16 Mayıs 2012 Çarşamba

Notes from AN INTEGRATIVE THEORY OF PREFRONTAL CORTEX FUNCTION



My notes from: AN INTEGRATIVE THEORY OF PREFRONTAL CORTEX FUNCTION (41 pages)

Earl K. Miller, Center for Learning and Memory, RIKEN-MIT Neuroscience Research Center and Department of Brain and Cognitive Sciences, MIT.
Jonathan D. Cohen, Center for the Study of Brain, Mind, and Behavior and Department of Psychology, Princeton

Abstract ... cognitive control stems from the active maintenance of patterns of activity in the prefrontal cortex that represent goals and the means to achieve them. They provide bias signals to other brain structures whose net effect is to guide the flow of activity along neural pathways that establish the proper mappings between inputs, internal states, and outputs needed to perform a given task.

INTRODUCTION ...
Simple behaviors can rely on relatively straightforward interactions between the brain’s input and output systems. Animals with fewer than a hundred thousand neurons (in the human brain there are 100 billion or more neurons) can approach food and avoid predators. For animals with larger brains, behavior is more flexible. ...
...
To deal with this multitude of possibilities and to curtail confusion, we have evolved mechanisms that coordinate lower-level sensory and motor processes along a common theme, an internal goal.
...
It is well positioned to coordinate a wide range of neural processes: The PFC is a collection of interconnected neocortical areas that sends and receives projections from virtually all cortical sensory systems, motor systems, and many subcortical structures. ...

The Role of the PFC in Top-Down Control of Behavior
The PFC is not critical for performing simple, automatic behaviors, such as our tendency to automatically orient to an unexpected sound or movement. These behaviors can be innate or they can develop gradually with experience as learning mechanisms potentiate existing pathways or form new ones. These “hardwired” pathways are advantageous because they allow highly familiar behaviors to be executed quickly and automatically (i.e. without demanding attention). However,these behaviors are inflexible, stereotyped reactions elicited by just the right stimulus. They do not generalize well to novel situations, and they take extensive time and experience to develop. These sorts of automatic behaviors can be thought of as relying primarily on “bottom-up” processing; that is, they are determined largely by the nature of the sensory stimuli and well-established neural pathways that connect these with corresponding responses.

By contrast, thePFC is important when “top-down” processing is needed; that is, when behavior must be guided by internal states or intentions. The PFC is critical in situations when the mappings between sensory inputs, thoughts, and actions either are weakly established relative to other existing ones or are rapidly changing. This is when we need to use the “rules of the game,” internal representations of goals and the means to achieve them. Several investigators have argued that this is a cardinal function of the PFC...
...
This illustrates one of the most fundamental aspects of cognitive control and goal-directed behavior: the ability to select a weaker, task-relevant response (or source of information) in the face of competition from an otherwise stronger, but task-irrelevant one. Patients with frontal impairment have difficulty with this task... ...
Overview of the Theory
We assume that the PFC serves a specific function in cognitive control: the active maintenance of patterns of activity that represent goals and the means to achieve them. They provide bias signals throughout much of the rest of the brain, affecting not only visual processes but also other sensory modalities, as well as systems responsible for response execution, memory retrieval, emotional evaluation, etc.

The aggregate effect of these bias signals is to guide the flow of neural activity along pathways that establish the proper mappings between inputs, internal states, and outputs needed to perform a given task. This is especially important whenever stimuli are ambiguous (i.e. they activate more than one input representation), or when multiple responses are possible and the task-appropriate response must compete with stronger alternatives. From this perspective, the constellation of PFC biases...can be viewed as the neural implementation of attentional templates, rules, or goals, depending on the target of their biasing influence.
...
When a behavior meets with success, reinforcement signals augment the corresponding pattern of activity by strengthening connections between the PFC neurons activated by that behavior. This process also strengthens connections between these neurons and those whose activity represents the situation in which the behavior was useful, establishing an association between these circumstances and the PFC pattern that supports the correct behavior.
...
But we believe that this general notion can explain many of the posited functions of the PFC. The biasing influence of PFC feedback signals on sensory systems may mediate its role in directing attention (Stuss &
Benson 1986; Knight 1984, 1997; Banich et al 2000), signals to the motor system may be responsible for response selection and inhibitory control (Fuster 1980, Diamond 1988), and signals to intermediate systems may support short-term (or working) memory (Goldman-Rakic 1987) and guide retrieval from long-term memory (Schachter 1997, Janowsky et al 1989, Gershberg & Shimamura 1995).

Without the PFC, the most frequently used (and thus best established) neural pathways would predominate or, where these don’t exist, behavior would be haphazard. Such impulsive, inappropriate, or disorganized behavior is a hallmark of PFC dysfunction in humans (e.g. Bianchi 1922, Duncan 1986, Luria 1969, Lhermitte 1983, Shallice & Burgess 1996, Stuss & Benson 1986).

