Dorsolateral prefrontal cortex

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Dorsolateral prefrontal cortex
Prefrontal1.png
An illustration of brain's prefrontal region
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Identifiers
Latin cortex praefrontalis dorsolateralis
MeSH D000087643
FMA 276189
Anatomical terms of neuroanatomy

The dorsolateral prefrontal cortex (DLPFC or DL-PFC) is an area in the prefrontal cortex of the primate brain. It is one of the most recently derived parts of the human brain. It undergoes a prolonged period of maturation which lasts into adulthood. [1] The DLPFC is not an anatomical structure, but rather a functional one. It lies in the middle frontal gyrus of humans (i.e., lateral part of Brodmann's area (BA) 9 and 46 [2] ). In macaque monkeys, it is around the principal sulcus (i.e., in Brodmann's area 46 [3] [4] [5] ). Other sources consider that DLPFC is attributed anatomically to BA 9 and 46 [6] and BA 8, 9 and 10. [1]

Contents

The DLPFC has connections with the orbitofrontal cortex, as well as the thalamus, parts of the basal ganglia (specifically, the dorsal caudate nucleus), the hippocampus, and primary and secondary association areas of neocortex (including posterior temporal, parietal, and occipital areas). [7] [8] The DLPFC is also the end point for the dorsal pathway (stream), [9] which is concerned with how to interact with stimuli.

An important function of the DLPFC is the executive functions, such as working memory, cognitive flexibility, [10] planning, inhibition, and abstract reasoning. [11] However, the DLPFC is not exclusively responsible for executive functions. All complex mental activity requires the additional cortical and subcortical circuits with which the DLPFC is connected. [12] The DLPFC is also the highest cortical area that is involved in motor planning, organization and regulation. [12]

Structure

As the DLPFC is composed of spatial selective neurons, it has a neural circuitry that encompasses the entire range of sub-functions necessary to carry out an integrated response, such as: sensory input, retention in short-term memory, and motor signaling. [13] Historically, the DLPFC was defined by its connection to: the superior temporal cortex, the posterior parietal cortex, the anterior and posterior cingulate, the premotor cortex, the retrosplenial cortex, and the neocerebellum. [1] These connections allow the DLPFC to regulate the activity of those regions, as well as to receive information from and be regulated by those regions. [1]

Function

Primary functions

The DLPFC is known for its involvement in the executive functions, which is an umbrella term for the management of cognitive processes, [14] including working memory, cognitive flexibility, [15] and planning. [16] A couple of tasks have been very prominent in the research on the DLPFC, such as the A-not-B task, the delayed response task and object retrieval tasks. [1] The behavioral task that is most strongly linked to DLPFC is the combined A-not-B/delayed response task, in which the subject has to find a hidden object after a certain delay. This task requires holding information in mind (working memory), which is believed to be one of the functions of DLPFC. [1] The importance of DLPFC for working memory was strengthened by studies with adult macaques. Lesions that destroyed DLPFC disrupted the macaques' performance of the A-not-B/delayed response task, whereas lesions to other brain parts did not impair their performance on this task. [1]

DLPFC is not required for the memory of a single item. Thus, damage to the dorsolateral prefrontal cortex does not impair recognition memory. [17] Nevertheless, if two items must be compared from memory, the involvement of DLPFC is required. People with damaged DLPFC are not able to identify a picture they had seen, after some time, when given the opportunity to choose from two pictures. [17] Moreover, these subjects also failed in Wisconsin Card-Sorting Test as they lose track of the currently correct rule and persistently organize their cards in the previously correct rule. [18] In addition, as DLPFC deals with waking thought and reality testing, it is not active when one is asleep. [18] Likewise, DLPFC is most frequently related to the dysfunction of drive, attention and motivation. [19] Patients with minor DLPFC damage display disinterest in their surroundings and are deprived of spontaneity in language as well as behavior. [19] Patients may also be less alert than normal to people and events they know. [19] Damage to this region in a person also leads to the lack of motivation to do things for themselves and/or for others. [19]

Decision making

The DLPFC is involved in both risky and moral decision making; when individuals have to make moral decisions like how to distribute limited resources, the DLPFC is activated. [20] This region is also active when costs and benefits of alternative choices are of interest. [21] Similarly, when options for choosing alternatives are present, the DLPFC evokes a preference towards the most equitable option and suppresses the temptation to maximize personal gain. [22]

Working memory

Working memory is the system that actively holds multiple pieces of transitory information in the mind, where they can be manipulated. The DLPFC is important for working memory; [23] reduced activity in this area correlates to poor performance on working memory tasks. [24] However, other areas of the brain are involved in working memory as well. [25]

