Beyond risk and ambiguity: Deciding under ignorance

In this study, we examined the neural basis of decision making under different types of uncertainty that involve missing information: ambiguity (vague probabilities) and sample space ignorance (SSI; unknown outcomes). fMRI revealed that these two different types of uncertainty recruit distinct neural substrates: Ambiguity recruits the left insula, whereas SSI recruits the anterior cingulate cortex, bilateral inferior parietal cortex, and the lateral orbitofrontal cortex. The finding of unique activations for different types of uncertainty may not necessarily be predicted within the reductive approach of modern theories of decision making under uncertainty, because these theories purport that humans reduce more complicated uncertain environments to subjectively formed less complicated ones (i.e., SSI to ambiguity). The predictions of the reductive view held only for ambiguityaverse individuals and not for ambiguity-tolerant individuals. Consequently, theories of decision making under uncertainty should include individual tolerance for missing information and how these individual differences modulate the neural systems engaged during decision making. Supplemental materials for this article may be downloaded from http://cabn.psychonomic-journals.org/content/supplemental.     

Action sets and decisions in the medial frontal cortex

Activations in human dorsomedial frontal and cingulate cortices are often present in neuroimaging studies of decision making and action selection. Interpretations have emphasized executive control, movement sequencing, error detection and conflict monitoring. Recently, however, experimental approaches, using lesions, inactivation, and cell recording, have suggested that these are just components of the areas' functions. Here we review these results and integrate them with those from neuroimaging. A medial superior frontal gyrus (SFG) region centred on the pre-supplementary motor area (pre-SMA) is involved in the selection of action sets whereas the anterior cingulate cortex (ACC) has a fundamental role in relating actions to their consequences, both positive reinforcement outcomes and errors, and in guiding decisions about which actions are worth making.     

The contribution of distinct subregions of the ventromedial frontal cortex to emotion,social behavior,and decision making

Damage to the ventromedial frontal cortex (VMFC) in humans is associated with deficits in decision making. Decision making, however, often happens while people are interacting with others, where it is important to take the social consequences of a course of action into account. It is well known that VMFC lesions also lead to marked alterations in patients’ emotions and ability to interact socially; however, it has not been clear which parts of the VMFC are critical for these changes. Recently, there has been considerable interest in the role of the VMFC in choice behavior during interpersonal exchanges. Here, we highlight recent research that suggests that two areas within or adjacent to the VMFC, the orbitofrontal cortex (OFC) and the anterior cingulate cortex (ACC), may play distinct but complementary roles in mediating normal patterns of emotion and social behavior. Converging lines of evidence from human, macaque, and rat studies now suggest that the OFC may be more specialized for simple emotional responses, such as fear and aggression, through its role in representing primary reinforcement or punishment. By contrast, the ACC may play a distinct role in more complex aspects of emotion, such as social interaction, by virtue of its connections with the discrete parts of the temporal lobe and subcortical structures that control autonomic responses.     

Neurocomputational mechanisms of reinforcement-guided learning in humans: A review

Adapting decision making according to dynamic and probabilistic changes in action-reward contingencies is critical for survival in a competitive and resource-limited world. Much research has focused on elucidating the neural systems and computations that underlie how the brain identifies whether the consequences of actions are relatively good or bad. In contrast, less empirical research has focused on the mechanisms by which reinforcements might be used to guide decision making. Here, I review recent studies in which an attempt to bridge this gap has been made by characterizing how humans use reward information to guide and optimize decision making. Regions that have been implicated in reinforcement processing, including the striatum, orbitofrontal cortex, and anterior cingulate, also seem to mediate how reinforcements are used to adjust subsequent decision making. This research provides insights into why the brain devotes resources to evaluating reinforcements and suggests a direction for future research, from studying the mechanisms of reinforcement processing to studying the mechanisms of reinforcement learning.     