Minimal Requirements for a Mechanism of Top-Down Control
There are several critical features of our theory. First, the PFC must provide a source of activity that can exert the required pattern of biasing signals to other structures. We can thus think of PFC function as “active memory in the service of control.” It follows, therefore, that the PFC must maintain its activity robustly against distractions until a goal is achieved, yet also be flexible enough to update its representations when needed. It must also house the appropriate representations, those that can select the neural pathways needed for the task. In sofar as primates are capable of tasks that involve diverse combinations of stimuli, internal states, and responses, representations in the PFC must have access to and be able to influence a similarly wide range of information in other brain regions. That is, PFC representations must have a high capacity for multimodality and integration. Finally, as we can acquire new goals and means, the PFC must also exhibit a high degree of plasticity. Of course, it must be possible to exhibit all these properties without the need to invoke some other mechanism of control to explain them, lest our theory be subject to perennial concerns of a hidden “homunculus.”
...
pattern of deficits following PFC damage as a loss of the ability to acquire and use behavior-guiding rules. ...
...
A GUIDED ACTIVATION THEORY OF PFC FUNCTION ...
A Simple Model of PFC Function ...
Guided Activation as a Mechanism of Cognitive Control
...the role of the PFC is modulatory rather than transmissive. That is, the pathway from input to output does not “run through” the PFC. Instead, the PFC guides activity flow along task-relevant pathways in more posterior and/or subcortical areas.
...
This distinction between modulation vs transmission is consistent with the classic pattern of neuropsychological deficits associated with frontal lobe damage. The components of a complex behavior are usually left intact, but the subject is not able to coordinate them in a task-appropriate way (for example, a patient who, when preparing coffee, first stirred and then added cream). ...

Active Maintenance in the Service of Control...
another critical feature of our theory: the importance of sustained activity as a mechanism of control...
...
executive control involves the active maintenance of a particular type of information: the goals and rules of a task.
...
selective attention and behavioral inhibition are two sides of the same coin: Attention is the effect of biasing competition in favor of task-relevant information, and inhibition is the consequence that this has for the irrelevant information.
...
In particular, we believe that PFC-mediated control is complemented by another form of control dependent on the hippocampal system. The hippocampus is important for binding together information into a memory of a specific episode (Eichenbaum et al 1999, McClelland et al 1995, Squire 1992, Zola-Morgan & Squire 1993). By contrast, we suggest that the PFC, like other neocortical areas, is more important for extracting the regularities across episodes—in the case of the PFC, those corresponding to goals and task rules, rather than episodic memories of actually performing the task. We further posit that the PFC uses “activity-based” control; that is, its ongoing activity specifies the pattern of neural pathways that are currently needed. If PFC activity changes, so does the selected pattern of pathways.

By contrast, the hippocampus may provide a form of “weighted-based” control; it helps consolidate permanent associative links between the pieces of information that define a long-term memory (Cohen & O’Reilly 1996, O’Reilly et al 1999, O’Reilly 2000). To use the railroad metaphor, the hippocampus is responsible for laying down new tracks and the PFC is responsible for flexibly switching between them. As noted below, interactions between the PFC and the hippocampus may provide a basis for understanding prospective forms of control, such as planning.

Updating of PFC Representations
In the real world, cognitive control is highly dynamic...is highly flexible. So long as suitable representations exist within the PFC, activating them can quickly invoke a goal or rule, which can be flexibly switched to others as circumstances demand. That is, it is easier and faster (and perhaps less costly) to switch between existing tracks than it is to lay new ones down.
...
PFC representations are selectively responsive (adaptive) to task-relevant stimuli (Rainer et al 1998b), yet they are resistant to interference (robust) from distractors (Miller et al 1996). Conversely, two hallmarks of damage to the PFC are perseveration (inadequate updating) and increased distractibility (inappropriate updating)
...
Recent modeling work suggests that hierarchical updating and the sequencing of actions may rely on interactions between the PFC and the basal ganglia.

13 Nisan 2012 Cuma

Quotations from "The Prefrontal Cortex: Minireview"

My quotations from: The Prefrontal Cortex: Minireview
Complex Neural Properties for Complex Behavior
Earl K. Miller*
Department of Brain and Cognitive Sciences and The Center for Learning and Memory
Massachusetts Institute of Technology
Cambridge, Massachusetts 02139
...
The PF cortex is a collection of cortical areas in the most anterior portion of the frontal lobes (Figure 1). It has long been associated with high-level, “executive” processes needed for voluntary goal-directed behavior. Its damage in humans does not produce a single, characteristic deficit. Rather, it results in disturbances in a variety of functions, including attention, memory, response selection, planning, and inhibitory control.


Studies of the neural basis of PF function in monkeys focused primarily on active short-term, or working memory. They have revealed that when a delay is imposed between a visual stimulus and a response based on it, many PF neurons show sustained stimulus-related activity (Goldman-Rakic, 1994; Fuster, 1995). Because sensory inputs are often fleeting, this short-term maintenance may be fundamental to many cognitive functions. However, complex behavior requires more than temporary storage. Relevant sensory inputs need to be selected and integrated with other information common to the goal at hand. Also needed are the executive mechanisms that determine, for example, which stimuli are relevant and should be selected. Much less is known about the neural basis of these higher functions.
...
In many views of cognition, the type of executive control thought to be mediated by the PF cortex is synonymous with attentional selection, that is, the ability to voluntarily focus awareness on certain sensory inputs, thoughts, or actions. Selection is necessary because higher-order cognitive functions are severely limited in capacity. Indeed, at a given moment we are aware of only a small fraction of available sensory information.

Given these capacity limitations, the ability to ignore distractions and select behaviorally relevant information is critical. Evidence that the lateral PF cortex is involved in selection processes comes from a number of studies, including a recent study by Rainer et al. (1998b)
...
Earlier target selection in the PF cortex suggests that the visual cortex may “inherit” target information from the PF cortex. That is, that the PF cortex may be a source of the top-down signals that mediate attentional selection in the visual cortex (Desimone and Duncan.)

