Inhibition

AVISHAI HENIK
Ben Gurion University of the Negev, Israel

THOMAS H. CARR
Michigan State University

I. The Role of Inhibition in Cognition and Behavior II. Attending to a Spatial Location III. Selection for Action and Conict Resolution IV. Priming and Retrieval from Memory V. Executive Control of Task Performance: Stopping an Ongoing Thought or Action VI. Developmental Aspects VII. Conclusions

GLOSSARY
anterior cingulate cortex A region of the medial frontal lobe involved in a variety of cognitive, motor, and emotionalmotivational functions, including Brodmanns area 24 and perhaps the closely associated area 25. attention A class of cognitive processes involving prioritization or selection among stimuli to be processed, tasks to be performed, or responses to produce. frontal eye eld A region centered at the intersection of the precentral sulcus with the superior frontal sulcus, that is, Brodmanns area 6. This region is involved in generation and control of eye movements. reaction time The time elapsed from onset of the imperative stimulus to initiation of the subjects response. saccade Rapid eye movement that changes the point of xation from one location to another. stimulus onset asynchrony The time elapsed from the onset of the rst stimulus (or the cue) to the onset of the imperative stimulus.

restrains its transmission from one place in the brain to another. This article addresses hypotheses about the role of inhibition at the systems level, considering the possible functional consequences of inhibitory processes for cognition and behavior. Ultimately, one might guess that at the neuronal level, inhibitory processes would be implemented via neuron-to-neuron communication in which release of inhibitory neurotransmitters reduces the probability of action potentials in postsynaptic target neurons of brain structures whose activity needs to be curtailed. As will be shown, however, reduction or suppression of activation is not the only way in which a systems-level inhibitory outcome can be achieved in cognition or behavior, although it is the most commonly proposed mechanism.

I. THE ROLE OF INHIBITION IN COGNITION AND BEHAVIOR

Cognition and the control of overt behavior rely on real-time orchestration of component cognitive processes or mental operations. Each operation achieves a step in the sequence of steps leading from stimulus to response, intention to action, or thought to thought. A major distinction is made between reexive or automatic operations and voluntary or controlled operations. The more automatic an operation is, the more able it is to occur without intention, needing only the appropriate stimulus conditions or information inputs to trigger it; to occur outside of

Inhibition refers to mechanisms by which the nervous system suppresses information, restricts its use, or

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Copyright 2002, Elsevier Science (USA). All rights reserved.

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conscious awareness, without being noticed phenomenologically; and to run in parallel with other mental operations. Automatic operations gain these properties either from genetic hardwiring or, more frequently, from repetition under relatively unchanging conditions. Hence, they sometimes occur as reexes, and they become quite prominent in familiar situations and well-practiced tasks. Controlled operations are the opposite. The more voluntary or controlled an operation is, the more its execution is intentional, conscious, and demanding of serial attention. Controlled processes become prominent when dealing with novel situations to which reexes and habits are poorly adapted or when pursuing particular goals in situations that are likely to trigger reexes and habits that would produce incorrect or inappropriate behavior if unrestrained. To achieve exibility in dealing with both the familiar, unchanging aspects of the world and with novel events, it is important to make automatic and controlled processes work in concert. Inhibition provides a tool for curbing or regulating automated responses in the service of controlled assessment and reaction. Just how fundamental a role is played by inhibition of automated responses can be appreciated by thinking about the developmental trajectory of reexes across the life span. One of the central principles of neurology is that disease processes affecting higher brain centers, especially the cerebral cortex, are revealed by reappearance of primitive reexes. The knee jerk of a normal person is tonically inhibited, but it can become hyperactive after damage to the spinal cord that interrupts descending inhibitory pathways from the motor cortex. The sucking reex of infants disappears after the nursing years but can reappear in a patient with Alzheimers disease. Presumably, with the development of the nervous system, these primitive reexes are inhibited throughout adulthood but may be disinhibited and reappear due to nervous system insult. Other reexes remain active throughout the life span. Among the most common are reexive orienting reactions that involve the automatic deployment of attention to a suddenly appearing visual stimulus or a loud sound. Because these are very common occurrences, such attentional reactions are frequent and do not always occur at convenient times. When controlled deployment of attention is required by task performance, disruptive reexive deployments may need to be inhibited.

Analogous problems can arise in controlling responses that are not reexes but have become automated through practice. Attention decit and hyperactivity disorder is a persistent individual difference characterized by an impulsive inability to resist engaging in prepotent or automated actions triggered by task-irrelevant stimuli in the environment or taskirrelevant thoughts in the mind. Intrusion of an unwanted or inappropriate automatic response occasionally occurs in normal children and adults as well. This can be seen in slips of action in which distraction of attention while carrying out a lowfrequency task can result in inadvertently executing a different task that is higher in frequency given the situation and stimulus environment. Other inhibitory functions are aimed at suppressing unwanted or incorrect perceptions and thoughts rather than controlling attention or inhibiting overt actions. These extraneous mental representations may become activated through relatively automatic processes of generalization because they are similar to correct perceptions and thoughts, or they may become activated because the environmental situation is ambiguous and admits multiple interpretations, only one of which is relevant to the task at hand. It is obvious that the wrong interpretation can be reached in an ambiguous situation, and if so it will need to be replaced. Less obvious, perhaps, is the possibility that all the interpretations of an ambiguous stimulus might be computed relatively automatically at an early stage of processing on every occasion, with one chosen for further processing at a later stage. In either case, contextually inappropriate interpretations will be active at some point in task performance and will need to be put aside or eliminated. This may involve inhibition. In this regard, Sir John Eccles wrote the following in 1977: I always think that inhibition is a sculpturing process. The inhibition, as it were, chisels away at the diffuse and rather amorphous mass of excitatory action and gives a more specic form to the neuronal performance at every stage of synaptic relay. In this article, we present several examples representing the types of inhibitory function we have just described. Some are examples of inhibition regulating the deployment of attention. Others are examples of inhibition enabling disengagement from ongoing or automatically triggered responding so that overt behavior can be under voluntary and strategic control.

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Still others are examples of inhibition tuning and sharpening the representation of the stimulus produced by perception or the interpretation of the situation produced by higher cognitive processes.