Controlling the integration of emotion and cognition: the role of frontal cortex in distinguishing helpful from hurtful emotional information

Emotion has been both lauded and vilified for its role in decision making. How are people able to ensure that helpful emotions guide decision making and irrelevant emotions are kept out of decision making? The orbitofrontal cortex has been identified as a neural area involved in incorporating emotion into decision making. Is this area's function specific to the integration of emotion and cognition, or does it more broadly govern whether emotional information should be integrated into cognition? The present research examined the role of orbitofrontal cortex when it was appropriate to control (i.e., prevent) the influence of emotion in decision making (Experiment 1) and to incorporate the influence of emotion in decision making (Experiment 2). Together, the two studies suggest that activity in lateral orbitofrontal cortex is associated with evaluating the contextual relevance of emotional information for decision making.     

Probing human and monkey anterior cingulate cortex in variable environments

Previous research has identified the anterior cingulate cortex (ACC) as an important node in the neural network underlying decision making in primates. Decision making can, however, be studied under a large variety of circumstances, ranging from the standard well-controlled lab situation to more natural, stochastic settings, in which multiple agents interact. Here, we illustrate how these different varieties of decision making studied can influence theories of ACC function in monkeys. Converging evidence from unit recordings and lesion studies now suggest that the ACC is important for interpreting outcome information according to the current task context to guide future action selection. We then apply this framework to the study of human ACC function and discuss its potential implications.     

Neurobiologische Korrelate der Psychotherapie von Zwangsst?rungen

Obsessive-compulsive disorders are associated with different neuropsychological impairments concerning executive functions, error monitoring, decision making and other basic functionalities. Contemporary neurocircuitry models emphasize the dysfunction of the fronto-striato-thalamo-cortical network related to parietal regions, the orbitofrontal cortex, the anterior cingulate cortex and other limbic structures. In brief, studies on the functional neuroanatomy of psychotherapy effects realized by positron emission tomography (PET) and functional magnetic resonance imaging (fMRI) are reported. Perspectives concern combined process-outcome research with repeated neuroimaging and highly resolved (daily) self-ratings focussing the nonlinear dynamics of psychotherapeutic change processes. New therapeutic approaches could explore the combination of psychotherapy with fMRI-based real-time neurofeedback and noninvasive neuromodulation with coordinated reset stimulation focusing on the desynchronization of pathologically oversynchronized neural populations and networks.     

Conflict monitoring and decision making: Reconciling two perspectives on anterior cingulate function

According to one influential account, the anterior cingulate cortex (ACC) serves to monitor for conflicts in information processing. According to another influential account, the ACC monitors action outcomes and guides decision making. Both of these perspectives are supported by an abundance of data, making it untenable to reject one view in favor of the other. Instead, the apparent challenge is to discover how the two perspectives might fit together within a larger account. In the present article, we consider the prospects for such a reconciliation. Juxtaposing the conflict-monitoring and decision-making accounts suggests an extension of the conflict-monitoring theory, by which conflict would act as a teaching signal driving a form of avoidance learning. The effect of this mechanism would be to bias behavioral decision making toward cognitively efficient tasks and strategies. We discuss evidence favoring this proposal and present an initial computational model, which lays the foundation for further development.     

The functional neuroanatomy of emotion and affective style

Recently, there has been a convergence in lesion and neuroimaging data in the identification of circuits underlying positive and negative emotion in the human brain. Emphasis is placed on the prefrontal cortex (PFC) and the amygdala as two key components of this circuitry. Emotion guides action and organizes behavior towards salient goals. To accomplish this, it is essential that the organism have a means of representing affect in the absence of immediate elicitors. It is proposed that the PFC plays a crucial role in affective working memory. The ventromedial sector of the PFC is most directly involved in the representation of elementary positive and negative emotional states while the dorsolateral PFC may be involved in the representation of the goal states towards which these elementary positive and negative states are directed. The amygdala has been consistently identified as playing a crucial role in both the perception of emotional cues and the production of emotional responses, with some evidence suggesting that it is particularly involved with fear-related negative affect. Individual differences in amygdala activation are implicated in dispositional affective styles and increased reactivity to negative incentives. The ventral striatum, anterior cingulate and insular cortex also provide unique contributions to emotional processing.     