Complex behavior, however, depends on more than selecting sensory information. To benefit from past experience, we must be able to select (recall) stored knowledge. Watanabe (1996) demonstrated this ability in PF neurons. He found that when monkeys could predict that a specific reward would appear (e.g., raisins, cabbage, etc.), the activity of many PF neurons reflected that expected reward. This ability to prospectively” code predicted events is thus critical for choosing among response alternatives.

Integration
Sensory processing is highly fragmented. Even within a modality, different stimulus attributes may be separately processed. The primate cortical visual system, for example, is thought to analyze the form and color information to identify a stimulus (i.e., what it is) largely from information about stimulus location (i.e., where it is). An area involved in complex behavior, however, needs to have access to diverse information.


Even actions almost invariably require satisfying multiple, diverse constraints. When I search for my coffee cup, for example, I have in mind not only what it looks like but also where it is likely to be. So, somewhere and somehow, diverse information such as what and where needs to come together. Given its role in organizing behavior and its extensive connections, the PF cortex seems a likely region where integration might be evident, particularly when integration is needed for behavior.
...
Other studies have also found that many PF neurons process both object identity and location.
...
Interconnections between different PF regions (Figure 1) could result in a population of PF neurons with multimodal properties.

Associative Learning, Rules, Context, and Cognitive Control
The complexity of primate behavior is partially attributable to the fact that primates can acquire new goals and manners of achieving them. Not surprisingly, the PF cortex is thought to be central to this ability. Its executive role in brain function has been hypothesized to result from the acquisition and representation of “rules” that guide goal-directed behavior.

Rule learning depends on forming arbitrary associations between disparate, but behaviorally related, information. We learn that “red” means “stop,” for example. The PF cortex seems well positioned to play a role in associative learning. It is interconnected with all sensory systems, with the motor system, and with limbic structures involved in long-term memory and affect; it is truly “association cortex.”
...
Thus, PF activity can reflect stimuli, associated actions, and expected consequences.
...
Cohen and colleagues have suggested how the ability of PF neurons to reflect conjunctions of behaviorally related information might result in mechanisms for executive control. They posit that control emanates from representation of context (Cohen and Servan-Schreiber, 1992). Context is the constellation of task-relevant information needed to guide a given behavior.
...
Top-down signals from the PF cortex are excitatory and represent the to-be-attended item. These signals increase activity of neurons that process the relevant information and, by virtue of the mutual inhibition, suppress activity of neurons processing irrelevant information. Context representations may act in a similar fashion. However, rather than convey only visual attributes, context representations are thought to be multimodal and include information about stimuli, actions, etc., that have become associated through experience. Thus, they can bias motor as well as sensory processing and also allow appropriate actions to be selected and executed.

Conclusions
Recent studies have shown that, consistent with their putative role in the highest level of cognitive function, PF neurons have complex response properties that are highly dependent on, and shaped by, task demands. They selectively process and integrate information needed for a common behavioral goal. Thus, its extensive connections and the ability of its neurons to be modified by experience may allow the PF cortex to play a role in knitting together behaviorally relevant associations, a process needed for acquisition of “rules” that guide goal-directed behavior. This may result in a representation of context, a template of a previously successful configuration of sensory and response-related information that biases processing in other brain regions in favor of task-relevant information. Such complicated and malleable activity would be expected for a region so closely linked with the complexity and flexibility that are the hallmarks of primate behavior.

10 Mart 2012 Cumartesi

Emotion Regulation

My quotations from:
Emotion Regulation: Conceptual Foundations Date: 3/22/06
James J. Gross (Stanford Uni), Ross A Thompson (Uni of California, Davis)
...
Emotion Regulation and Related Processes
...
emotion regulation refers to the heterogeneous set of processes by which emotions are themselves regulated. Emotion regulatory processes may be automatic or controlled, conscious or unconscious, and may have their effects at one or more points in the emotion generative process

Because emotions are multi-componential processes that unfold over time, emotion regulation involves changes in “emotion dynamics” (Thompson, 1990), or the latency, rise time, magnitude, duration, and offset of responses in behavioral, experiential, or physiological domains. Emotion regulation may dampen, intensify, or simply maintain emotion, depending on an individual’s goals. Emotion regulation also may change the degree to which emotion response components cohere as the emotion unfolds, such as when large changes in emotion experience and physiological responding occur in the absence of facial behavior.

One as-yet unresolved issue is whether emotion regulation refers to intrinsic processes (Fred regulates his own emotions: emotion regulation in self), to extrinsic processes (Sally regulates Bob’s emotions: emotion regulation in other), or to both. In general, researchers in the adult literature typically focus on intrinsic processes (Gross, 1998). By contrast, researchers in the developmental literature focus more on extrinsic processes.


Core Features of Emotion Regulation
...
Emotion Regulation and Related Constructs
..., we see emotion regulation as subordinate to the broader construct of affect regulation. Under this broad heading fall all manner of efforts to influence our valenced responses (Westen, 1994).
... affect regulation includes (among other things) four overlapping constructs: (a) coping, (b) emotion regulation, (c) mood regulation, and (d) psychological defenses.
...
Emotion Regulation Strategies
...These five points represent five families of emotion regulation processes: situation selection, situation modification, attentional deployment, cognitive change, and response modulation (Gross, 1998b).

