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Wednesday, June 21, 2023

Purkinje cell

From Wikipedia, the free encyclopedia
Purkinje cell
PurkinjeCell.jpg
Drawing of pigeon Purkinje cells (A) by Santiago Ramon y Cajal
Details
PronunciationOften pronounced as /pɜːrˈkɪni/ pur-KIN-jee; but Czech pronunciation is (Czech: [ˈpurkɪɲɛ] (listen) cells
LocationCerebellum
Shapeflat dendritic arbor
Functioninhibitory projection neuron
NeurotransmitterGABA
Presynaptic connectionsParallel fibers and Climbing fibers
Postsynaptic connectionsCerebellar deep nuclei
Identifiers
MeSHD011689
NeuroNames365
NeuroLex IDsao471801888
TA98A14.1.07.404
FMA67969

Purkinje cells, or Purkinje neurons, are a class of GABAergic inhibitory neurons located in the cerebellum. They are named after their discoverer, Czech anatomist Jan Evangelista Purkyně, who characterized the cells in 1839.

Structure

Neurons (Purkinje cells) located in the cerebellum
 
Transverse section of a cerebellar folium. (Purkinje cell labeled at center top.)
 
Silver stain of cerebellum showing Purkinje cells.
 
Purkinje cells. Bielschowsky stain.
 
Confocal microscope image of cerebellar Purkinje cells expressing tdTomato

These cells are some of the largest neurons in the human brain (Betz cells being the largest), with an intricately elaborate dendritic arbor, characterized by a large number of dendritic spines. Purkinje cells are found within the Purkinje layer in the cerebellum. Purkinje cells are aligned like dominos stacked one in front of the other. Their large dendritic arbors form nearly two-dimensional layers through which parallel fibers from the deeper-layers pass. These parallel fibers make relatively weaker excitatory (glutamatergic) synapses to spines in the Purkinje cell dendrite, whereas climbing fibers originating from the inferior olivary nucleus in the medulla provide very powerful excitatory input to the proximal dendrites and cell soma. Parallel fibers pass orthogonally through the Purkinje neuron's dendritic arbor, with up to 200,000 parallel fibers forming a Granule-cell-Purkinje-cell synapse with a single Purkinje cell. Each Purkinje cell receives approximately 500 climbing fiber synapses, all originating from a single climbing fiber. Both basket and stellate cells (found in the cerebellar molecular layer) provide inhibitory (GABAergic) input to the Purkinje cell, with basket cells synapsing on the Purkinje cell axon initial segment and stellate cells onto the dendrites.

Purkinje cells send inhibitory projections to the deep cerebellar nuclei, and constitute the sole output of all motor coordination in the cerebellar cortex.

Molecular

The Purkinje layer of the cerebellum, which contains the cell bodies of the Purkinje cells and Bergmann glia, express a large number of unique genes. Purkinje-specific gene markers were also proposed by comparing the transcriptome of Purkinje-deficient mice with that of wild-type mice. One illustrative example is the Purkinje cell protein 4 (PCP4) in knockout mice, which exhibit impaired locomotor learning and markedly altered synaptic plasticity in Purkinje neurons. PCP4 accelerates both the association and dissociation of calcium (Ca2+) with calmodulin (CaM) in the cytoplasm of Purkinje cells, and its absence impairs the physiology of these neurons.

Development

Mammalian embryonic research has detailed the neurogenic origins of Purkinje cells. During early development Purkinje cells arise in the ventricular zone in the neural tube, the nervous system´s precursor in the embryo. All cerebellar neurons derive from germinal neuroepithelia from the ventricular zone. Purkinje cells are specifically generated from progenitors in the ventricular neuroepithelium of the embryonic cerebellar primordium. The first cells generated from the cerebellar primordium form a cap over a diamond-shaped cavity of the developing brain called the fourth ventricle forming the two cerebellar hemispheres. The Purkinje cells that develop later are those of the cerebellum's center-lying section called the vermis. They develop in the cerebellar primordium that covers the fourth ventricle and below a fissure-like region called the isthmus of the developing brain. Purkinje cells migrate toward the outer surface of the cerebellar cortex and form the Purkinje cell layer.