There is an ongoing discussion if the DLPFC is specialized in a certain type of working memory, namely computational mechanisms for monitoring and manipulating items, or if it has a certain content, namely visuospatial information, which makes it possible to mentally represent coordinates within the spatial domain. [23]

There have also been some suggestions that the function of the DLPFC in verbal and spatial working memory is lateralised into the left and right hemisphere, respectively. Smith, Jonides and Koeppe (1996) [26] observed a lateralisation of DLPFC activations during verbal and visual working memory. Verbal working memory tasks mainly activated the left DLPFC and visual working memory tasks mainly activated the right DLPFC. Murphy et al. (1998) [27] also found that verbal working memory tasks activated the right and left DLPFC, whereas spatial working memory tasks predominantly activated the left DLPFC. Reuter-Lorenz et al. (2000) [28] found that activations of the DLPFC showed prominent lateralisation of verbal and spatial working memory in young adults, whereas in older adults this lateralisation was less noticeable. It was proposed that this reduction in lateralisation could be due to recruitment of neurons from the opposite hemisphere to compensate for neuronal decline with ageing. Overall, the DLPFC is complex and yet not fully understood.

Secondary functions

The DLPFC may also be involved in the act of deception and lying, [29] which is thought to inhibit normal tendency to truth telling. Research also suggests that using TMS on the DLPFC can impede a person's ability to lie or to tell the truth. [30]

Additionally, supporting evidence suggests that the DLPFC may also play a role in conflict-induced behavioral adjustment, for instance when an individual decides what to do when faced with conflicting rules. [31] One way in which this has been tested is through the Stroop test, [32] in which subjects are shown a name of a color printed in colored ink and then are asked to name the color of the ink as fast as possible. Conflict arises when the color of the ink does not match the name of the printed color. During this experiment, tracking of the subjects' brain activity showed a noticeable activity within the DLPFC. [32] The activation of the DLPFC correlated with the behavioral performance, which suggests that this region maintains the high demands of the task to resolve conflict, and thus in theory plays a role in taking control. [32]

DLPFC may also be associated with human intelligence. However, even when correlations are found between the DLPFC and human intelligence, that does not mean that all human intelligence is a function of the DLPFC. In other words, this region may be attributed to general intelligence on a broader scale as well as very specific roles, but not all roles. For example, using imaging studies like PET and fMRI indicate DLPFC involvement in deductive, syllogistic reasoning. [33] Specifically, when involved in activities that require syllogistic reasoning, left DLPFC areas are especially and consistently active. [33]

The DLPFC may also be involved in threat-induced anxiety. [34] In one experiment, participants were asked to rate themselves as behaviorally inhibited or not. Those who rated themselves as behaviorally inhibited, moreover, showed greater tonic (resting) activity in the right-posterior DLPFC. [34] Such activity is able to be seen through electroencephalogram (EEG) recordings. Individuals who are behaviorally inhibited are more likely to experience feelings of stress and anxiety when faced with a particularly threatening situation. [34] In one theory, anxiety susceptibility may increase as a result of present vigilance. Evidence for this theory includes neuroimaging studies that demonstrate DLPFC activity when an individual experiences vigilance. [34] More specifically, it is theorized that threat-induced anxiety may also be connected to deficits in resolving problems, which leads to uncertainty. [34] When an individual experiences uncertainty, there is increased activity in the DLPFC. In other words, such activity can be traced back to threat-induced anxiety.

Social cognition

Among the prefrontal lobes, the DLPFC seems to be the one that has the least direct influence on social behavior, yet it does seem to give clarity and organization to social cognition. [11] The DLPFC seems to contribute to social functions through the operation of its main specialty – the executive functions – for instance, when handling complex social situations. [11] Social areas in which the role of the DLPFC is investigated are, amongst others, social perspective taking [8] and inferring the intentions of other people, [8] or theory of mind; [11] the suppression of selfish behavior, [8] [35] and commitment in a relationship. [36]

Relation to neurotransmitters

As the DLPFC undergoes long maturational changes, one change that has been attributed to the DLPFC for making early cognitive advances is the increasing level of the neurotransmitter dopamine in the DLPFC. [1] In studies where adult macaques' dopamine receptors were blocked, it was seen that the adult macaques had deficits in the A-not-B task, as if the DLPFC was taken out altogether. A similar situation was seen when the macaques were injected with MPTP, which reduces the level of dopamine in the DLPFC. [1] Even though there have been no physiological studies about involvement of cholinergic actions in sub-cortical areas, behavioral studies indicate that the neurotransmitter acetylcholine is essential for working memory function of the DLPFC. [37]