II. ATTENDING TO A SPATIAL LOCATION

Orienting of visual attention to a point of interest is commonly accompanied by overt movements of the head, eyes, or body. Attending may originate at will, such as when we decide to look at a particular location where something of interest is expected, or it may originate reexively without intention when something captures our attention, such as when we orient to a ash of light in the dark or to a movement in the periphery of our vision. In everyday life there are constantly competing demands on attention by the outside world as well as from internally generated goals. The need for mechanisms to arbitrate between these competing demands is straightforward: This must be done so that they can be integrated, prioritized, or selected among to provide coherent and adaptive behavior. Michael Posner developed a paradigm widely employed to study visual spatial attention. Figure 1 shows the basic features of this paradigm. In a typical experiment, the subject is rst presented with three boxes on a computer screen. A trial begins with a xation cross at the middle box. After a short interval, one of the boxes may ash briey, and a target (an asterisk) appears in one of the peripheral boxes. The subject is asked to respond as quickly as possible, by pressing a key, to the appearance of the target. Reaction time (RT) is measured (in milliseconds) from target onset until the subjects response. It is possible to study covert attention by asking the participants not to move their eyes and by measuring keypress responses (rather than saccade latencies).

Figure 2 Typical time course for effects of a peripheral nonpredictive luminance cue (box 50% cue) and a central predictive arrow cue. The task is a simple RT key press response to the appearance of the target. Mean detection RTs are presented as a function of cuetarget interval (SOA) for valid and invalid cue conditions.

Figure 1 Basic paradigm for spatial attention, showing an exogenous cue and a valid trial.

The cue may summon attention to the target location, in which case it is a valid cue, or it may summon attention to the wrong location, in which case it is an invalid cue. For volitional goal-directed shifts of attention, often called endogenous shifts of attention, a central arrow serves as a cue and predicts where the target is likely to occur in most trials. That is, in 80% of the trials the target will appear at the valid cue location and in 20% of the trials the target will occur at the invalid cue location. For reexive stimulus-driven shifts of attention, often called exogenous shifts of attention, the peripheral luminance change that serves as a cue has no predictive value with respect to where the target will occur (e.g., in 50% of the trials the target will occur at the valid cue location and in 50% of the trials the target will appear at the invalid cue location). In order to measure the effectiveness of the cue in summoning attention, researchers have manipulated the time interval between cue onset and target onset [the stimulus onset asynchrony (SOA)]. The typical effects of the two types of cues are depicted in Fig. 2. The top panel of Fig. 2 shows the time course of a

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nonpredictive peripheral luminance cue on summoning reexive exogenous attention, and the bottom of Fig. 2 shows the time course of a predictive central arrow. These results were achieved in a covert attention experiment, with no movement of the eyes. The two cues are similar in the sense that in both types of cues the facilitory effects begin at 50 msec. However, there are differences. With endogenous cueing, the facilitory effect appears to be more sustained. With peripheral cueing, the advantage at the cued location changes, after a few hundred milliseconds, into an inhibition resulting in longer RTs for the cued location. No such inhibition is seen with endogenous cueing. These features of Fig. 2 will be discussed next, each related to a different mechanism of inhibition generated by the deployment of spatial attention.

A. Noise Reduction and Suppression of Competing Stimuli

Figure 2 shows that for about 200 msec after an exogenous cue and for 600 msec or longer following an endogenous cue, RT to detect the onset of a target is shorter when a target appears at the valid cue location compared to the invalid cue location. This RT effect indicates that attention has been drawn to the location of the cue, thereby facilitating the detection of the target at the valid location compared to the invalid location. It is generally agreed that the presence of attention due to valid cueing increases the signal-to-noise ratio or sensitivity of information processing at the cued location. Of course, this could happen either if the signal from the attended location were enhanced (a facilitatory process) or if noise that might interfere with that signal were reduced or suppressed (an inhibitory process). The available evidence supports a combination of these two effects. First, valid cueing can increase sensitivity even when a single target appears in an otherwise blank visual eld. Since there are no other stimuli around to produce any noise or distraction, it would appear that attention is enhancing the signal from the attended location. However, the effect of cueing a target in an otherwise blank eld is often small and sometimes disappears. The impact of cueing is usually greater when the target is accompanied by distracting stimuli at other spatial locations. From such evidence it has been argued that the primary contribution of spatial attention, especially in real-world visual environments, is to reduce noise or cross talk from unattended

stimuli that if unsuppressed would interfere with processing the attended stimulus. Direct evidence for inhibition of cross talk-based interference from unattended stimuli comes from single-cell recording in extrastriate cortical areas of the ventral object-processing pathway of monkeys. When two stimuli are present in the receptive eld of a cell in one of these areas (e.g., V4 or TEO), the cells response is less than if only one of them was present. This reduction in activity shows that the two stimuli interfere with each anothers ability to activate the cell. However, if the monkey attends to one of the stimuli for the purpose of performing an experimental task, the cells response returns to approximately the same level as if the attended stimulus were presented alone. This restoration of responsiveness shows that the unattended stimulus is being ltered out of the cells receptive eld and hence no longer interferes with processing the other stimulus. Analogous evidence that humans have a similar attention-driven mechanism of noise reduction through suppression of competing stimuli comes from functional magnetic resonance imaging (fMRI) experiments reported by Kastner and colleagues.

B. Inhibition of Return
A second important feature of Fig. 2, (seen only in the top), is that when SOAs following an exogenous cue are longer than 2300 msec RT to detect a target is longer when a target appears at the valid location compared to the invalid location. That is, facilitation is transformed into inhibition. This is a standard outcome of experiments with nonpredictive cues, in which subjects have no reason to use the cue to guide attention and would prefer to keep attention focused at xation or spread diffusely across the display. Michael Posner and Yoav Cohen analyzed the effect of nonpredictive cues at the longer SOA durations, now known as the inhibition of return (IOR) phenomenon. Facilitation changes to inhibition (i.e., IOR) 200300 msec after cue onset. IOR lasts for about 3 or 4 sec; it works in environmental rather than in retinal coordinates, it is not generated by endogenous cues unless the oculomotor system has been activated, and it declines as the distance from the original cued location increases. What is the explanation for this counterintuitive transformation of facilitation into inhibition? Exogenous cueing commonly produces a reexive shift of attention to the cued location or object (producing the