Neural Correlates of Emotional Reactivity in Sensation Seeking

ABSTRACT— High sensation seeking has been linked to increased risk for drug abuse and other negative behavioral outcomes. This study explored the neurobiological basis of this personality trait using functional magnetic resonance imaging (fMRI). High sensation seekers (HSSs) and low sensation seekers (LSSs) viewed high- and low-arousal pictures. Comparison of the groups revealed that HSSs showed stronger fMRI responses to high-arousal stimuli in brain regions associated with arousal and reinforcement (right insula, posterior medial orbitofrontal cortex), whereas LSSs showed greater activation and earlier onset of fMRI responses to high-arousal stimuli in regions involved in emotional regulation (anterior medial orbitofrontal cortex, anterior cingulate). Furthermore, fMRI response in anterior medial orbitofrontal cortex and anterior cingulate was negatively correlated with urgency. Finally, LSSs showed greater sensitivity to the valence of the stimuli than did HSSs. These distinct neurobiological profiles suggest that HSSs exhibit neural responses consistent with an overactive approach system, whereas LSSs exhibit responses consistent with a stronger inhibitory system.     

Changes in cue-induced, prefrontal cortex activity with video-game play

Brain responses, particularly within the orbitofrontal and cingulate cortices, to Internet video-game cues in college students are similar to those observed in patients with substance dependence in response to the substance-related cues. In this study, we report changes in brain activity between baseline and following 6 weeks of Internet video-game play. We hypothesized that subjects with high levels of self-reported craving for Internet video-game play would be associated with increased activity in the prefrontal cortex, particularly the orbitofrontal and anterior cingulate cortex. Twenty-one healthy university students were recruited. At baseline and after a 6-week period of Internet video-game play, brain activity during presentation of video-game cues was assessed using 3T blood oxygen level dependent functional magnetic resonance imaging. Craving for Internet video-game play was assessed by self-report on a 7-point visual analogue scale following cue presentation. During a standardized 6-week video-game play period, brain activity in the anterior cingulate and orbitofrontal cortex of the excessive Internet game-playing group (EIGP) increased in response to Internet video-game cues. In contrast, activity observed in the general player group (GP) was not changed or decreased. In addition, the change of craving for Internet video games was positively correlated with the change in activity of the anterior cingulate in all subjects. These changes in frontal-lobe activity with extended video-game play may be similar to those observed during the early stages of addiction.     

Charting the maturation of the frontal lobe: an electrophysiological strategy

Tracking the functional development of specific regions of the prefrontal cortex in children using event-related potentials (ERPs) is challenging for both technical and conceptual reasons. In this paper we outline our strategy for studying frontal lobe development and present preliminary results from children aged 7-17 years and young adults using ERPs functionally associated with anterior cingulate and prefrontal cortex, especially the orbitofrontal, ventral, and medial portions. Our analysis of contingent negative variation, error-related negativity, and novelty P300 data show that the ERPs associated with these regions are still maturing into late adolescence, and that their amplitude has significant correlations with behavioral capacities.     

Face processing in different brain areas,and critical band masking

Neurophysiological evidence is described showing that some neurons in the macaque inferior temporal visual cortex have responses that are invariant with respect to the position, size, view, and spatial frequency of faces and objects, and that these neurons show rapid processing and rapid learning. Critical band spatial frequency masking is shown to be a property of these face‐selective neurons and of the human visual perception of faces. Which face or object is present is encoded using a distributed representation in which each neuron conveys independent information in its firing rate, with little information evident in the relative time of firing of different neurons. This ensemble encoding has the advantages of maximizing the information in the representation useful for discrimination between stimuli using a simple weighted sum of the neuronal firing by the receiving neurons, generalization, and graceful degradation. These invariant representations are ideally suited to provide the inputs to brain regions such as the orbitofrontal cortex and amygdala that learn the reinforcement associations of an individual's face, for then the learning, and the appropriate social and emotional responses generalize to other views of the same face. A theory is described of how such invariant representations may be produced by self‐organizing learning in a hierarchically organized set of visual cortical areas with convergent connectivity. The theory utilizes either temporal or spatial continuity with an associative synaptic modification rule. Another population of neurons in the cortex in the superior temporal sulcus encodes other aspects of faces such as face expression, eye‐gaze, face view, and whether the head is moving. These neurons thus provide important additional inputs to parts of the brain such as the orbitofrontal cortex and amygdala that are involved in social communication and emotional behaviour. Outputs of these systems reach the amygdala, in which face‐selective neurons are found, and also the orbitofrontal cortex, in which some neurons are tuned to face identity and others to face expression. In humans, activation of the orbitofrontal cortex is found when a change of face expression acts as a social signal that behaviour should change; and damage to the human orbitofrontal and pregenual cingulate cortex can impair face and voice expression identification, and also the reversal of emotional behaviour that normally occurs when reinforcers are reversed.     

