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Orbitofrontal cortex

Dr. Simon Moss

Overview

The orbitofrontal cortex is located at the front and sides of the brain, towards the bottom rather than top of prefrontal regions. More precisely, the orbitofrontal cortex entails Brodmann areas 10, 11, and 47 (Kringelbach, 2005). In general, this region enables individuals to adapt their behavior in response to unexpected rewards or adversities.

To illustrate, suppose individuals need to decide which of several courses of action to pursue, such as which house to purchase, which holiday to choose, or which comment to express. These individuals may imagine the various alternatives. These images are represented in various association regions of the cortex but transmitted to the orbitofrontal cortex. The orbitofrontal cortex then, in essence, calculates or represents the emotions that individuals would experience if they pursued each alternative. For example, if the features of one house would evoke a blend of excitement but disappointment, the orbitofrontal cortex transmits these emotions to other regions. These regions then evoke the bodily states, such as increased blood flow to the legs, that underpin these emotions. Thus, in essence, individuals experience or anticipate the emotions that each alternative will elicit (for more information about this premise, see the somatic marker hypothesis). These emotions then bias the decisions that individuals reach.

Accordingly, the orbitofrontal cortex enables individuals to anticipate whether some course of action will evoke positive or negative emotions, both immediately and in the future. In this sense, the orbitofrontal cortex also underpins moral behavior. That is, when the orbitofrontal cortex is activated, individuals choose the courses of action that tend to be rewarded, rather than punished, by other people.

Occasionally, the actions that individuals choose do not elicit the anticipated outcomes. That is, the emotions and bodily states that individuals actually experience after some action deviate from the expected emotions and bodily states. This discrepancy is then transmitted back to the orbitofrontal cortex to improve future predictions. For example, suppose a holiday is not as exciting as expected. The next time that individuals envisage a similar holiday, the level of excitement they anticipate will be diminished. In this sense, the orbitofrontal cortex facilitates the capacity of individuals to adapt to unexpected outcomes.

Because the orbitofrontal cortex evokes the emotions and bodily states that hypothetical activities would elicit, this region also facilitates empathy. That is, this region enables individuals to experience the emotions that someone else may feel in a specific setting. This empathy can facilitate social behavior. In contrast, when this region is inhibited or compromised, individuals may behave inappropriately. Indeed, damage to the orbitofrontal cortex may underpin some forms of psychopathy.

The orbitofrontal cortex is closely connected to many other regions. For example, because this region is closely connected to the limbic system, especially the amygdala, the orbitofrontal cortex is sometimes regarded as part of the expanded limbic system (Nauta, 1979). The orbitofrontal cortex is also closely connected to the dorsolateral prefrontal cortex. Dopamine projections from the nucleus accumbens and ventral tegmental area also project significantly to the orbitofrontal cortex

Evidence of the roles and functions of the orbitofrontal cortex

Iowa Gambling Task

The roles and functions of the orbitofrontal cortex were initially clarified by studies that utilized the Iowa Gambling Task (for more information, see measures of risky decision making). The Iowa Gambling Task is usually administered over computer (see Bechara, Damasio, Tranel, Damasio, 1997). Four decks of cards are presented, face down, labeled A, B, C, and D, each comprising 40 cards. Participants begin with a specific amount of money, such as $1000. This amount is specified on the screen, sometimes above the decks.

The participants then select a card from any of the four decks, often by clicking a mouse onto a specific deck. Each card can generate a reward, a penalty, or both. After a card is selected, this reward and penalty appears on the screen for several seconds. To illustrate, one card might generate a reward of $100, a penalty of $150, and thus a net loss of $50. Their overall amount would then shift from $1000 to $950. Participants then repeat this process many times, such as 100 trials.

Unbeknownst to participants, the probability of a reward and penalty as well as the distribution of these rewards and penalties varies across the decks. That is, two of the decks, over time, are more likely to incur losses than are the other two decks. Specifically, the two disadvantageous decks, on average, might lose $25. The two advantageous decks, on average might gain $25.