...the first four emotion regulation families may be considered antecedent-focused, in that they occur before appraisals give rise to full-blown emotional response tendencies, and may be contrasted with response-focused emotion regulation, which occurs after the responses are generated (Gross & Munoz, 1995).

Situation Selection
The most forward-looking approach to emotion regulation is situation selection. This type of emotion regulation involves taking actions that make it more (or less) likely that one will end up in a situation one expects will give rise to desirable (or undesirable) emotions.

Another barrier to effective situation selection is appropriately weighing short-term benefits of emotion regulation versus longer-term costs. For example, a shy person may feel much better in the short term if she avoids social situations. However, this short-term relief may
come at the cost of longer term social isolation.

Situation Modification
Potentially emotion-eliciting situations – such as the approach of the terrifying barber in
the example above – do not inevitably lead to emotional responses. After all, one can always ask to wait until a less frightening barber is free. Such efforts to directly modify the situation so as to alter its emotional impact constitute a potent form of emotion regulation.

Attentional Deployment
Situation selection and situation modification help shape the individual’s situation. However, it also is possible to regulate emotions without actually changing the environment. Situations have many aspects, and attentional deployment refers to how individuals direct their attention within a given situation in order to influence their emotions.

Distraction focuses attention on different aspects of the situation, or moves attention away from the situation altogether, such as when an infant shifts its gaze from the emotioneliciting
stimulus to decrease stimulation (Rothbart, this volume; Stifter & Moyer, 1991).

Distraction also may involve changing internal focus, such as when individuals invoke thoughts or memories that are inconsistent with the undesirable emotional state (Watts, this volume), or when an actor calls to mind an emotional incident in order to portray that emotion convincingly.

Concentration draws attention to emotional features of a situation. Wegner and Bargh (1997) have termed this “controlled starting” of emotion. When attention is repetitively directed to one’s feelings and their consequences, this is referred to as rumination. Ruminating on sad
events leads to longer and more severe depressive symptoms (Just & Alloy, 1997; Nolen-
Hoeksema, 1993).

Cognitive Change
Even after a situation has been selected, modified, and attended to, an emotional response
is by no means a foregone conclusion. Emotion requires that percepts be imbued with meaning and that individuals evaluate their capacity to manage the situation. As described above, appraisal theorists have described the cognitive steps needed to transform a percept into something that elicits emotion. Cognitive change refers to changing how one appraises the situation one is in so as to alter its emotional significance, either by changing how one thinks about the situation or about one’s capacity to manage the demands it poses.

One form of cognitive change that has received particular attention is reappraisal (Gross,
2002; John & Gross, this volume; Ochsner & Gross, this volume). This type of cognitive change involves changing a situation’s meaning in a way that alters its emotional impact. Leading subjects to reappraise negatively valenced films has been shown to result in decreased negative emotion experience.

Response Modulation
In contrast with other emotion regulatory processes, response modulation occurs late in
the emotion generative process, after response tendencies have been initiated. Response
modulation refers to influencing physiological, experiential, or behavioral responding as directly as possible. Attempts at regulating the physiological and experiential aspects of emotion are common.

Another common form of response modulation involves regulating emotion-expressive
behavior (Gross, John, & Richards, in press). A person may wish to regulate expressive behavior for many reasons, ranging from an assessment that it would be best to hide one’s true feelings from another person (e.g., hiding one’s fear when standing up to a bully) to direct prompts from a parent (e.g., in the barbershop example).

23 Şubat 2012 Perşembe

Ego-control and ego-resiliency

My quotations from:

Ego-control and ego-resiliency: Generalization
of self-report scales based on personality
descriptions from acquaintances,
clinicians, and the self
2004

Tera D. Letzringa,*, Jack Blockb, David C. Fundera
a Department of Psychology, University of California, Riverside, CA 92521, USA
b University of California, Berkeley, USA

Abstract
Ego-control refers to the inhibition/expression of impulse and ego-resiliency (ER) to the dynamic capacity to contextually modify one_s level of ego-control in response to situational affordances (Block, J., 1950, 2002; Block, J.H., 1951; Block & Block, 1980).
...

1. Introduction
Ego-control (EC) and ego-resiliency (ER) are conceptualized as central personality constructs for understanding motivation, emotion, and behavior (Block, J., 1950, 2002; Block, J.H., 1951; Block & Block, 1980). Broadly conceived and summarily characterized, EC refers to a meta-dimension of impulse inhibition/expression and ER refers to a meta-dimension of the dynamic capacity to contextually modify one’s level of control in response to situational demands and affordances.

Overcontrolled individuals characteristically contain impulse and affect across situations, even when
doing so may not be necessary. On the other hand, undercontrolled individuals characteristically express impulse and affect across situations, even when doing so may be inappropriate. Highly ego-resilient individuals are characteristically able to modify their level of control, either up or down, as may be appropriate or necessary according to the situational context. Individuals with a low level of ego-resiliency are more restricted to the same level of impulse containment or expression regardless of situational demands.

1.1. Ego-control
According to theory developed by Jack and Jeanne Block over several decades, the individual difference dimension of ego-control varies from overcontrol to undercontrol (Block, 2002; Block & Block, 1980). Overcontrolled individuals are conceptualized as relatively inhibited in action and affect-expressiveness to the point of at times being excessively constrained. They have difficulty making decisions, may unnecessarily delay gratification or deny themselves pleasure, are tightly organized,
are insulated from environmental distractions, and are able to continue even repetitive tasks for long periods of time.