Decision making in the Balloon Analogue Risk Task (BART): Anterior cingulate cortex signals loss aversion but not the infrequency of risky choices

The inferior frontal gyrus/anterior insula (IFG/AI) and anterior cingulate cortex (ACC) are key regions involved in risk appraisal during decision making, but accounts of how these regions contribute to decision making under risk remain contested. To help clarify the roles of these and other related regions, we used a modified version of the Balloon Analogue Risk Task (Lejuez et al., Journal of Experimental Psychology: Applied, 8, 75-84, 2002) to distinguish between decision-making and feedback-related processes when participants decided to pursue a gain as the probability of loss increased parametrically. Specifically, we set out to test whether the ACC and IFG/AI regions correspond to loss aversion at the time of decision making in a way that is not confounded with either reward-seeking or infrequency effects. When participants chose to discontinue inflating the balloon (win option), we observed greater ACC and mainly bilateral IFG/AI activity at the time of decision as the probability of explosion increased, consistent with increased loss aversion but inconsistent with an infrequency effect. In contrast, we found robust vmPFC activity when participants chose to continue inflating the balloon (risky option), consistent with reward seeking. However, in the cingulate and in mainly bilateral IFG regions, blood-oxygenation-level-dependent activation decreased when participants chose to inflate the balloon as the probability of explosion increased, findings that are consistent with a reduced loss aversion signal. Our results highlight the existence of distinct reward-seeking and loss-averse signals during decision making, as well as the importance of distinguishing between decision and feedback signals.     

The cognitive and neural basis of option generation and subsequent choice

Decision-making research has thoroughly investigated how people choose from a set of externally provided options. However, in ill-structured real-world environments, possible options for action are not defined by the situation but have to be generated by the agent. Here, we apply behavioral analysis (Study 1) and functional magnetic resonance imaging (Study 2) to investigate option generation and subsequent choice. For this purpose, we employ a new experimental task that requires participants to generate options for simple real-world scenarios and to subsequently decide among the generated options. Correlational analysis with a cognitive test battery suggests that retrieval of options from long-term memory is a relevant process during option generation. The results of the fMRI study demonstrate that option generation in simple real-world scenarios recruits the anterior prefrontal cortex. Furthermore, we show that choice behavior and its neural correlates differ between self-generated and externally provided options. Specifically, choice between self-generated options is associated with stronger recruitment of the dorsal anterior cingulate cortex. This impact of option generation on subsequent choice underlines the need for an expanded model of decision making to accommodate choice between self-generated options.     

Using fMRI to investigate

Using fMRI, we investigated the functional organization of prefrontal cortex (PFC) as participants briefly thought of a single just-experienced item (i.e., refreshed an active representation). The results of six studies, and a meta-analysis including previous studies, identified regions in left dorsolateral, anterior, and ventrolateral PFC associated in varying degrees with refreshing different types of information (visual and auditory words, drawings, patterns, people, places, or locations). In addition, activity increased in anterior cingulate with selection demands and in orbitofrontal cortex when a nonselected item was emotionally salient, consistent with a role for these areas in cognitive control (e.g., overcoming “mental rubbernecking”). We also found evidence that presenting emotional information disrupted an anterior component of the refresh circuit. We suggest that refreshing accounts for some neural activity observed in more complex tasks, such as working memory, long-term memory, and problem solving, and that its disruption (e.g., from aging or emotion) could have a broad impact.     