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early facilitation in Fig. 2). Even when people are asked not to pay attention to the cued location or it is in their favor to ignore the cued location, they nd it difcult to avoid reacting to a peripheral luminance change and often cannot refrain from orienting to this kind of cue. As much as such efcient and rapid orienting is important for predatory and defensive behavior, voluntary control of reexive orienting and the ability to strategically search the environment are also critical for survival. It appears that IOR is a mechanism that enables the organism to disengage from reexive orienting and switch to the control of a more voluntary attentional system. How does it work? It seems that a location (or an object) that was recently cued or searched is tagged and IOR biases attention away from responding to events occurring at the tagged locations. Avoidance of tagged locations encourages search in new spatial locations. Accordingly, IOR seems to be a mechanism that supports efcient foraging behavior, which involves strategic search of the environment and use of knowledge about previous searched locations or objects. Several lines of evidence point to the midbrain superior colliculus (SC) as the neural substrate for the implementation of IOR. IOR is abnormal in patients with damage to the SC. This has been shown in patients with midbrain degeneration due to progressive supranuclear palsy and in a patient with unilateral lesion to the SC. IOR is preserved in the presence of hemianopia, in which only the extrageniculate pathways are available to process visual information. It is present in newborn infants in whom the geniculostriate pathway is not yet developed. It is generated asymmetrically in the temporal and nasal visual elds. The temporal hemield has stronger collicular representation than the nasal hemield; accordingly, for monocular presentations of stimuli, IOR is larger for stimuli presented to the temporal than to the nasal hemield. However, it seems that cortical structures play a role in the generation of IOR. In particular, the parietal lobe has been suggested as a structure that conveys the spatial coordinates of the tagged locations to the SC.

restored by ablation of the dorsal midbrain contralateral to the cortical lesion. He noted, This initial hemianopia is apparently due to depression of function of the colliculus ipsilateral to the cortical lesion, a depression maintained by inux of inhibition from the crossed colliculus. Thus, removal of the contralateral tectum, or splitting of the collicular commisure, abolishes this inhibition and allows the return of function in the ipsilateral colliculus, and with it the recovery from hemianopia. This phenomenon is called the Sprague effect. A reversed version of the Sprague effect can be found with lesions to the frontal lobe, which will be described later. The SC is involved in triggering reexive saccades, whereas cortical mechanisms are needed for generating voluntary saccadic eye movements under strategic guidance. Early work showed that patients with frontal cortex lesions have difculty executing saccades in response to verbal commands. When these patients scan pictures their eye movements are controlled by external stimuli rather than by instructions. In addition, lesions of the frontal lobes produce a decit in inhibiting reexive glances. The patients seem unable to resist moving their eyes in the direction of a peripheral stimulus even when instructed to prevent such eye movements. It appears that the frontal eye elds (FEFs) are critical components of the frontal circuitry underlying inhibition of reexive saccades and endogenous control of eye movements. Unilateral lesions of the FEF may shorten latencies of reex saccades to targets in the contralesional eld. Moreover, single cell recordings show that the FEF contains cells that respond in temporal correlation with purposive saccades even in the absence of an exogenous visual stimulus. In line with these reports, Henik, Rafal, and Rhodes found that lesions to the FEF have opposite effects on endogenously activated and visually guided saccades to external stimuli. In a typical trial of an experiment examining saccades in FEF patients, subjects saw a visual cue (get ready) followed by a target (go). The cue was informative (a small arrow) or neutral (a double-headed arrow). There were two types of target go signals: an asterisk appearing at the periphery (used to elicit reexive, exogenously triggered saccades) or a large arrowhead in the center of the display pointing left or right (used to measure voluntary, endogenously generated saccades). The efcacy of saccade

C. Inhibition of Reexive Orienting

As suggested previously, other cortical structures in addition to the parietal lobes regulate activation of the SC. In 1966, James Sprague reported that occipitotemporal cortical ablation in cats produced stable hemianopia. However, visually guided behavior was

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preparation was measured as facilitation in saccade latency in the informative cue condition compared to the neutral cue condition. Patients with FEF lesions presented slower endogenous saccades (in response to the central arrow) to the contralesional eld, whereas exogenously triggered saccades (in response to the peripheral targets) were faster to the contralesional eld (a reversed Sprague effect). These results indicate that the FEF is involved in generating endogenous saccades and therefore lesions in this region increase their latency. In addition, the FEF is involved in visually guided saccades through inhibitory connections to the SC. It seems that the FEF has the opposite effect on the extrageniculate visual system from occipital lesions. That is, whereas occipital lesions reduce activation of the ipsilesional SC (producing increased activation of the contralateral SC), FEF lesions disinhibit the ipsilesional colliculus, producing suppression of collicular function contralateral to the cortical lesion. Taken together, this evidence indicates hierarchical control over eye movements. A relatively automatic system depending on SC is stimulus driven, responding to the appearance of new objects in the visual eld and to movement, especially in the periphery. This system is also sensitive to sound via interactions between superior and inferior colliculus. Barry Stein and colleagues have shown that reexive saccades and head turning are elicited when a movement that is subthreshold for triggering an orienting response is accompanied by a sound from the same spatial location that is also too weak by itself to cause orienting. The second level of control is cortical, depending on FEF. It exerts endogenous control rather than responding reexively to stimulus inputs, implementing voluntary eye movements, and inhibiting the responses of the automatic system when such responses would conict with the intended voluntary movement. Interactions between automatic orienting and voluntary control have been studied extensively in the antisaccade task. This task requires subjects to move their eyes away from a stimulus that appears abruptly in the visual eldFa stimulus that would ordinarily attract a reexive saccade. Endogenous control is far from perfect in this difcult task. Errors occur in which the eyes move toward the abruptly appearing stimulus, and changes in error rate with experimental manipulations and subject characteristics can be used to diagnose the effectiveness of endogenous control. More evidence is provided by the latencies of correct responses away from the stimulus, which are slower than either reexive saccades toward the same stimulus

or voluntary saccades toward stimuli that do not elicit reexive saccades (either because they are already present in the visual eld or because they are new but their onset is ramped up slowly to eliminate visualonset transients). In general, factors that increase the perceptual salience of the stimulus, such as brightness, tend to increase errors and slow latencies, as do factors that reduce the subjects ability to concentrate on the task, such as attentional load, and factors that reduce the subjects ability to predict when effortful control will be needed, such as randomly varying foreperiods prior to stimulus onset. Furthermore, patients with frontal lobe damage have great difculty with the antisaccade task.