Nevertheless, the losses that each deck incurs vary across trials and, therefore, is difficult to decipher at first. For one of the advantageous decks, each card generates a gain of $50, and one in two cards incurs a loss of $50. For the other advantageous deck, each card generates a gain of $50, and one in ten cards incurs a loss of $250. For one of the disadvantageous decks, each card generates a gain of $100, and one in ten cards incurs a loss of $1250. Finally, for the other disadvantageous decks, each card generates a gain of $100, and one in every two cards incurs losses that range from of $150 to $350, although variations to these schemes can be included.

At the outset, most participants will choose the disadvantageous decks, because the gains are higher. Over time, usually within the first 40 trials, participants learn to choose the advantageous decks.

Studies have shown that participants with lesions in the ventromedial prefrontal cortex often do not learn to choose the advantageous decks (Bechara, Damasio, Tranel, & Anderson, 1994). Over time, with more precision in their instruments, researchers showed that impairments confined to a specific region of the ventromedial prefrontal cortex?-the orbitofrontal cortex?-is especially likely to compromise performance on this task (e.g., Bechara, 2004). Thus, the orbitofrontal cortex enables individuals to adapt their choices in response to unexpected punishments.

Anticipation of rewards and punishments

These studies also showed that lesions to the orbitofrontal cortex would manifest in physiological responses. That is, when this region is damaged, individuals are not as likely to exhibit the physiological responses, such as increased skin conductance, before potential punishments. In contrast, when this region is intact, individuals may exhibit signs of threat before they choose a disadvantageous pack (Bechara, Damasio, Tranel, & Anderson, 1994).

These studies imply that perhaps the orbitofrontal cortex, in essence, enables individuals to anticipate the emotions and bodily states of specific choices (Rolls, 2000). These individuals do not only anticipate the immediate outcome of a behavior, such as whether or not they will be criticized, but future outcomes as well. To illustrate, suppose some act may generate immediate pleasure but could also culminate in problems in the future. The orbitofrontal cortex, in essence, can integrate all these outcomes to represent an average or amalgam of these emotions.

Furthermore, because the orbitofrontal cortex integrates the possible emotions of many outcomes, choices will also depend on the likelihood of each outcome. For example, suppose one course of action, such as purchasing a house, is very likely to elicit moderate excitement. Nevertheless, suppose ths house may be haunted and thus, although unlikely, could also elicit considerable fear. Because the moderate excitement is more plausible, the orbitofrontal cortex will ensure that individuals primarily anticipate this emotion. This region, therefore, is especially important when individuals need to decide between likely but modest rewards and unlikely but sizeable rewards (see Rogers et al., 1999).

Unfulfilled expectations and reversal learning

To ensure that individuals anticipate emotions and bodily states accurately, the orbitofrontal cortex must be responsive to outcomes that deviate from expectations. Indeed, any unexpected outcomes are transmitted to the orbitofrontal cortex, which then adapts accordingly. If an event is not as enjoyable as predicted, the orbitofrontal cortex will ensure that individuals anticipate some problems with similar events in the future.

Accordingly, the orbitofrontal cortex enables individuals to adapt to unexpected information (Rolls, 2000& Schnider et al., 2005). Indeed, when this region is impaired, individuals do not adapt effectively. For example, suppose individuals learn that a specific action, such as pressing the right of two buttons, tends to attract rewards. These individuals may continue to press this button even after this option no longer attracts rewards (e.g., Nahum et al., 2009). That is, the orbitofrontal cortex enables individuals to extinguish or abstain from behaviors that are no longer viable (see also Boulougouris, V., & Robbins, 2009& Burke, Takahashi, Correll, Leon Brown, & Schoenbaum, 2009).

Both unexpected rewards and adverities activate the orbitofrontal cortex, as shown by PET (Schnider et al., 2005) as well as EEG studies (Schnider et al., 2007). In contrast, some regions are sensitive only to unexpected rewards or adversities, but not both.

One study, conducted by Nahum, Simon, Sander, Lazeyras, and Schnider (2011), represents an excellent example of this role and function of the orbitofrontal cortex. In this study, participants needed to undertake a reversal learning task. Specifically, on each trial, a pair of faces appeared, side by side. A couple of seconds later, a target, such as a spider, would appear on the nose of one face. Participants needed to press one of two buttons to predict the face on which this target would appear.