Purkinje cells are born during the earliest stages of cerebellar neurogenesis. Neurogenin2, together with neurogenin1, are transiently expressed in restricted domains of the ventricular neuroepithelium during the time-window of Purkinje cell genesis. This spatio-temporal distribution pattern suggests that neurogenins are involved in the specification of phenotypically heterogeneous Purkinje cell subsets, ultimately responsible for constructing the framework of the cerebellar topography.

There is evidence in mice and humans that bone marrow cells either fuse with or generate cerebellar Purkinje cells, and it is possible that bone marrow cells, either by direct generation or by cell fusion, could play a role in repair of central nervous system damage. Further evidence points yet towards the possibility of a common stem cell ancestor among Purkinje neurons, B-lymphocytes and aldosterone-producing cells of the human adrenal cortex.

Function

The Purkinje cell protein 4 (PCP4) is markedly immunoreactive in the Purkinje cells of the human cerebellum. From top to bottom 40X, 100X and 200X microscopic magnifications. The immunohistochemistry was performed based on published methods.
 
Microcircuitry of the cerebellum. Excitatory synapses are denoted by (+) and inhibitory synapses by (-).
MF: Mossy fiber.
DCN: Deep cerebellar nuclei.
IO: Inferior olive.
CF: Climbing fiber.
GC: Granule cell.
PF: Parallel fiber.
PC: Purkinje cell.
GgC: Golgi cell.
SC: Stellate cell.
BC: Basket cell.

Purkinje cells show two distinct forms of electrophysiological activity:

  • Simple spikes occur at rates of 17 – 150 Hz (Raman and Bean, 1999), either spontaneously or when Purkinje cells are activated synaptically by the parallel fibers, the axons of the granule cells.
  • Complex spikes are slow, 1–3 Hz spikes, characterized by an initial prolonged large-amplitude spike, followed by a high-frequency burst of smaller-amplitude action potentials. They are caused by climbing fiber activation and can involve the generation of calcium-mediated action potentials in the dendrites. Following complex spike activity, simple spikes can be suppressed by the powerful complex spike input.

Purkinje cells show spontaneous electrophysiological activity in the form of trains of spikes both sodium-dependent and calcium-dependent. This was initially shown by Rodolfo Llinas (Llinas and Hess (1977) and Llinas and Sugimori (1980)). P-type calcium channels were named after Purkinje cells, where they were initially encountered (Llinas et al. 1989), which are crucial in cerebellar function. Activation of the Purkinje cell by climbing fibers can shift its activity from a quiet state to a spontaneously active state and vice versa, serving as a kind of toggle switch. These findings have been challenged by a study suggesting that such toggling by climbing-fiber inputs occurs predominantly in anaesthetized animals and that Purkinje cells in awake behaving animals, in general, operate almost continuously in the upstate. But this latter study has itself been challenged and Purkinje cell toggling has since been observed in awake cats. A computational model of the Purkinje cell has shown intracellular calcium computations to be responsible for toggling.

Findings have suggested that Purkinje cell dendrites release endocannabinoids that can transiently downregulate both excitatory and inhibitory synapses. The intrinsic activity mode of Purkinje cells is set and controlled by the sodium-potassium pump. This suggests that the pump might not be simply a homeostatic, "housekeeping" molecule for ionic gradients. Instead, it could be a computation element in the cerebellum and the brain. Indeed, a mutation in the Na+
-K+
pump causes rapid onset dystonia parkinsonism; its symptoms indicate that it is a pathology of cerebellar computation. Furthermore, using the poison ouabain to block Na+
-K+
pumps in the cerebellum of a live mouse induces ataxia and dystonia. Numerical modeling of experimental data suggests that, in vivo, the Na+
-K+
pump produces long quiescent punctuations (>> 1 s) to Purkinje neuron firing; these may have a computational role. Alcohol inhibits Na+
-K+
pumps in the cerebellum and this is likely how it corrupts cerebellar computation and body co-ordination.

Clinical significance

In humans, Purkinje cells can be harmed by a variety of causes: toxic exposure, e.g. to alcohol or lithium; autoimmune diseases; genetic mutations causing spinocerebellar ataxias, gluten ataxia, Unverricht-Lundborg disease, or autism; and neurodegenerative diseases that are not known to have a genetic basis, such as the cerebellar type of multiple system atrophy or sporadic ataxias.