III. SELECTION FOR ACTION AND CONFLICT RESOLUTION

The antisaccade task involves more than inhibiting the reexive response toward the abruptly appearing stimulus. The subject must successfully implement another response at the same time the reex is being inhibited. This second response is less well learned or less natural than the one that must be inhibited. Hence, the competition created by the automatic response is powerful, and resolving the conict in favor of the less well-learned response is difcult. The antisaccade task is one example of a common method for studying the regulatory processes of cognitive control. This method involves creating a conict situation in which the subject has to respond to one stimulus or to one aspect of the stimulus and ignore another stimulus or another aspect of the stimulus. In these situations, the subject needs to focus on the target (a stimulus or an aspect of a stimulus) and ignore all the rest of the display. Failures in attention are commonly revealed in two ways: (i) reduction in efciency of responding to the target when the irrelevant features of the display are present and (ii) indications for processing of the irrelevant material, especially when it clearly interferes with processing of the target. The two most widely used paradigms for studying this type of selection are Stroop color naming and negative priming.

A. Stroop Color Naming

J. Ridley Stroop, a theologist with a side interest in psychology, sought an experimental method that would enable him to measure the interference of one stimulus dimension on attempts to process another. In

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1935, he published a seminal paper describing three experiments. The second experiment asked subjects to name the color of the ink of color words (e.g., the word red printed in green ink) or the color of colored squares. The rst condition is called incongruent and the second neutral. Stroop found that the words interfered with naming the color. That is, RT to the incongruent condition was slower than RT to the neutral condition. Later, researchers added a congruent condition to the task (e.g., the word green printed in green ink). The congruent condition enabled one to look at facilitation, which is the difference in RT between congruent and neutral trials. It is commonly found that although facilitation in the congruent condition is small and on many occasions not signicant, interference in the incongruent condition is robust and signicant. Since 1935, this type of conict has been studied extensively using variations of Stroops color-naming task and in other Stroop-like situations with a wide variety of different stimuli and task demands. All studies converge on the conclusion that people cannot suppress the irrelevant dimension if it is heavily practiced and consequently overlearned (like reading words). Hence, the Stroop effect is considered a powerful example of automatic processing. Nevertheless, several studies have shown that readers can modulate and partially control the impact of the word. Increasing the proportion of congruent trials relative to incongruent trials produces a larger Stroop effect. Moreover, even when the numbers of congruent and incongruent trials are kept constant while the proportion of neutral trials changes, the effect can be altered. It seems that the expectation to face a relatively large proportion of conict trials prompts the adoption of a strategy that helps reduce interference. In addition, it seems that language competence may modulate the effect. When bilinguals are tested, under certain conditions they can reduce the effect in their rst language but not in their second language. Note that they experience interference in both their rst and their second language, but they are better able to control reading (i.e., reduce Stroop interference) in the language in which they are more competent. Neuropsychological studies of brain-injured individuals suggest that the left frontal lobe is crucial to successful performance of the Stroop task. In particular, it has been reported that injury to the left dorsolateral prefrontal cortex results in enlarged Stroop interference. This result suggests that the Stroop interference presented by noninjured individuals is an underestimate of the potential interference.

Stroop interference presented by the noninjured individuals is the product of automatic intrusion of the irrelevant word and their ability (admittedly not perfect) to inhibit this reading. In addition, it points to the involvement of the left dorsolateral prefrontal cortex in the control processes by which Stroop interfence is modulated. Consistent with this lesion evidence, neuroimaging studies of blood ow and changes in blood oxygenation during task performance show that the incongruent condition of the Stroop task activates left dorsolateral prefrontal cortex more than the neutral or congruent conditions. Even more noticeable in neuroimaging studies is differential activation of the anterior cingulate gyrus. Barch, Braver, Sabb, and Noll suggested that this medialfrontal structure is also active in a variety of other tasks in which selections must be made among competing stimuli, stimulus properties, and responses to them. Of course, the crucial question in the present context is whether resolution of conict in the Stroop task involves inhibition. Many theorists interpret the task in this way, although a well-known computational model of Stroop performance suggested by Cohen, Dunbar, and McClelland is able to account for many aspects of performance in the Stroop task by facilitation of the less automated process of color naming rather than inhibition of the more automated process of word reading. The phenomenon discussed next offers a more demanding and therefore more analytic test.

B. Negative Priming
In the incongruent or conict condition of Stroop color naming, an additional slowing of performance is observed, over and above the usual interference, if the color name to be produced on any given trial is the same as the color word that had to be ignored on the immediately preceding trial. This effect, which has been dubbed negative priming, can be found in a wide variety of task situations in which two stimuli occur on each trial, one to be ignored and the other requiring a response. When a just-ignored distractor becomes the target on the next trial, responding is slowed relative to not having ignored the current target item in the recent past. Similar slowing occurs when there is just one stimulus on each trial to which subjects must produce a newly learned arbitrary response. Suppose that subjects must say car whenever they see bike or a picture of a bike, plane whenever they see car or a picture of a car, and boat whenever they see plane

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or a picture of a plane. Now suppose that as a prime the subject sees car and correctly says plane. If the succeeding target is bike, production of the correct response carFthe overlearned response that had to be avoided when processing the primeFwill be slower than if the prime was an unrelated stimulus that required an unrelated arbitrary response, such as planeboat. The dominant interpretation of negative priming invokes a selective process that occurs while processing the prime. To respond appropriately, subjects must select against the distracter item that is the nonimperative component of the prime (or, in a task such as that of Shiu and Kornblums just described, they must select against the overlearned but contextually inappropriate response that the prime tends to elicit). The act of selection leaves the distractor or the overlearned response in a state that makes it more difcult to process if it recurs as the target. Two sorts of hypotheses have been offered about how this act of selection could be implemented. According to the distractor inhibition hypothesis, attentional operations actively inhibit either the primes perceptually activated mental representation, in order to prevent it from competing for access to response selection operations, or the link between the primes mental representation and the action ordinarily associated with the prime, in order to make that response unavailable. Inhibition takes time to dissipate. Therefore, if the inhibited representation or response link is needed soon thereafter for processing a target, more time and effort will be required for its activation. George Houghton and Steven Tipper constructed a simulation model embodying such processes. According to an alternative proposal, the episodic retrieval hypothesis of W. Trammell Neill and colleagues, attentional operations mark the distractor item with a do not respond to this stimulus tag or the overlearned response with a do not produce this response tag. The tag provides an instruction that guides decision and response selection, and it remains a part of the experience of having processed the prime that is stored in episodic memory. When the target appears it acts as a cue to retrieve this memory. The tag that served the subject well when the tagged stimulus or response needed to be ignored causes confusion and interferes with performance when the tagged item becomes the target. Note that the episodic retrieval hypothesis does not propose inhibition in the classic sense of reduction or suppression of activation. Its inhibitory process is

symbolic rather than analog, acting through an inuence on a mental representations informational content rather than its level of activity. Considerable effort has been expended attempting to distinguish these two underlying mechanisms by which an overtly inhibitory behavioral outcome might arise. Arguments have begun to appear that both mechanisms may be at work, and partly as a solution to the dilemma, in 1998 Milliken et al. made an important attempt to reinterpret negative priming in terms of the difculty of deciding whether retrieved episodic memories of past processing should or should not be used to guide current performance rather than whether they have been inhibited or tagged with negative content. This debate illustrates a crucial point mentioned earlier. Inhibition of behavior (i.e., slowing of overt performance) does not necessarily signal inhibition of mental processing, if what one means by inhibition of mental processing is reduction or suppression of activation. Considerable theoretical analysis and empirical investigation are often required to make this determination.