Integrating orbitofrontal cortex into prefrontal theory: common processing themes across species and subdivisions

Currently, many theories highlight either representational memory or rule representation as the hallmark of prefrontal function. Neurophysiological findings in the primate dorsolateral prefrontal cortex indicate that both features may characterize prefrontal processing. Neurons in the dorsolateral prefrontal cortex encode information in working memory, and this information is represented when relevant to the rules governing performance in a task. In this review, we discuss recent reports of encoding in primate and rat orbitofrontal regions indicating that these features also characterize activity in the orbitofrontal subdivision of the prefrontal cortex. These data indicate that (1) neural activity in the orbitofrontal cortex links the current incentive value of reinforcers to cues, rather than representing the physical features of cues or associated reinforcers; (2) this incentive-based information is represented in the orbitofrontal cortex when it is relevant to the rules guiding performance in a task; and (3) incentive information is also represented in the orbitofrontal cortex in working memory during delays when neither the cues nor reinforcers are present. Therefore, although the orbitofrontal cortex appears to be uniquely specialized to process incentive or motivational information, it may be integrated into a more global framework of prefrontal function characterized by representational encoding of performance-relevant information.     

Dopamine D1 receptors in the anterior cingulate cortex regulate effort-based decision making

The anterior cingulate cortex (ACC) has been implicated in encoding whether or not an action is worth performing in view of the expected benefit and the cost of performing the action. Dopamine input to the ACC may be critical for this form of effort-based decision making; however, the role of distinct ACC dopamine receptors is yet unknown. Therefore, we examined in rats the effects of an intra-ACC D1 and D2 receptor blockade on effort-based decision making tested in a T-maze cost-benefit task. In this task, subjects could either choose to climb a barrier to obtain a high reward in one arm or a low reward in the other arm without a barrier. Unlike vehicle-treated rats, rats with intra-ACC infusion of the D1 receptor antagonist SCH23390 exhibited a reduced preference for the high-cost- high-reward response option when having the choice to obtain a low reward with little effort. In contrast, in rats with intra-ACC infusion of the D2 receptor antagonist eticlopride, the preference for the high-cost-high-reward response option was not altered relative to vehicle-treated rats. These data provide the first evidence that D1 receptors in the ACC regulate effort-based decision making.     

Neurocognitive mechanisms of cognitive control: the role of prefrontal cortex in action selection, response inhibition, performance monitoring, and reward-based learning

Convergent evidence highlights the differential contributions of various regions of the prefrontal cortex in the service of cognitive control, but little is understood about how the brain determines and communicates the need to recruit cognitive control, and how such signals instigate the implementation of appropriate performance adjustments. Here we review recent progress from cognitive neuroscience in examining some of the main constituent processes of cognitive control as involved in dynamic decision making: goal-directed action selection, response activation and inhibition, performance monitoring, and reward-based learning. Medial frontal cortex is found to be involved in performance monitoring: evaluating outcome vis-a-vis expectancy, and detecting performance errors or conflicting response tendencies. Lateral and orbitofrontal divisions of prefrontal cortex are involved in subsequently implementing appropriate adjustments.     

Brain abnormalities in antisocial individuals: implications for the law

With the increasing popularity in the use of brain imaging on antisocial individuals, an increasing number of brain imaging studies have revealed structural and functional impairments in antisocial, psychopathic, and violent individuals. This review summarizes key findings from brain imaging studies on antisocial/aggressive behavior. Key regions commonly found to be impaired in antisocial populations include the prefrontal cortex (particularly orbitofrontal and dorsolateral prefrontal cortex), superior temporal gyrus, amygdala-hippocampal complex, and anterior cingulate cortex. Key functions of these regions are reviewed to provide a better understanding on how deficits in these regions may predispose to antisocial behavior. Objections to the use of imaging findings in a legal context are outlined, and alternative perspectives raised. It is argued that brain dysfunction is a risk factor for antisocial behavior and that it is likely that imaging will play an increasing (albeit limited) role in legal decision-making.