Gluten ataxia is an autoimmune disease triggered by the ingestion of gluten. The death of Purkinje cells as a result of gluten exposure is irreversible. Early diagnosis and treatment with a gluten-free diet can improve ataxia and prevent its progression. Less than 10% of people with gluten ataxia present any gastrointestinal symptom, yet about 40% have intestinal damage. It accounts for 40% of ataxias of unknown origin and 15% of all ataxias.

The neurodegenerative disease spinocerebellar ataxia type 1 (SCA1) is caused by an unstable polyglutamine expansion within the Ataxin 1 protein. This defect in Ataxin 1 protein causes impairment of mitochondria in Purkinje cells, leading to premature degeneration of the Purkinje cells. As a consequence, motor coordination declines and eventually death ensues.

Some domestic animals can develop a condition where the Purkinje cells begin to atrophy shortly after birth, called cerebellar abiotrophy. It can lead to symptoms such as ataxia, intention tremors, hyperreactivity, lack of menace reflex, stiff or high-stepping gait, apparent lack of awareness of foot position (sometimes standing or walking with a foot knuckled over), and a general inability to determine space and distance. A similar condition known as cerebellar hypoplasia occurs when Purkinje cells fail to develop in utero or die off before birth.

The genetic conditions ataxia telangiectasia and Niemann Pick disease type C, as well as cerebellar essential tremor, involve the progressive loss of Purkinje cells. In Alzheimer's disease, spinal pathology is sometimes seen, as well as loss of dendritic branches of the Purkinje cells. Purkinje cells can also be damaged by the rabies virus as it migrates from the site of infection in the periphery to the central nervous system.

Etymology

Purkinje cells are named after the Czech scientist Jan Evangelista Purkyně, who discovered them in 1839.

Cognitive effects of bilingualism

Bilingualism, a subset of multilingualism, means having proficiency in two or more languages. A bilingual individual is traditionally defined as someone who understands and produces two or more languages on a regular basis. A bilingual individual's initial exposure to both languages may start in early childhood, e.g. before age 3, but exposure may also begin later in life, in monolingual or bilingual education. Equal proficiency in a bilingual individuals' languages is rarely seen as it typically varies by domain. For example, a bilingual individual may have greater proficiency for work-related terms in one language, and family-related terms in another language.

Being bilingual has been linked to a number of cognitive benefits. Research on how a bilingual individual's first language (L1) and second language (L2) interact shows that both languages have an influence on the function of one another and on cognitive function outside of language. Research on executive functions like working memory, perception, and attentional and inhibitory control, suggests that bilinguals can benefit from significant cognitive advantages over monolingual peers in various settings. There are also age-related benefits which seem to protect against cognitive decline in older adults.

Throughout the history of research into the cognitive advantages of bilingualism, views have shifted from a subtractive to an additive perspective: it is now believed that being bilingual adds to an individual's abilities rather than subtracting from it.

There is, however, strong disagreement over how findings on this subject should be interpreted. Systematic reviews and meta-analyses of executive functioning studies have failed to find compelling evidence for cognitive advantages in healthy adults or in participants across a broader age range. Moreover, the distribution of effect sizes in meta-analyses suggest publication bias, or that the reporting of bilingualism effects on executive functioning give a distorted view of the evidence.

History

According to the Singapore Management University (SMU) School of Social Sciences, research before the 1960s on bilingual individuals varied but commonly supported the idea that there were disadvantages to bilingualism. Researchers believed that bilinguals would have smaller vocabularies and stunted cognitive abilities. They also thought that children learning two languages at a young age would struggle differentiating and building proficiency in two languages to become competent in either. The idea that being bilingual was harmful to a child's linguistic and cognitive development, persisted. According to a historical review in "The Journal of Genetic Psychology," various researchers held these beliefs, noting a "problem of bilingualism" or the "handicapping influence of bilingualism." Following studies reported that bilinguals performed worse in IQ tests and suffered in most aspects of language development. These perspectives on bilingualism may have come from studies that did not control for socioeconomic status (SES) and gave IQ tests to non-proficient speakers of a second language in that second language. Many of these studies also used unstandardized and subjective definitions of bilingualism and of a bilingual individual, labeling someone as bilingual or monolingual through assumptions based on parent national origin, or based on family name.