IV. PRIMING AND RETRIEVAL FROM MEMORY

It is often possible to retrieve information from memory at will (though sometimes, of course, such intentional attempts at retrieval fail). At the same time, there are often occasions when thoughts come to mind without any apparent intention. Here again, we are dealing with two fundamental aspects of cognition: controlled and automatic processing. In the domain of retrieval of information from memory (general knowledge as well as specic episodes of experience), these two aspects of cognition have been investigated by exploiting priming effects. Priming consists of an alteration in the speed or accuracy of responding to a stimulus such as a word or object due to a previous encounter with that stimulus or with related stimuli.

A. Repetition Priming
Repetition priming refers to the change in responding to a word or an object as a result of a previous encounter with that same item, either in the same task or in a different task. Responding to words or objects is typically improved due to this previous experience (usually more so when the task as well as the stimulus item remains the same). For example, if a word appears for a second time in a task that requires reading the target aloud (naming task) or deciding whether the

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target is a word or a nonword (lexical decision task), responding is faster and more accurate than for words appearing for the rst time at the same level of overall practice in the task. In addition, repetition priming can inuence the probability of producing a previously encountered item as a response when a task allows multiple possible answers on each trial. For example, suppose that in the study phase of a two-phase experiment subjects are asked to study a series of words. After a delay that can range from minutes to hours they are given three-letter word stems and asked to complete each stem with the rst word that comes to mind (e.g., gre_ for green). In this word stem completion task, the repetition priming effect appears when subjects respond more frequently with words that had been studied earlier than with words that were not encountered in the study phase. As can be seen from the examples, repetition priming is commonly examined using tasks that do not require conscious recollection of past experience with the stimulus item to complete. Therefore, it potentially represents a nonconscious or unintended effect of that experience. Such effects are called implicit memory in the repetition priming literature. When it can be documented that repetition priming has occurred without conscious recollection, the priming is called implicit memory, and as such it represents another example of automatic processing. Brain imaging studies using positron emission tomography (PET) and fMRI have shown that priming is accompanied by reduced neural activity within areas that were initially activated to perform the task. Reduced neural activity was found in occipital visual cortex (Brodmanns area 19), left frontal cortex, and inferior temporal cortex. Whereas the reduced neural activity in visual cortex is specic to visually presented stimuli, the activity reductions in left frontal cortex occur regardless of cue modality (visual or auditory). The reduced activity within the left frontal cortex suggests an amodal priming effect that represents access to the meaning of the word rather than to its visual or orthographic representation. Analogous modality-specic and amodal effects have been observed with objects. Single cell recording in monkeys provides evidence about the neural mechanisms that might mediate these repetition priming effects, at least for nonverbal stimuli. Repeated experience with the same visual stimulus leads to suppression of neuronal responses in subpopulations of visual neurons. This repetition suppression was found in the delayed matching-tosample task. In these studies a monkey was presented

with a sample stimulus followed by a sequence of test stimuli. The animal was rewarded for indicating which test stimulus matched the sample. For example, the monkey was presented with the sequence A y B y C y A and was supposed to respond to the nal A. Under these conditions, the common type of neural response was suppressive, and it was graded by similarity. The more similar the test stimulus was to the sample, the more the neural response was suppressed. Moreover, repetition suppression was associated with item repetition, whether the repeated item was the target or a distractor. Repetition suppression was found to be stimulus specic and long-lasting. In addition to visual cortex, this effect was recorded in the inferior temporal cortex and also in some regions of the prefrontal cortex. Robert Desimone suggested that the reduction in cortical activation in human neuroimaging studies such as those described earlier was due to a repetition suppression effect such as that documented in monkeys. This repetition suppression effect at the neuronal level would result in a decrease in the total number of activated cells (and hence in a reduced demand for oxygenated blood, producing the signal change measured in PET and fMRI). This reduced population of neurons, according to Desimone, provides a sharpened stimulus representation. The prime tunes the population of neurons so that a selective subpopulation that carries the critical features of the stimulus gives a robust response when the stimulus recurs, whereas other neurons, which are probably related to other stimuli, are suppressed. The more selective representation allows for a more efcient responding upon the next encounter with the stimulus. Desimones argument recalls the words of Sir John Eccles quoted earlier.

B. Semantic Priming
Semantic priming arises because the brain makes use of relations among similar or related stimuli in addition to using past experiences with the same stimulus. In the basic version of the semantic priming paradigm, subjects are presented with two successive stimuli called the prime and the target. They are usually asked to respond overtly only to the target. When words are the stimuli, the task may be naming or lexical decision. Supposing that the target is the word nurse, the prime can be a related word (e.g., doctor), an unrelated word (e.g., bread), or a neutral stimulus (e.g., a row of Xs). Under these conditions, the