At the other extreme, undercontrolled individuals characteristically express affect and impulses relatively immediately and directly even when doing so may be socially or personally inappropriate. They are relatively unable to delay gratification, have fluctuating emotions, and are spontaneous, easily distracted, and relatively unbound by social customs (Block, 2002; Funder & Block, 1989).

The consequences of characteristic overcontrol or undercontrol may be adaptive or maladaptive depending on circumstances. Overcontrol may facilitate disciplined and directed behavior, which can be advantageous in some situations. In other contexts, where delaying gratification and pleasure is unwarranted or psychologically undesirable, overcontrol is likely to be detrimental to personal and often societal fruition.

In parallel, undercontrol can facilitate the expression of warmth, friendliness, and spontaneity, which are likely to be advantageous in promoting intimacy and the enjoyment of life. However, undercontrol can be maladaptive when it leads to erratic, unorganized, or dangerous behavior.

1.2. Ego-resiliency
According to the Blocks_ theorizing, ego-resiliency is the ability to adapt one_s level of control temporarily up or down as circumstances dictate (Block, 2002; Block & Block, 1980). As a result of this adaptive flexibility, individuals with a high level of resiliency are more likely to experience positive affect, and have higher levels of selfconfidence and better psychological adjustment than individuals with a low level of resiliency (Block & Kremen, 1996; Klohnen, 1996). When confronted by stressful circumstances, individuals with a low level of resiliency may act in a stiff and perseverative manner or chaotically and diffusely, and in either case, the resulting behavior is likely to be maladaptive (Block & Kremen, 1996).

3 Şubat 2012 Cuma

How to Regulate Emotions

The crux of this article from the point of daily application is:

“ Emotional processing can also be modulated through deliberate and conscious application of top-down executive control over processing of an emotional stimulus”.

You can find how this is done:
1- One page outline made of quotations from the original article
2- 4 Pages of my notes made of quotations from the original article
3- Original article itself.

1111111111111111111111111111111111111111111111111111111111111111111111
Emotional processing in anterior cingulate and medial prefrontal cortex
Amit Etkin1,2, Tobias Egner3 and Raffael Kalisch4

In summary, the pattern of anatomical connectivity supports an important role for the sgACC, pgACC and adACC in interacting with the limbic system, including its effector regions, and for the adACC and pdACC in communicating with other dorsal and lateral frontal areas that are important for top-down forms of regulation [72].

Fear conditioning and extinction in humans

Thus, the dorsal ACC and mPFC seem to function generally in the appraisal and expression of fear or anxiety.

To elucidate how fear is regulated, we next discuss activations associated with extinction of learned fear
. In extinction, the CS is repeatedly presented in the absence of reinforcement, leading to the formation of a CS(conditioned stimulus)–no US(unconditioned stimulus) association (or extinction memory) that competes with the original fear memory for control over behavior [28–30]. Hence, extinction induces conflicting appraisals of, and response tendencies to, the CS because it now signals both threat and safety, a situation that requires regulation, as outlined above.

Emotional conflict regulation
The circuitry we find to be specific for regulation of emotional conflict (ventral ACC and mPFC and amygdala) is very similar to that involved in extinction.

Top-down control of emotion
During emotional conflict regulation, emotional processing is spontaneously modulated in the absence of an explicit instruction to regulate emotion. Emotional processing can also be modulated through deliberate and conscious application of top-down executive control over processing of an
emotional stimulus.
The best-studied strategy for the latter type of regulation is reappraisal, a cognitive technique whereby appraisal of a stimulus is modified to change its ability to elicit an emotional reaction [42]. Reappraisal involves both the initial emotional appraisal process and
the reappraisal process proper, whereby an additional positive appraisal is created that competes with the initial negative emotional appraisal.

These data suggest that controlled top-down regulation, like emotional conflict regulation, uses ventral ACC and mPFC areas to inhibit negative emotional processing in the amygdala, thus dampening task interference.

The ventral ACC and mPFC might thus perform a generic negative emotion inhibitory function that can be recruited by other regions (e.g. dorsal ACC and mPFC and lateral PFC) when there is a need to suppress limbic reactivity [10].

22222222222222222222222222222222222222222222222222222222222222222222
Emotional processing in anterior cingulate and medial prefrontal cortex
Amit Etkin1,2, Tobias Egner3 and Raffael Kalisch4
1 Department of Psychiatry and Behavioral Sciences, Stanford University, Stanford, CA, USA
2 Sierra Pacific Mental Illness, Research, Education, and Clinical Center (MIRECC) at the Veterans Affairs Palo Alto Health Care System, Palo Alto, CA, USA
3 Department of Psychology & Neuroscience and Center for Cognitive Neuroscience, Duke University, Durham, NC, USA
4 Institute for Systems Neuroscience and NeuroImage Nord, University Medical Center Hamburg-Eppendorf (UKE), Hamburg, Germany