Researchers began to change tone in the late 1950s/early 1960s, when Lenneberg, Chomsky, and Halle co-founded the field of biolinguistics and explored the role of biology in language. Their ideas led others to consider the role of human development more. In 1962, a turning point came about from a study that emphasized the importance of controlling for factors like age, sex, and SES, as well as of having a standardized measure for bilingualism when selecting a sample of bilinguals to be studied. Researchers carefully matched bilingual to monolingual participants and found that the bilinguals appeared to have significant advantages to that of their monolingual peers, outperforming in both verbal and non-verbal tests, more specifically in the non-verbal tests. In continuation of this study, research after this point began to shift focus, investigating areas of cognitive development and aptitude like perception and executive functioning. In 1967, publication of Lenneberg's seminal book, Biological Foundations of Language, first introduced the idea of a critical period of language acquisition, now better known as a sensitive period and further influenced bilingualism ideas. In 1977 the American Institutes for Research published an influential study which discussed bilingualism as it relates to education - how it affects a child's performance compared to peers. This study played a large role in our understanding of multilingualism and the effects that it has on the brain.

Since the late 1970s, researchers have found more cognitive benefits of bilingualism, including better attention, task-switching, and protection against aging declines. Over time, the prevalence of bilinguals in the United States has also increased. While the United States Census Bureau does not directly poll for bilingualism, they do poll for household languages. Non-English-speaking households are also surveyed for English proficiency. A 2012 interpretation of Census Bureau results observed that 11% of the population was bilingual in 1980, 14% in 1990, and 20% in 2012.

Cognitive advantages

Executive function

Executive function is the group of high-level cognitive processes that assists in goal-oriented tasks, such as problem solving, mental flexibility, attentional control, inhibitory control, and task switching. Much of the current research on cognitive effects of bilingualism studies the potential relationship between bilingualism and executive function; monolinguals and bilinguals may have executive function differences. Studies that match age and background factors like SES have found that bilinguals, who experience the mental demands of learning multiple languages, may have greater executive control and experience other cognitive benefits than their monolingual peers. Executive function may also have effects for older adults. Adults who learned a second language at a young age show better control and inhibitory processing than monolingual adults, as well as greater protection from cognitive decline such as dementia. Notably, some scientists continue to debate whether a bilingual advantage in executive functioning exists, with some studies and meta-analyses coming to opposite conclusions.

Bimodal bilinguals

Unimodal bilinguals are individuals who are proficient in two spoken languages. In contrast, bimodal bilinguals are individuals who are proficient in languages in differing modalities (i.e. a spoken and signed language). Studies have indicated that unimodal and bimodal bilinguals face different factors and requirements for their language use. In a monolingual conversational situation, unimodal bilinguals must suppress their use of one of their languages more significantly than bimodal bilinguals, while using the other language. This is because the use of a spoken word in another language would be confusing to the listener, whereas the use of a signed word would be less distracting to a listener, even when they do not know the meaning of the sign.

In 1998, David Green proposed his "inhibitory control model", which references a bilingual's constant need to suppress one language while using another. Because this task requires suppressing a source of distraction, this kind of control is then applied to other tasks. This assertion was bolstered by a study comparing unimodal bilinguals and bimodal bilinguals. Because bimodal bilinguals can express themselves in both languages at the same time, they may require less inhibition. This idea was supported by the results of the study; only unimodal bilinguals were found to have an advantage, as measured by the flanker task (a cognitive task that measures attentional focus and inhibition). Bimodal bilinguals also switch languages less frequently, because they are more likely to use both languages at once than to completely switch from one to the other. For this reason, the researchers of this study hypothesized that it may be the switching between languages that gives unimodal bilinguals the advantage.

Bilingualism vs. executive control

Hakuta and Diaz addressed an origin question about the potential bilingual advantage: do children with greater cognitive abilities tend to learn more than one language, or could knowing more than one language contribute to enhanced cognition? They administered Raven's Progressive Matrices, a set of non-verbal tests designed to measure cognitive ability, to a bilingual sample of children. Results showed their sample had a higher correlation with the degree of bilingualism, or language proficiency, and test scores. Accordingly, this study suggested that the "egg came first," or that bilingualism predicts cognitive ability through performance. Furthermore, native bilinguals often learn a second language due to family environment, where use of the two languages is necessary. It is less likely that children learn second languages in monolingual households through individual, innate cognitive abilities.

The bilingualism advantage controversy

The idea that bilingualism enhances executive function is not universally accepted among researchers. Mixed experimental results have led to bitter, opposing views.