Clinical significance

Schizophrenia

Schizophrenia may be partially attributed to a lack in activity in the frontal lobe. [18] The dorsolateral prefrontal cortex is especially underactive when a person has chronic schizophrenia.[ citation needed ] Schizophrenia is also related to lack of dopamine neurotransmitter in the frontal lobe.[ citation needed ] [18] The DLPFC dysfunctions are unique among the schizophrenia patients as those that are diagnosed with depression do not tend to have the same abnormal activation in the DLPFC during working memory-related tasks. [24] Working memory is dependent upon the DLPFC's stability and functionality, thus reduced activation of the DLPFC causes schizophrenic patients to perform poorly on tasks involving working memory. The poor performance contributes to the added capacity limitations in working memory that is greater than the limits on normal patients. [38] [ citation needed ] The cognitive processes that deal heavily with the DLPFC, such as memory, attention, and higher order processing, are the functions that once distorted contribute to the illness. [24] [ page needed ]

Depression

Along with regions of the brains such as the limbic system, the dorsolateral prefrontal cortex deals heavily with major depressive disorder (MDD). The DLPFC may contribute to depression due to being involved with the disorder on an emotional level during the suppression stage. [39] While working memory tasks seem to activate the DLPFC normally, [40] its decreased grey matter volume correlates to its decreased activity. The DLPFC may also have ties to the ventromedial prefrontal cortex in their functions with depression. [39] This can be attributed to how the DLPFC's cognitive functions can also involve emotions, and the VMPFC's emotional effects can also involve self-awareness or self-reflection. Damage or lesion to the DLPFC can also lead to increased expression of depression symptoms. [41]

Stress

Exposure to severe stress may also be linked to damage in the DLPFC. [42] More specifically, acute stress has a negative impact on the higher cognitive function known as working memory (WM), which is also traced to be a function of the DLPFC. [42] In an experiment, researchers used functional magnetic resonance imaging (fMRI) to record the neural activity in healthy individuals who participated in tasks while in a stressful environment. [42] When stress successfully impacted the subjects, their neural activity showed reduced working memory related activity in the DLPFC. [42] These findings not only demonstrate the importance of the DLPFC region in relation to stress, but they also suggest that the DLPFC may play a role in other psychiatric disorders. In patients with post-traumatic stress disorder (PTSD), for example, daily sessions of right dorsolateral prefrontal repetitive transcranial magnetic stimulation (rTMS) at a frequency of 10 Hz resulted in more effective therapeutic stimulation. [43]

Substance use

Substance use disorders (SUD) may correlate with dorsolateral prefrontal cortex dysfunction. [44] Those who recreationally use drugs have been shown to engage in increased risky behavior, possibly correlating with a dysfunction of the DLPFC. The executive controlling functions of the DLPFC in individuals who recreationally use drugs may have a weaker connection from risk factoring areas such as the anterior cingulate cortex and insula. [44] This weaker connection is even shown in healthy subjects, such as a patient who continued to make risky decisions with a disconnect between their DLPFC and insula. Lesions of the DLPFC may result in irresponsibility and freedom from inhibitions, [45] and the use of drugs can invoke the same response of willingness or inspiration to engage in the daring activity.

Alcohol

Alcohol creates deficits on the function of the prefrontal cortex. [46] As the anterior cingulate cortex works to inhibit any inappropriate behaviors through processing information to the executive network of the DLPFC, [46] as noted before this disruption in communication can lead to these actions being made. In a task known as Cambridge risk task, SUD participants have been shown to have a lower activation of their DLPFC. Specifically in a test related to alcoholism, a task called the Wheel of Fortune (WOF) had adolescents with a family history of alcoholism present lower DLPFC activation. [44] Adolescents that have had no family members with a history of alcoholism did not exhibit the same decrease of activity. [ citation needed ]

See also

Related Research Articles

Working memory is a cognitive system with a limited capacity that can hold information temporarily. It is important for reasoning and the guidance of decision-making and behavior. Working memory is often used synonymously with short-term memory, but some theorists consider the two forms of memory distinct, assuming that working memory allows for the manipulation of stored information, whereas short-term memory only refers to the short-term storage of information. Working memory is a theoretical concept central to cognitive psychology, neuropsychology, and neuroscience.

