociation in human prefrontal cortex. The associative engine and neural structures that support imprecise, ambiguous, abstract, indeterminate representations are lateralized in the right prefrontal cortex, while the inference engine and neural structures that support precise, unambiguous, determinant representations are lateralized in the left prefrontal cortex. At the brain level, successful design of artifacts requires a delicate balance between the two hemispheres of prefrontal cortex.

The effect of partner-directed emotion in social exchange decision-making.

Despite the prevalence of studies examining economic decision-making as a purely rational phenomenon, common sense suggests that emotions affect our decision-making particularly in a social context. To explore the influence of emotions on economic decision-making, we manipulated opponent-directed emotions prior to engaging participants in two social exchange decision-making games (the Trust Game and the Prisoner's Dilemma). Participants played both games with three different (fictional) partners and their tendency to defect was measured. Prior to playing each game, participants exchanged handwritten "essays" with their partners, and subsequently exchanged evaluations of each essay. The essays and evaluations, read by the participant, were designed to induce either anger, sympathy, or a neutral emotional response toward the confederate with whom they would then play the social exchange games. Galvanic skin conductance level (SCL) showed enhanced physiological arousal during anger induction compared to both the neutral and sympathy conditions. In both social exchange games, participants were most likely to defect against their partner after anger induction and least likely to defect after sympathy induction, with the neutral condition eliciting intermediate defection rates. This pattern was found to be strongest in participants exhibiting low cognitive control (as measured by a Go/no-Go task). The findings indicate that emotions felt toward another individual alter how one chooses to interact with them, and that this influence depends both on the specific emotion induced and the cognitive control of the individual.

Differential impact of beliefs on valence and arousal.

Many cognitive accounts of emotional processing assume that emotions have representational content that can be influenced by beliefs and desires. It is generally thought that emotions also have non-cognitive, affective components, including valence and arousal. To clarify the impact of cognition on these affective components we asked participants to rate sentences along cognitive and affective dimensions. For the former case, participants rated the believability of the material. For the latter case, they provided valence and arousal ratings. Across two experiments, we show that valence and arousal are differently influenced by beliefs, suggesting that these two largely independent affective components of emotion differ in their cognitive penetrability. While both components depended upon overall comprehension of sentence meaning, only valence was influenced by the consistency of the sentences with participants' beliefs (i.e., whether it was believable or unbelievable). We discuss the implications of these findings for understanding cognition-emotion relationships.

Transitive inference reasoning is impaired by focal lesions in parietal cortex rather than rostrolateral prefrontal cortex.

Transitive inference reasoning involves the examination and comparison of a given number of relational pairs in order to understand overall group hierarchy (e.g., A>B, B>C, C>D; therefore is A>D?). A number of imaging studies have demonstrated the role of the parietal cortex for resolving transitive inferences. Some studies also identify the rostrolateral prefrontal cortex as being critical for "relational integration" processes supporting transitive reasoning. To clarify this issue, we carried out a transitive inference study involving neurological patients with focal lesions to the rostrolateral prefrontal (n=5) or parietal cortices (n=7), as well as normal controls (n=6). The patients and controls were statistically matched on age, education, pre-injury IQ, general memory, working memory, and performance/full IQ, though the rostrolateral patients did score significantly higher than the normal controls on verbal IQ. Results indicate that patients with focal lesions to the parietal cortex were impaired in the task relative to both the patients with focal lesions to rostrolateral prefrontal cortex and the control group, and there was no difference in task performance between the rostrolateral prefrontal and the control groups. This result continued to hold after controlling for verbal IQ as a covariate. These findings point to a critical role for the parietal cortex, rather than the rostrolateral prefrontal, in transitive inference. Since the groups performed similarly on a working memory task, working memory cannot fully account for the result, suggesting a specific role of parietal cortex in transitive inference.

Lesions to right prefrontal cortex impair real-world planning through premature commitments.

While it is well accepted that the left prefrontal cortex plays a critical role in planning and problem-solving tasks, very little is known about the role of the right prefrontal cortex. We addressed this issue by testing five neurological patients with focal lesions to right prefrontal cortex on a real-world travel planning task, and compared their performance with the performance of five neurological patients with focal lesions to left prefrontal cortex, five neurological patients with posterior lesions, and five normal controls. Only patients with lesions to right prefrontal cortex generated substandard solutions compared to normal controls. Examination of the underlying cognitive processes and strategies revealed that patients with lesions to right prefrontal cortex approached the task at an excessively precise, concrete level compared to normal controls, and very early locked themselves into substandard solutions relative to the comparison group. In contrast, the behavior of normal controls was characterized by a judicious interplay of concrete and abstract levels/modes of representations. We suggest that damage to the right prefrontal system impairs the encoding and processing of more abstract and vague representations that facilitate lateral transformations, resulting in premature commitment to precise concrete patterns, and hasty albeit substandard conclusions (because the space of possibilities has not been properly explored).