Negative emotional stimuli activate a broad network of brain regions, including the medial prefrontal (mPFC) and anterior cingulate (ACC) cortices. An early influential view dichotomized these regions into dorsal–caudal cognitive and ventral–rostral affective subdivisions. In this review, we examine a wealth of recent research on negative emotions in animals and humans, using the example of fear or anxiety, and conclude that, contrary to the traditional dichotomy, both subdivisions make key
contributions to emotional processing. Specifically, dorsal– caudal regions of the ACC and mPFC are involved in appraisal and expression of negative emotion, whereas ventral–rostral portions of the ACC and mPFC have a regulatory role with respect to limbic regions involved in generating emotional responses. Moreover, this new framework is broadly consistent with emerging data on other negative and positive emotions.
...
Glossary
Appraisal: evaluation of the meaning of an internal or external stimulus to the organism. Only stimuli that are appraised as motivationally significant will induce an emotional reaction, and the magnitude, duration and quality of the emotional reaction are a direct result of the appraisal process. Moreover,
appraisal can be automatic and focus on basic affective stimulus dimensions such as novelty, valence or value, or expectation discrepancy, or may be slower and sometimes even require controlled conscious processing, which permits a more sophisticated context-dependent analysis.

Fear conditioning: learning paradigm in which a previously neutral stimulus, termed the conditioned stimulus (CS), is temporally paired with a non-learned aversive stimulus, termed the unconditioned stimulus (US). After pairing, the CS predicts the US and hence elicits a conditioned response (CR). For example, pairing of a tone with a foot shock results in elicitation of fear behavior during
subsequent responses to a non-paired tone.

Extinction: learning process created by repeatedly presenting a CS without pairing with an US (i.e. teaching the animal that the CS no longer predicts the US) after fear conditioning has been established. This results in formation of an extinction memory, which inhibits expression of, but does not erase, the original fear memory.

Reappraisal: specific method for explicit emotion regulation whereby a conscious deliberate effort is engaged to alter the meaning (appraisal) of an emotional stimulus. For example, a picture of a woman crying can be reappraised from a negative meaning to a positive one by favoring an
interpretation that she is crying tears of joy.

Regulation: general process by which conflicting appraisals and response tendencies are arbitrated between to allow selection of a course of action. Typically, regulation is thought to have an element of inhibition and/or enhancement for managing competing appraisals and response tendencies.
...
Box 1. Anatomy of the ACC and mPFC
...
In summary, the pattern of anatomical connectivity supports an important role for the sgACC, pgACC and adACC in interacting with the limbic system, including its effector regions, and for the adACC and pdACC in communicating with other dorsal and lateral frontal areas that are important for top-down forms of regulation [72].
...
Fear conditioning and extinction in humans
The paradigms used in the acquisition and extinction of learned fear are particularly valuable for isolating the neural substrates of fear processing because the anticipatory fear or anxiety triggered by the previously neutral conditioned stimulus (CS) can be dissociated from the reaction to the aversive unconditioned stimulus (US) per se. This is not possible in studies that, for example, use
aversive images to evoke emotional responses. Furthermore, comparison between fear conditioning and fear extinction facilitates an initial coarse distinction between regions associated with either the appraisal of fear-relevant stimuli and generation of fear responses (fear conditioning), for the inhibitory regulation of these processes (extinction).
...
sympathetic nervous system activity correlates positively primarily with dorsal ACC and mPFC regions and negatively primarily with ventral ACC and mPFC regions, which supports a role for the dorsal ACC and mPFC in fear expression (c).
...
These processes are intermixed with, and supported by, learning processes, namely, acquisition, consolidation and storage of a fear memory (CS–US association), and retrieval of the fear memory on subsequent CS presentations.
...
Thus, the dorsal ACC and mPFC seem to function generally in the appraisal and expression of fear or anxiety.
...
To elucidate how fear is regulated, we next discuss activations associated with extinction of learned fear. In extinction, the CS is repeatedly presented in the absence of reinforcement, leading to the formation of a CS–no US association (or extinction memory) that competes with the original fear memory for control over behavior [28–30]. Hence, extinction induces conflicting appraisals of, and response tendencies to, the CS because it now signals both threat and safety, a situation that requires regulation, as outlined above.
...
Emotional conflict regulation
...
The circuitry we find to be specific for regulation of emotional conflict (ventral ACC and mPFC and amygdala) is very similar to that involved in extinction. ... Much like the relationship between improved emotional conflict regulation and decreased conflict evaluation-related activation in the dorsal ACC and mPFC, more successful extinction is associated with decreased CS-driven activation in the dorsal ACC and mPFC of humans and rodents [40,41]. Thus, the most parsimonious explanation for these data is that emotional conflict evaluation-related functions involve overlapping neural mechanisms with appraisal and expression of fear, and that regulation of emotional conflict also involves circuitry that overlaps with fear extinction.
...
Top-down control of emotion
During emotional conflict regulation, emotional processing is spontaneously modulated in the absence of an explicit instruction to regulate emotion. Emotional processing can also be modulated through deliberate and conscious application of top-down executive control over processing of an
emotional stimulus. The best-studied strategy for the latter type of regulation is reappraisal, a cognitive technique whereby appraisal of a stimulus is modified to change its ability to elicit an emotional reaction [42]. Reappraisal involves both the initial emotional appraisal process and
the reappraisal process proper, whereby an additional positive appraisal is created that competes with the initial negative emotional appraisal. Thus, we would expect reappraisal to involve the dorsal ACC and mPFC regions that we observed to be important for emotional conflict detection (Figure 2a). Consistent with this prediction, a meta-analysis found that reappraisal was reliably associated with activation in the dorsal ACC and mPFC (Figure 2b) [43].