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semantic priming or relatedness effect emerges. This effect can be described as a greater speed and accuracy of performance in the response to a target word when it is presented after a semantically related prime word than when it is presented after an unrelated prime word or after a neutral stimulus. This effect has been documented in a variety of situations. The semantic priming effect can occur when people are asked to pay attention to the prime and also when they do not pay attention to the prime, or even when they are unaware of its identity and do not phenomenologically realize that a prime has occurred. That is, semantic priming effects still exist when the target is presented very briey and masked by visual noise presented immediately following the prime so that people believe they have seen only the visual noise and are not aware of the presence of the prime. James Neely studied automatic and controlled processes by examining the ability of subjects to switch between semantic categories in an arbitrary fashion. He asked his subjects to think of parts of a building when the prime was the category name body (e.g., bodydoor). Subjects were able to follow the instructions when the time between the prime and the target (i.e., SOA) was long enough (e.g., 750 msec). That is, they responded faster to door following body relative to door following an unrelated prime (e.g., tree) or even a neutral prime (e.g., XXXX). However, they were not able to switch from one category to another when the SOA was short (e.g., 250 msec). What was the fate of the rejected category? That is, when the prime was body and the subject made an effort to think of the category building, what happened to parts of the body such as arm? Arm appearing after body was facilitated at short SOAs (o300 msec) and inhibited at long SOAs (between 500 and 2000 msec)Fthat is, RTs for related trials (bodyarm) were longer than RTs for neutral trials and equivalent to those for unrelated trials (birdarm). Hence, priming can be achieved both by unintentional automatic activation, as shown by priming from masked words of which subjects are unaware, and from consciously perceptible words at short SOAs even when subjects are trying to think of unrelated words. However, priming can also be achieved by intentionally focusing on a concept and generating possibilities following some rule, even an arbitrary rule as in Neelys switch condition, although such intentional focusing takes more time than automatic activation. Thus, these ndings support the existence of two mechanisms of semantic priming. The rst is

automatic, nonconscious, and can work without attention. By analogy to repetition priming, one might wonder if it is mediated by reduction of neural response in structures that store semantic knowledge. The second is voluntary, conscious, and occupies attention. One might expect that this mechanism would be associated with neural structures involved in the executive control operations of working memory, and that invoking this mechanism would increase neural activation in those structures. To date, however, we do not know of any neuroimaging studies of semantic priming.

C. Inhibitory Semantic Priming and the CenterSurround Theory of Retrieval Operations

Neelys evidence for inhibition of related words in his switch condition is one of the few instances in the literature in which semantic relatedness between successive stimuli harms performance in speeded tasks, such as naming or lexical decision. Another instance was reported by Dale Dagenbach and colleagues. They found that in certain circumstances lexical decisions following semantically related primes are slower than lexical decisions following unrelated primesFan absolute inhibition effect associated with semantic relatedness. This inhibitory priming occurs as a consequence of attempting to retrieve the meaning of a perceptually presented word when the meaning is weakly activatedFeither because the word is masked and hence perceptual input is easily confused with other input or because the word is newly learned and hence its representation in semantic memory is weak and easily confused with or overwhelmed by other representations. In either case, the weakly activated meaning is likely to suffer interference because other representations are activated that are similar but incorrect. Dagenbach and colleagues proposed that in such circumstances, the attempt to retrieve the weakly activated semantic code is accompanied by active inhibition of the related information that is producing the interference. This inhibition reects the operation of a center-surround attentional mechanism, which works to facilitate a semantic code on which it is focused or centered while inhibiting surrounding codes. These are codes that are similar or related to but different from the desired code and are competing with it for retrieval. The center-surround hypothesis predicts that repetition priming should be facilitatory at the same time that semantic priming is

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inhibitoryFa prediction that has been conrmed. Analogous ndings have been reported by Steven Lehmkuhle and colleagues in the domain of spatial rather than semantic processing, suggesting that inhibition of similar or closely related representations may be a generalizable strategy for conict resolution in the central nervous system.

V. EXECUTIVE CONTROL OF TASK PERFORMANCE: STOPPING AN ONGOING THOUGHT OR ACTION

Sometimes we start thinking about something or start to perform an action, only to realize it does not t with our primary goal or our general plan at the moment or that our goal has changed and it is no longer appropriate. Such situations call for a change in the course of thought or action. We must stop the current thought or action in order to be able to switch to another one. An experimental method for studying this common example of executive control is the stop signal paradigm developed by Gordon Logan. In this paradigm subjects are engaged in a primary task. In a relatively small proportion of trials they are signaled to stop before executing the response to the primary task. For example, participants can be asked to respond in each trial by pressing a button to an X and another button to an O. In addition, they are asked to withhold their response if a tone is presented at any point during a trial. The tone is presented in 25% of the trials at various delays after onset of the primary stimulus. Performance in this task can be successfully modeled as a race between a go process (perceiving and responding to the primary stimulus) and a stop process. The stop process is conceived to be a separate sequence of mental operations that involves perceiving the stop signal and intervening in the ongoing sequence of operations involved in the go process or primary task. If participants nish the stop process before the go process, they inhibit their response to the primary stimulus. If they nish the go process before the stop process, they produce a response to the primary stimulus, failing to inhibit it. RT to the go signal can be measured directly, whereas stop-signal RT (the time needed to cancel the planned response) cannot be measured directly (since its behavioral signature is the absence of action and hence there is nothing overt to measure). However, Logans race model provides quantitative methods for estimating the stop-signal RT. Research applying this model shows that young adults can stop a wide variety of actions (key presses,

hand movements, squeezes, and speech) very quickly, with an estimated latency from the stop signal of about 200 msec. Williams and colleagues reported that the ability to stop improves developmentally throughout childhood (i.e., stop-signal RT decreases) and then remains approximately constant across much of adulthood, falling off slightly but nonsignicantly in old age. Interestingly, speed of responding in most tasks that would be used as a primary task in this paradigmFthe go processFalso improves throughout childhood, but peak performance in young adulthood is followed by signicant slowing beginning in middle adulthood. Thus, there appears to be a difference between the developmental trajectory of the execution of primary tasks and that of the type of inhibitory control represented by stopping. This supports Logans hypothesis that processes governing inhibition are separate from those governing execution of speeded primary processing. Other research, however, shows that in old age the probability of stopping successfully does deteriorate, even if stop-signal RT remains fairly constant on those trials in which the stopping process succeeds. This suggests that old age may involve a loss of concentration in which inhibitory control processes are less likely to be implemented appropriately, even though they may still work effectively once deployed. Although this conclusion is consistent with the available data from the stopping paradigm, there is more to the relationship between age and inhibition. As will be discussed later, a major theory of cognitive aging proposes that most inhibitory functions decline in old age. In addition to developmental changes, the ability to inhibit ones actions in the stopping paradigm is related to some personal characteristics and individual differences. Stop-signal RT varies with impulsivity, being longer for more impulsive individuals. Hyperactive children have trouble stopping. Their stopsignal RT is longer than that of normal controls and they fail to inhibit responding on many occasions. This pattern is not due to a failure to detect the stop signal itself but rather to a decit in the inhibitory mechanism that implements stopping. Moreover, stimulant medication (methylphenidate) that improves behavioral symptoms of hyperactive children also improves their stopping performance. Event-related potentials suggest two loci at which the stop process exerts its impactFa central locus in frontal cortex that acts on motor planning and execution operations and a more peripheral process that acts on descending motor commands after they have left cortex. Evidence on the frontal locus comes

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from application of the stop-signal paradigm to examine gaze control in the FEF of monkeys by Jeffrey Schall and colleagues. Recording from single neurons in FEF showed that movement-related activity, which began to increase when a signal to move the eyes was presented, decreased when a signal to cancel the saccade was presented. The activity associated with this inhibition began to decrease before the stop-signal RT was over. It seems that the preparation of a movement is a controlled process composed of both execution and inhibition processes. FEF is involved in the generation of saccades, as mentioned earlier; in addition, neurons in the FEF can specify saccade cancellationFa specic and well-documented example of inhibitory function.