<span class="mw-page-title-main">Precuneus</span> Region of the parietal lobe of the brain

In neuroanatomy, the precuneus is the portion of the superior parietal lobule on the medial surface of each brain hemisphere. It is located in front of the cuneus. The precuneus is bounded in front by the marginal branch of the cingulate sulcus, at the rear by the parieto-occipital sulcus, and underneath by the subparietal sulcus. It is involved with episodic memory, visuospatial processing, reflections upon self, and aspects of consciousness.

<span class="mw-page-title-main">Frontal lobe</span> Part of the brain

The frontal lobe is the largest of the four major lobes of the brain in mammals, and is located at the front of each cerebral hemisphere. It is parted from the parietal lobe by a groove between tissues called the central sulcus and from the temporal lobe by a deeper groove called the lateral sulcus. The most anterior rounded part of the frontal lobe is known as the frontal pole, one of the three poles of the cerebrum.

<span class="mw-page-title-main">Brodmann area 9</span> Part of the frontal cortex in the brain of humans and other primates

Brodmann area 9, or BA9, refers to a cytoarchitecturally defined portion of the frontal cortex in the brain of humans and other primates. Its cytoarchitecture is referred to as granular due to the concentration of granule cells in layer IV. It contributes to the dorsolateral and medial prefrontal cortex.

<span class="mw-page-title-main">Brodmann area 46</span> Brain area

Brodmann area 46, or BA46, is part of the frontal cortex in the human brain. It is between BA10 and BA45.

In the philosophy of mind, neuroscience, and cognitive science, a mental image is an experience that, on most occasions, significantly resembles the experience of "perceiving" some object, event, or scene but occurs when the relevant object, event, or scene is not actually present to the senses. There are sometimes episodes, particularly on falling asleep and waking up, when the mental imagery may be dynamic, phantasmagoric, and involuntary in character, repeatedly presenting identifiable objects or actions, spilling over from waking events, or defying perception, presenting a kaleidoscopic field, in which no distinct object can be discerned. Mental imagery can sometimes produce the same effects as would be produced by the behavior or experience imagined.

<span class="mw-page-title-main">Mesocortical pathway</span>

The mesocortical pathway is a dopaminergic pathway that connects the ventral tegmentum to the prefrontal cortex. It is one of the four major dopamine pathways in the brain. It is essential to the normal cognitive function of the dorsolateral prefrontal cortex, and is thought to be involved in cognitive control, motivation, and emotional response.

<span class="mw-page-title-main">Prefrontal cortex</span> Part of the brain responsible for personality, decision-making, and social behavior

In mammalian brain anatomy, the prefrontal cortex (PFC) covers the front part of the frontal lobe of the cerebral cortex. It is the association cortex in the frontal lobe. The PFC contains the Brodmann areas BA8, BA9, BA10, BA11, BA12, BA13, BA14, BA24, BA25, BA32, BA44, BA45, BA46, and BA47.

<span class="mw-page-title-main">Lobes of the brain</span> Parts of the cerebrum

The lobes of the brain are the four major identifiable regions of the human cerebral cortex, and they comprise the surface of each hemisphere of the cerebrum. The two hemispheres are roughly symmetrical in structure, and are connected by the corpus callosum. Some sources include the insula and limbic lobe but the limbic lobe incorporates parts of the other lobes. The lobes are large areas that are anatomically distinguishable, and are also functionally distinct. Each lobe of the brain has numerous ridges, or gyri, and furrows, sulci that constitute further subzones of the cortex. The expression "lobes of the brain" usually refers only to those of the cerebrum, not to the distinct areas of the cerebellum.

Frontal lobe epilepsy (FLE) is a neurological disorder that is characterized by brief, recurring seizures arising in the frontal lobes of the brain, that often occur during sleep. It is the second most common type of epilepsy after temporal lobe epilepsy (TLE), and is related to the temporal form in that both forms are characterized by partial (focal) seizures.

<span class="mw-page-title-main">Executive functions</span> Cognitive processes necessary for control of behavior

In cognitive science and neuropsychology, executive functions are a set of cognitive processes that support goal-directed behavior, by regulating thoughts and actions through cognitive control, selecting and successfully monitoring actions that facilitate the attainment of chosen objectives. Executive functions include basic cognitive processes such as attentional control, cognitive inhibition, inhibitory control, working memory, and cognitive flexibility. Higher-order executive functions require the simultaneous use of multiple basic executive functions and include planning and fluid intelligence.