This reappraisal meta-analysis, interestingly, did not implicate a consistent role for the ventral ACC and mPFC [43], which suggests that reappraisal does not primarily work by suppressing the processing of an undesired emotional stimulus. Nevertheless, activity in the ventral ACC and mPFC in some instances is negatively correlated with activity in the amygdala in paradigms in which reappraisal resulted in downregulation of amygdalar activity in response to negative pictures [44,45]. ...
These data suggest that controlled top-down regulation, like emotional conflict regulation, uses ventral ACC and mPFC areas to inhibit negative emotional processing in the amygdala, thus dampening task interference. The ventral ACC and mPFC might thus perform a generic negative emotion inhibitory function that can be recruited by other regions (e.g. dorsal ACC and mPFC and lateral PFC) when there is a need to suppress limbic reactivity [10]. This would be a prime example of parsimonious use of a basic emotional circuitry, conserved between rodents and humans (Box 2), for the purpose of higher-level cognitive functions possible only in humans.
...
Amygdala–ACC and –mPFC functional connectivity
...
In addition, during regulation tasks, connectivity was restricted to the ventral ACC and mPFC and was primarily negative (Figure 2d). These data thus lend further support to our proposal of a dorso–ventral separation in terms of negative emotion generation (appraisal and expression) and inhibition (regulation).
...
Integration with other perspectives on ACC and mPFC function and other emotions
Although less developed than the literature on fear and anxiety, studies on other emotions are broadly consistent with our formulation of ACC and mPFC function. On the negative emotion appraisal and expression side, direct experience of pain, or empathy for others experiencing
pain, activates the dorsal ACC and mPFC [49], and lesions of the dACC also serve as treatment for chronic pain [50]. Similarly, increased sensitivity to a range of negative emotions is associated with greater engagement of the dorsal ACC and mPFC, including disgust [51] and rejection [52], and transcranial-magnetic-stimulation-induced disruption of the dmPFC interferes with anger processing [53]. Uncertainty or ambiguity, which can induce anxiety and relates to emotional conflict, leads to activation in the dACC and dmPFC [54].
...
Positive emotion, which can serve to regulate and diminish negative emotion, has been associated in a metaanalysis with activation in the sgACC, vmPFC and pgACC [58]. Extinction of appetitive learning activates the vmPFC [59], much as extinction of learned fear does. The evaluation of positive stimuli and reward is more complicated. For instance, Rushworth and co-workers proposed that the processes carried out by the adACC are mirrored by similar contributions to reinforcement-guided decision-making from the orbitofrontal cortex, with the distinction that the adACC is concerned with computing reinforcement value of actions whereas the orbitofrontal cortex is concerned with gauging the reinforcement values of stimuli [60].

Taken together, these data broadly support our dorsal–ventral distinction along appraisal–expression versus regulation lines, with respect specifically to negative emotion.
...

21 Ocak 2012 Cumartesi

understanding the interaction between cognition and emotion

Short Notes from:
Cognitive and emotional influences in anterior cingulate cortex
George Bush, Phan Luu and Michael I. Posner

Anterior cingulate cortex (ACC) is a part of the brain’s limbic system. Classically, this region has been related to affect, on the basis of lesion studies in humans and in animals.
...
ACC might be the brain’s error detection and correction device.
...
ACC is a part of a circuit involved in a form of attention that serves to regulate both cognitive and emotional processing. Neuroimaging studies showing that separate areas of ACC are involved in cognition and emotion.
...
(1) Cingulate cortex includes specific processing modules for sensory, motor, cognitive and emotional information.
(2) As a whole, cingulate cortex integrates input from various sources (including motivation, evaluation of error, and representations from cognitive and emotional networks).
(3) Cingulate cortex acts by influencing activity in other brain regions and modulating cognitive, motor, endocrine and visceral responses.
...

cingulate cortex encompasses numerous specialized subdivisions that subserve a vast array of cognitive, emotional, motor, nociceptive and visuospatial functions.
...
Its two major subdivisions subserve distinct functions. These include a dorsal cognitive division (ACcd; areas 24b9-c9 and 329) and a rostral–ventral affective division (ACad; rostral areas 24a–c and 32, and ventral areas 25 and 33).
...
The cognitive subdivision is part of a distributed attentional network. It maintains strong reciprocal interconnections with lateral prefrontal cortex (BA 46/9), parietal cortex (BA 7), and premotor and supplementary motor areas6. Various functions have been ascribed to the ACcd, including modulation of attention or executive functions by influencing sensory or response selection (or both); monitoring competition, complex motor control, motivation, novelty, error detection and working memory; and anticipation of cognitively demanding tasks (see Refs 1,3,5,6,14–17 for reviews).


The affective subdivision, by contrast, is connected to the amygdala, periaqueductal gray, nucleus accumbens, hypothalamus, anterior insula, hippocampus and orbitofrontal cortex6, and has outflow to autonomic, visceromotor and endocrine systems. The ACad is primarily involved in assessing the salience of emotional and motivational information and the regulation of emotional responses5,6,15,18.
...
The cognitive division (ACcd) has been activated (Fig. 2a) by cognitively demanding tasks that involve stimulus– response selection in the face of competing streams of information, including Color Stroop and Stroop-like tasks, divided- attention tasks, verbal- and motor-response selection tasks and many working-memory tasks. The affective division (ACad) has been activated (Fig. 2a) by affect-related tasks, including studies of emotional processing in normal healthy volunteers and symptom provocation studies in a number of psychiatric disorders (anxiety, simple phobia and obsessive–compulsive disorder). It has also been activated repeatedly by induced sadness in normal subjects
and in individuals with major depression.
...
Combining functional imaging, anatomical and behavioral methods will be important for understanding the interaction between cognition and emotion.
...
These studies suggest that both dorsal ACC and areas of the lateral prefrontal cortex operate together during tasks that involve high levels of mental effort.
...
According to this hypothesis, the cognitive division of the ACC serves to monitor crosstalk or conflict
between brain areas, and this computation signals the need for control processes. Lateral areas of the cortex are then activated to provide control operations, which might include increasing or inhibiting neural activity within distinct brain areas so as to eliminate the confusion between modules.