VI. DEVELOPMENTAL ASPECTS A. Development of Reaching and the A Not B Error

Studies of reaching behavior in human infants and monkeys suggest that development of such behavior involves both the ability to plan and execute sequences of action and the ability to inhibit certain reexive actions or dominant response tendencies. Piaget suggested that infants have difculty in understanding objects and their properties, including spatial relations. However, it seems that infants, even as young as 5 months old, can understand the object concept but they have difculty demonstrating this understanding by their reaching behavior. Reaching behavior of infants has been studied by Adele Diamond and colleagues using several paradigms. In one paradigm infants were presented with a Plexiglas box with a building block inside or outside the box. Infants were able to retrieve the object when a direct line of reach was possible (the object was in front of the box touching its front wall or in the middle of the box, away from its front wall). However, when the object was placed inside the box touching its front wall infants were unsuccessful in retrieving it. To retrieve the object in the latter situation there is a need to execute a sequence of two movements in order to avoid touching the front wall of the boxFone away from the object and a second in the direction of the object. Moreover, when the infants touched the front of the box they reexively grasped it or withdrew their hands. Sevenmonth-old infants rarely continued their reaching toward the object. In contrast, 10-month-old infants were much less likely to show these reexive behaviors

upon touching the edge of the box. These infants were able to retrieve the object in these circumstances. It seems that the older infants developed the ability to execute a reach that requires a change of direction and to inhibit reexive reactions of the hand. Infants also have difculty detouring around a barrier to retrieve an object. Here, the infants task is to retrieve the object from a transparent box that has an open side. Infants 6.57 months old reach straight through the side at which they are looking at the object. If they see the object through the open side, they can retrieve it. Otherwise, they cannot retrieve the object and they do not try alternative reaching behaviors. Toward the end of the rst year of life they can look through a closed side but retrieve the object from any open side of the box. In order to develop this ability infants need to inhibit the tendency to reach according to the line of sight. In another paradigm the infant is presented with an object in one of two places, A or B. The locations are covered to hide the object from sight and the infant is then allowed to search for it by lifting the covers. After the infant retrieves the object, the object is again placed in one of the two locations and the search task continues. Suppose the object is rst hidden at location A and the infant retrieves it successfully. If the object is then hidden at the second location B, the infant will often try to retrieve the object from A, even though the infant watched the experimenter hiding the object at B. This is the A not B error. Infants continue to make the A not B error from about 7.5 to 12 months of age, as long as the delay between hiding and retrieval is incremented as the infant gets older. Perhaps the most striking aspect of the phenomenon is that infants appear to know that the object is at B despite the fact that they reach toward A. Visual habituation and other visual memory tests indicate that infants remember the correct location. Sometimes, in the search task, the infant xates B while he or she is reaching toward A. Moreover, infants show the A not B error even when the covers are transparent and the object can be seen. In order to prevent this type of error there is a need to hold details of the task and the situation briey in short-term memory, and there is a need to inhibit the tendency to reach to A. The tendency to reach to A develops because reaching to A was reinforced earlier by successful nding of the object, and it increases with the number of times the infant has retrieved the object from location A before it is hidden at location B. The ability to meet both of these needs imposed by the task improves during infancy. Short-term memory improves as evidenced by the longer retention interval between

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hiding and allowing the infant to reach for the object needed to elicit the A not B error, and inhibition of the incorrect response tendency improves as evidenced by the eventual disappearance of the A not B error. It seems from this evidence that during the second half of the rst year of life infants begin to gain control over their reaching actions. They can inhibit interfering automatic tendencies and demonstrate planned and goal-directed control over manual behavior. The gradual ascendance of planned and goal-directed behavior over reexive or practiced stimulus-action routines has been modeled both by Diamond and colleagues and Stuart Marcovitch and Philip Zelazo as changes with age in the relative strength or dominance of competing systems, one like working memory and the other like conditioned or procedural learning. Evidence from lesions and single-cell recordings suggests that the increase in dominance of the working memory-like system is achieved, at least in part, through the maturation of several components of frontal cortex: the supplementary motor area (SMA) and the dorsolateral prefrontal cortex. Reexive grasping is released in adult humans following lesions of the SMA, and the same is true in the case of lesioned monkeys. Lesions of the dorsolateral prefrontal cortex in monkeys produce the A not B error and difculties inhibiting the urge to reach straight ahead to retrieve an object.

B. Development of Selection and Conict Resolution

It is possible to view the previously mentioned studies as conict situations in which habitual or endogenous tendencies compete for control of behavior. Throughout early childhood there is development in the ability to resolve such conict and select among competing stimuli, stimulus properties, and responses. Several researchers have suggested that central to this achievement is the development of the ability to effectively inhibit stimuli or associations that are irrelevant to the task. Moreover, it has been suggested that such a development relies on maturation of the frontal lobes. The Stroop task and its many variations have played a major part in the study of this development. For example, when preschool children are asked to say day to a picture of a moon and night to a picture of a sun, their accuracy is reduced relative to a neutral condition (e.g., responding day or night to a checkerboard). In addition, their accuracy decreases across the experimental session. Older children are

able to maintain above-chance accuracy throughout a session. Similar trends have been found in latency of responding. Other conict situations present similar results and document continued development of inhibitory function during childhood. For example, when Stroop color naming is examined across the school years, intrusion of the irrelevant word in place of the color name decreases in the incongruent or conict condition, as does the impact of the irrelevant word on latency of correct responding. Combining this evidence on conict resolution with the evidence on stopping discussed previously suggests a general improvement in the ability to inhibit prepotent and automated responses from infancy to adulthood. This ability frees the system from stimulusdriven control, enabling strategic control and planning to play a more dominant part in behavior. Moreover, it seems that such development is dependent on high levels of cortical maturation. Other trends in development are consistent with this idea that inhibitory function depends on cortical input. One example is IOR, discussed earlier. The SC, which seems to be the generator of IOR, is already developed in infancy. However, there appears to be a need for the parietal lobes to provide this system with the spatial coordinates necessary for producing IOR. Although in some circumstances IOR can be observed even in infants only a few days old, the appearance of robust and widespread IOR in the eye movement patterns of infants awaits parietal development, rather than depending only on maturation of the SC.