In general, the spider would appear on the same face for 4 to 6 trials. Thus, participants learnt to choose this face. However, at one point, the spider would shift to the other face for 4 to 6 trials. Furthermore, in one condition, instead of a spider, the target was merely a disk.

Whenever the target shifted, and thus deviated from the expectations of participants, the orbitofrontal cortex, especially the lateral region, was activated. This region was activated regardless of whether the target was a spider or disk. That is, the valence of this expectation did not affect the level of activation. Presumably, activation of the orbitofrontal cortex enables individuals to shift their behavior in response to unexpected outcomes.

In contrast, the anterior insular cortex was activated only when anticipated spiders, but not disks, appeared. Presumably, this region represents the anticipation of disgusting or threatening objects.

Counterfactual thinking

After individuals experience some outcome, they often imagine the consequences that could have ensued had another sequence of events unfolded, called counterfactual thinking. If they imagine a less favorable event, their mood often improves. If they imagine a more favorable event, they often derive key insights. As Ursu and Carter (2005) showed in a functional MRI study, the orbito-frontal cortex is especially likely to mediate counterfactual thinking. This finding is unsurprising, given the orbito-frontal cortex enables individuals to experience the emotions they would feel had they engaged in various hypothetical activities.

Theory of mind

To socialize effectively, individuals need to understand the perspectives and preferences of other people. They need to appreciate the feelings and accommodate the desires of another person. This capacity is called theory of mind. The orbitofrontal cortex facilitates this capacity (Stone et al., 1998).

Two distinct capacities have been differentiated: cognitive and affective (e.g., Shamay-Tsoory, Harari, Aharon-Peretz, & Levkovitz, 2010). Cognitive theory of mind represents the ability of individuals to appreciate the beliefs and assumptions of other people, similar to perspective taking. Affective theory of mind represents the capacity of individuals to appreciate the emotions and feelings of other people, similar to empathy. The orbitofrontal cortex seems to underpin affective theory of mind in particular (e.g., Shamay-Tsoory et al., 2004). That is, the orbitofrontal cortex enables individuals to experience the emotions that someone else may be feeling.

To assess these two facets of theory of mind, the Yoni task is often used (e.g., Kalbe et al., 2010& Shamay -Tsoory, Harari, Aharon-Peretz, & Levkovitz, 2010). Specifically, a schematic picture of a face, called Yoni, appears in the middle of a screen. The face may be smiling, sad, or neutral. The eyes may be pointing in various directions. A picture also appears in each corner. The picture may be an object, like a car, or an object and face, such as a strawberry next to a sad face. Finally, a sentence appears at the top, such as "Yoni is thinking of ---".

Participants must point to the corner that seems to correspond to the sentence. For example, suppose the sentence is "Yoni is thinking of ---", and Yoni's eyes are directed to the top left corner, the participant would point to this corner.

In some trials, the stimuli and sentences do not correspond to any emotions. For example, the sentence "Yoni is thinking of ---" does not allude to his feelings. Furthermore, Yoni is neither smiling nor frowning. Performance on these trials represents only cognitive theory of mind. On other trials, the stimuli and sentences do correspond to emotions. For example, the sentence may be "Yoni likes ---" and Yoni seems to be smiling and peering towards a particular corner. Performance on these trials represents affective theory of mind.

In addition, the difficulty of these trials varies. In some trials, participants do not only need to decipher the thoughts or emotions of Yoni. Instead, they also need to decipher the thoughts and emotions of other faces, located in the corners. To illustrate, in one trial Yoni might be smiling and orienting his eyes to the top left corner. In this corner, a face may be smiling as well and peering at a toy. The sentence might read "Yoni loves the toy that --- loves". Participants would need to point to the face in the top left corner.

Shamay-Tsoory, Harari, Aharon-Peretz, and Levkovitz (2010) administered this task to patients with lesions in the orbitofrontal cortex as well as a control group. When participants needed to decipher the emotions of both Yoni and other faces, lesions to orbitofrontal cortex delayed reaction times and thus compromised performance. However, these lesions did not compromise performance in the other conditions. These findings imply that perhaps the orbitofrontal cortex is pertinent to affective theory of mind, especially in more complex settings.