Frontostriatal circuits are neural pathways that connect frontal lobe regions with the striatum and mediate motor, cognitive, and behavioural functions within the brain. They receive inputs from dopaminergic, serotonergic, noradrenergic, and cholinergic cell groups that modulate information processing. Frontostriatal circuits are part of the executive functions. Executive functions include the following: selection and perception of important information, manipulation of information in working memory, planning and organization, behavioral control, adaptation to changes, and decision making. These circuits are involved in neurodegenerative disorders such as Alzheimer's disease and Parkinson's disease as well as neuropsychiatric disorders including schizophrenia, depression, obsessive compulsive disorder (OCD), and in neurodevelopmental disorder such as attention-deficit hyperactivity disorder (ADHD).

<span class="mw-page-title-main">Ventromedial prefrontal cortex</span> Body part

The ventromedial prefrontal cortex (vmPFC) is a part of the prefrontal cortex in the mammalian brain. The ventral medial prefrontal is located in the frontal lobe at the bottom of the cerebral hemispheres and is implicated in the processing of risk and fear, as it is critical in the regulation of amygdala activity in humans. It also plays a role in the inhibition of emotional responses, and in the process of decision-making and self-control. It is also involved in the cognitive evaluation of morality.

In psychology and neuroscience, executive dysfunction, or executive function deficit, is a disruption to the efficacy of the executive functions, which is a group of cognitive processes that regulate, control, and manage other cognitive processes. Executive dysfunction can refer to both neurocognitive deficits and behavioural symptoms. It is implicated in numerous psychopathologies and mental disorders, as well as short-term and long-term changes in non-clinical executive control. Executive dysfunction is the mechanism underlying ADHD paralysis, and in a broader context, it can encompass other cognitive difficulties like planning, organizing, initiating tasks and regulating emotions. It is a core characteristic of ADHD and can elucidate numerous other recognized symptoms.

The effects of sleep deprivation on cognitive performance are a broad range of impairments resulting from inadequate sleep, impacting attention, executive function and memory. An estimated 20% of adults or more have some form of sleep deprivation. It may come with insomnia or major depressive disorder, or indicate other mental disorders. The consequences can negatively affect the health, cognition, energy level and mood of a person and anyone around. It increases the risk of human error, especially with technology.

A neurological look at race is multifaceted. The cross-race effect has been neurologically explained by there being differences in brain processing while viewing same-race and other-race faces. There is a debate over the cause of the cross-race effect.

<span class="mw-page-title-main">Fronto-cerebellar dissociation</span>

Fronto-cerebellar dissociation is the disconnection and independent function of frontal and cerebellar regions of the brain. It is characterized by inhibited communication between the two regions, and is notably observed in cases of ADHD, schizophrenia, alcohol use disorder, and heroin use. The frontal and cerebellar regions make distinctive contributions to cognitive performance, with the left-frontal activations being responsible for selecting a response to a stimulus, while the right-cerebellar activation is responsible for the search for a given response to a stimulus. Left-frontal activation increases when there are many appropriate responses to a stimulus, and right-cerebellar activation increases when there is a single appropriate response to a stimulus. A person with dissociated frontal and cerebellar regions may have difficulties with selecting a response to a stimuli, or difficulties with response initiation. Fronto-cerebellar dissociation can often result in either the frontal lobe or the cerebellum becoming more active in place of the less active region as a compensatory effect.

<span class="mw-page-title-main">Ventrolateral prefrontal cortex</span> Part of the prefrontal cortex of the brain

The ventrolateral prefrontal cortex (VLPFC) is a section of the prefrontal cortex located on the inferior frontal gyrus, bounded superiorly by the inferior frontal sulcus and inferiorly by the lateral sulcus. It is attributed to the anatomical structures of Brodmann's area (BA) 47, 45 and 44.

The anti-saccade (AS) task is a way of measuring how well the frontal lobe of the brain can control the reflexive saccade, or eye movement. Saccadic eye movement is primarily controlled by the frontal cortex.

The parieto-frontal integration theory (P-FIT) considers intelligence to relate to how well different brain regions integrate to form intelligent behaviors. The theory proposes that large scale brain networks connect brain regions, including regions within frontal, parietal, temporal, and cingulate cortices, underlie the biological basis of human intelligence. These regions, which overlap significantly with the task-positive network, allow the brain to communicate and exchange information efficiently with one another. Support for this theory is primarily based on neuroimaging evidence, with support from lesion studies. The P-FIT is influential in that it explains the majority of current neuroimaging findings, as well as increasing empirical support for cognition being the result of large-scale brain networks, rather than numerous domain-specific processes or modules. A 2010 review of the neuroscience of intelligence described P-FIT as "the best available answer to the question of where in the brain intelligence resides".

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