C. Cognitive Aging
Lynn Hasher, Rose Zacks, and colleagues have amassed a considerable body of evidence that inhibitory functions decline in old age. As a consequence, the cognitive processes of older adults are increasingly susceptible to interference from irrelevant or unwanted perceptions, thoughts, and tendencies toward action that are more successfully ignored by younger adults. Hasher, Zacks, and May suggested that the deleterious impact of interference appears to occur in addition to the generalized slowing observed in a wide variety of speeded task performances. Older adults produce more errors and slower correct responses in antisaccade tasks. They suffer greater interference in conict resolutions tasks such as Stroop color naming, and they nd it more difcult to inhibit primary task performance in the stopping paradigm. They suffer greater proactive interference in short-term memory

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tasks. They make more errors and retrieve answers more slowly in associative learning tasks when more than one response item is associated with a stimulus item (and in analogous fan effect fact-learning tasks in which multiple facts are learned about a single entity or topic item). They show less evidence of intentional or controlled forgetting in directed forgetting tasks, and they show less evidence of inhibiting irrelevant or inappropriate senses of ambiguous words and interpretations of ambiguous referring expressions in sentence reading and text comprehension. In many area of cognitive activity, older adults are susceptible to what Hasher, Zacks, and colleagues call mental clutter. Given the role played by prefrontal cortex in a number of the inhibitory functions reviewed in this article, one might wonder whether aging is particularly damaging to the integrity or efciency of processing in frontal tissue. There is considerable evidence to support such an idea. There are factors that moderate the impact of aging on cognitive activities with strong inhibitory components. One is practice, with task- and stimulus-specic practice being the most effective. Another is the level of circadian arousal. It is well-known that arousal levels vary systematically with time of day, and that individual differences in the time of occurrence of periods of optimal arousal create morning people and evening people. These individual differences extend to task performance. Both younger and older adults perform a wide variety of tasks better during their optimal periods of arousal than during off-peak periods, and the impact of variation in circadian arousal is greater for older adults. Indeed, if testing is done in the morning when older adults are more likely to be in an optimal period and younger adults are more likely to be in an off-peak period, age differences in task performance are minimized and in some cases nearly eliminated. Testing in the evening, when older adults are likely to be in an off-peak period and younger adults in an optimal period, exaggerates age differences. Thus, circadian variation in locus coeruleus activity and right-prefrontal activity related to vigilance and preparation interact with other aginginduced changes in cortical efcacy. Finally, there are a number of tasks in which performance does not appear to decline with age, at least in healthy adults free of nervous system insult such as stroke or Alzheimers disease. Many of these are related to language use. Vocabulary scores continue to increase with age, although word-nding problems during real-time speech production and

question answering may also increase. Sentence completion scores in ll-in-the-blank cloze tasks are not affected by age, nor is the accuracy of semantic categorization. All these language tasks that show little or no decline in old age are also affected little if at all by variation in circadian arousal. Perhaps the most surprising of the spared language functions is the ability to select one out of several possible and hence competing syntactic structures for application during real-time sentence production, which has been studied by Douglas Davidson in Zacks laboratory. Here, the integrity of older adults performance extends to speed as well as accuracy, violating both of the two bestestablished outcomes of cognitive agingFsusceptibility to interference in conict resolution situations and generalized slowing in speeded performances of many kinds. Thus, as has often been argued, language skills represent a domain-specic specialization whose operating principles seem to depart from those in many other areas of human cognition.

VII. CONCLUSIONS
Inhibitory functions are part and parcel of the cognitive processes that generate and control behaviors designed to provide specic solutions to dealing with environmental demands. In reviewing examples of inhibitory functions, we have seen that inhibition helps create coherent experience of the world along with the exibility and efciency required for skilled behavior. The ability to inhibit prepotent or reexive attentional and behavioral reactions and to stop unnecessary or inappropriate behavior develops throughout childhood. These developments free us from interference by otherwise dominant tendencies. This, in turn, enables us to exercise choice and intention over our actions. It seems that such changes are achieved through the development of various brain structures such as regions of the frontal lobes, including anterior cingulate, SMA, dorsolateral prefrontal cortex, and FEF. In doing their jobs, these control structures may interact or cooperate with regions of orbitofrontal cortex and amygdala involved in emotional regulation and sensitivity to delayed and long-term reinforcement contingencies. Note, however, that the ability of higher cortical structures to control reexive behaviors by inhibiting them is not the only development that takes place. In some functional domains, inhibition already being produced by lower brain structures is made accessible to the inuence of higher levels in the system.

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An example of this type of development is IOR, which appears to occur in retinal coordinates early in development when SC alone is responsible for it, but it occurs in environmental and object-based coordinates once parietal cortex becomes sufciently mature to contribute. Here, inhibition is already produced by certain brain structures but the development of cortical mechanisms allows this inhibition to be modulated by the higher brain mechanisms. In addition, we presented results from memory and language processing suggesting that inhibition helps focus on an object or a concept by sharpening the activated representation. Moreover, it seems that this sharpening can sometimes proceed automatically, with no involvement of attention, and can be achieved without awareness, at least in some circumstances. In other circumstances, deployment of inhibitory tuning mechanisms appears to be under the control of intentions to process a particular kind of information. In conclusion, inhibitory processes are ubiquitous in human cognition and vary in terms of the levels at which they operate and in terms of their relationship to various mental operations carried out in order to produce behavior. See Also the Following Articles
ANTERIOR CINGULATE CORTEX d ATTENTION d COGNITIVE PSYCHOLOGY, OVERVIEW d CONSCIOUSNESS d EMOTION d HOMEOSTATIC MECHANISMS d NEUROFEEDBACK d NEUROTRANSMITTERS d STRESS: HORMONAL AND NEURAL ASPECTS d SUPERIOR COLLICULUS d UNCONSCIOUS, THE

Suggested Reading
Barch, D. M., Braver, T. S., Sabb, F. W., and Noll, D. C. (2000). Anterior cingulate and the monitoring of response conict:

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