Furthermore, in this study, participants completed a multidimensional measure of empathy (Davis, 1983). Two of the subscales correspond to more cognitive forms of empathy, such as perspective taking and the fantasy scale, in which people imagine themselves from the perspective of other people. Two of the subscales correspond to more affective forms of empathy, such as empathic concern and personal distress in tense situations. Interestingly, only cognitive empathy was impaired when the orbitofrontal cortex was damaged.

Finally as Kalbe et al. (2010) showed when TMS is used to inhibit the dorsolateral prefrontal cortex, cognitive rather than affective theory of mind actually improves. According to Kalbe et al. (2010), when individuals need to decipher the beliefs of other people, the dorsolateral prefrontal cortex is usually activated. This system then inhibits the orbitofrontal cortex. Accordingly, affective cues are disregarded?=cues that might sometimes actually facilitate performance. When the dorsolateral prefrontal cortex is inhibited, this problem dissipates.

Psychopathy

Shamay-Tsoory, Harari, Aharon-Peretz, and Levkovitz (2010) argue that psychopathy could also represent impairments in affective, but not cognitive, theory of mind (for definitions and discussions of psychopathy, see measures of psychopathy). That is, people with psychopathy may not appreciate the emotions, anxieties, fears, and sorrow of another person, increasing the likelihood they will behave callously. Nevertheless, psychopaths are often skilled in social situations, implying they can readily decipher the assumptions and beliefs of other people. Accordingly, impairment to the orbitofrontal cortex could partly underpin psychopathy.

Many studies offer evidence that supports this possibility (e.g., Blair et al., 2006). As Shamay-Tsoory, Harari, Aharon-Peretz, and Levkovitz (2010) showed, affective but not cognitive theory of mind, especially in the difficult condition, is inversely related to measures of psychopathy. Indeed, individuals who exhibited the hallmarks of psychopathy seemed to show the same profile of results as individuals with lesions in the orbitofrontal cortex.

Furthermore, people with psychopathy exhibit deficits in the Iowa Gambling Task (Blair et al., 2001& Van Honk et al., 2002)-a deficit that often indicates dysfunction in the orbitofrontal cortex. Anderson, Bechara, Damasio, Tranel, and Damasio (1999) assume that impairments in these regions may be associated more with Factor 2 than Factor 1 psychopathy.

Moral decisions

When individuals need to consider a moral dilemma, such as whether to help or exploit another person, the lateral orbito-frontal cortex is activated in some, but not all people. Emonds, Declerck, Boone, Vandervliet, and Parizel (2011) used fMRI to distinguish two primary motivations of individuals to cooperate. That is, some people feel an inherent drive to cooperate, partly governed by a need to behave morally and to comply with social norms. Brain regions such as the lateral orbito-frontal cortex, the inferior parietal lobe, and the anterior superior temporal sulcus were shown to underpin this motivation. In contrast, some people cooperate with other individuals primarily because this behavior will maximize their returns. This more calculative drive is underpinned by the dorsolateral prefrontal cortex, the temporo-parietal junction near the posterior superior temporal sulcus, and the precuneus.

Extraversion

Extraversion is positively related to the volume of the medial orbitofrontal cortex, critical to the processing of reward information. This finding aligns to the proposition that extraversion partly emanates from an amplified sensitivity to rewards (DeYoung, Hirsh, Shane, Papademetris, Rajeevan, & Gray, 2010)

Testosterone

Testosterone has been shown to inhibit circuits that include the orbitofrontal cortex. Conceivably, when these circuits are inhibited, individuals often disregard the emotional consequences of their behavior. They will sometimes enact their entrenched responses. They may, for example, behave aggressively even when this behavior will elicit many negative emotions later. This mechanism could partly explain the relationship between testosterone and aggression (Mehta & Beer, 2010).

Variations across the orbitofrontal cortex

Sometimes researchers differentiate the medial and lateral regions of the orbitofrontal cortex. Indeed, the medial regions tend to be associated with more rewarding expectations, and the ventral regions tend to be associated with more punishing expectations.

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