2 Ocak 2012 Pazartesi

Short Notes from Conflict and Cogntive Control in the Brain

Short Notes for Interested People from

Conflict and Cogntive Control in the brain
Vincent van Veen1 and Cameron S. Carter2
1Department of Psychology, University of Pittsburgh, and 2Departments of Psychology and Psychiatry, University of California, Davis

ABSTRACT—Recent research from cognitive psychology and cognitive neuroscience has suggested that the control mechanisms by which people are able to regulate task performance can be dissociated into evaluative and executive components.
One process, implemented in the anterior cingulate cortex of the brain, monitors the amount of conflict that occurs during information processing; another process, implemented in the dorsolateral prefrontal cortex, is involved with maintaining the requirements of the task at hand and with biasing information processing in favor of appropriate responses.
...
Whenever one performs a task, one has to make sure that one selects the relevant information (stimuli, actions) and not get distracted by stimuli or thoughts that are irrelevant to the task. Such distraction might lead to inappropriate actions, such as errors. How the brain manages to do this is the central question in this paper: specifically, how people manage to pay more attention after they have either made an error or almost made an error.

One of the key aspects of cognitive control is how flexible it is.
The issue of how people monitor and correct for errors has become a popular topic of inquiry in cognitive research;
...
ATTENTIONAL CONTROL AND THE PREFRONTAL CORTEX
Many phenomena from cognitive and social psychology are thought to depend on automatic processes. Processes can be automatic when they are innate or highly practiced, because they are part of a strongly activated schema, or because they are imposed by a powerful social context or a strong motivational or emotional state, among other reasons.

Of all the stimuli that one encounters, only a limited set is relevant to what one is trying to accomplish; nevertheless, both relevant and irrelevant stimuli compete for access to the response system. Such competition can be difficult to overcome when the processing of irrelevant information is relatively automatic. When the irrelevant information is associated with a response, reliance on automatic processing might facilitate the task if this response is desirable. However, when the irrelevant, automatically processed information is associated with an inappropriate response, the resulting conflict between appropriate and inappropriate responses might be difficult to overcome.
...
This is thought to occur because word reading is relatively more automatic than color naming; therefore the word is hard to ignore, and it activates the associated response. During incongruent trials, the responses associated with the color compete with those associated with the word. When conflicts occur, attentional control is needed to overcome the conflict by selecting the relevant information and suppressing the processing of irrelevant information. Control is often conceptualized as the ability to represent and maintain the task requirements, to support the processing of information relevant to the goals of the current task, and to suppress irrelevant information. Many neuropsychological and neuroimaging studies have shown that an important brain structure that supports this ability is the prefrontal cortex (PFC ).
...
The PFC is also involved in the control of other forms of higher-order cognition. For instance, many neurophysiological and neuroimaging studies have shown that
this region is able to maintain task-relevant information for brief periods of time, which has supported the notion that this region constitutes a neural basis for working memory. Also, the PFC is involved in task preparation and switching between different tasks. Therefore, it is thought to be involved in many aspects of the executive control over our thoughts and actions.
...
PERFORMANCE MONITORING
The notion that control regulates other types of information processing begs the question, how is control itself regulated? Without a clear answer to this question, the notion of attentional control becomes homuncular, posing the same questions as it is supposed to answer. Therefore, an answer to this question is fundamental to understanding the flexible nature of attentional control.

We and other researchers have argued that
the amount of conflict occurring in the information-processing system plays a central role in regulating how much control is exerted and when control is withdrawn. The anterior cingulate cortex (ACC) of the brain, located on the medial surface of the frontal lobe, has (possibly among other functions) the role of detecting situations in which there are conflicts within the information-processing stream and which therefore require attention to be resolved
...
This activation of the incorrect response is followed by activation of the correct response. If the initial incorrect activation does not manage to reach response threshold and the correct response manages to override the incorrect response, the response to the trial ends up being correct. Errors in speeded-response tasks are typically fast, impulsive responses based on incomplete stimulus evaluation. The timing of response activation during error trials does not differ much from that during correct trials, the main exception being that during error trials the initial activation of the incorrect response does reach response threshold. The subsequent activation of the correct response is manifested as the tendency to ‘‘correct’’ the error
...
FUTURE DIRECTIONS
Many studies, including subliminal-priming studies, have suggested that conflict can occur in the absence of awareness; however, studies have not been conclusive as to whether this conflict also engages the ACC or whether such conflict is associated with a subsequent increase in attention.
...
Similarly, these areas have also been suggested to play a role in cognitive dissonance; Harmon-Jones (2004) has hypothesized that the ACC detects conflicts (dissonance) between actions and attitudes and alerts the PFC to reduce this dissonance.