The research methodologies of both groups for and against bilingualism advantages have been disputed. Some opposing researchers argue that correlations between bilingualism and executive function are inconsistent due to a reliance on findings with small sample sizes, publication bias, and non-uniform and definitions. Furthermore, they argue that bilinguals are not consistently better at all executive function tasks and existing studies only compare bilinguals to monolinguals, and not bilinguals of different proficiencies. Conversely, researchers in support of a bilingualism advantage note that while tests of executive function may undermine the robustness of findings, opposing findings come from researchers outside of the bilingualism field, and new neuroimaging technologies robustly verify and build upon historical findings of a bilingualism advantage.

Parallel activation of both languages

Bilinguals often have more knowledge in certain language domains, leading to a dominant "L1" language, and a less dominant "L2" language. Research shows that when a bilingual individual proficient in both uses only L1 or L2, both languages are simultaneously active, phonologically and semantically, and share overlapping neural representations This activation is shown by electrophysiological measures of performance when listening to speech, reading words in either language or even planning speech in either language. Bilingualism studies historically looked at languages that share the Roman alphabet, where co-activation of languages may make more intuitive sense. However, co-activation of L1 and L2 has also been reported in bilinguals whose two languages have different scripts (writing systems), lexical forms (e.g. Japanese and English), and modality (e.g. English and American Sign Language).

The words "red, green, and blue" are shown in colors not associated with their names.
The Stroop task, which shows color names in a different color, tests inhibition and distraction.

The fact that both languages are constantly activated means that they potentially compete for cognitive resources; bilinguals need a way to control the competition, to not use the wrong language at the wrong time. Inhibition refers to being able to ignore irrelevant information and therefore not be distracted by non-target stimuli. Inhibition may be the executive control system that allows successful linguistic selection even when both languages are co-activated in bilinguals. De Groot & Christofells (2006) proposed a distinction between two types of inhibition that may occur: global inhibition and local inhibition. Global inhibition refers to suppression of an entire language system, e.g. inhibiting Spanish when speaking English, and local inhibition refers to inhibition of a more specific competing vocabulary, e.g. the translation of the same word or phrase. Local inhibition mostly affects linguistic performance whereas global inhibition affects both linguistic and cognitive performance.

Inhibiting language in different ways may impact non-linguistic and linguistic cognitive processing. For example, a test that is widely used to assess this executive function is the Stroop task, where the word for a color is printed in a different color than the name (e.g. the word 'red' printed in blue ink). This causes interference and distraction; reaction times are measured to see how distracted the individual is by the incongruent word and color. Bilinguals compared to monolinguals have shown better Stroop task performance, suggesting that bilinguals have a more developed inhibition process, potentially due to the constant inhibition of their non-target language.

Abilities related to executive function

Bilingual individuals have also shown advantages in metalinguistic ability, which seems closely tied to executive function. Metalinguistic awareness is the understanding of the separation between language's structure and its meaning. For example, being able to judge the grammaticality of a sentence regardless of whether it is sensical, or being able to separate the set of sounds comprising a word from the word's meaning. Metalinguistic awareness involves inhibition, or the ability to suppress distracting information such as semantics. Metalinguistic ability could also be exercised by being bilingual, given that a bilingual individual has to suppress their knowledge of another language system when operating in one of their languages.

Improved theory of mind, which is the ability to understand someone else's thinking, has also been implicated in bilingualism. A meta-analysis in 2018 found that bilingual children performed better on a theory of mind test compared to monolingual children.

Benefits in older age

There has been a growing research interest in the benefits of bilingualism against age-related cognitive decline. Klein & Viswanathan found that the normal decrease in attention control observed in older adults was reduced in bilinguals, suggesting that bilingualism may protect against age-related cognitive deficits. Elderly bilinguals show better task-switching skill, can ignore unrelated information, and can choose the best phrasing. Bilingualism may be one of the environmental factors which contributes to 'cognitive reserve'. Cognitive reserve is the idea that engaging in stimulating physical or mental activity can maintain cognitive functioning in healthy aging and delay the onset of dementia-related memory losses. Contributing factors include education, occupational status, higher socioeconomic class, and doing physical, intellectual and social activities.

To test the protection of bilingualism against Alzheimer's disease (AD), Bialystok et al. (2007) examined hospital records of monolingual and bilingual patients who were diagnosed with various types of dementia. After controlling for various cognitive factors, the researchers found bilinguals experienced later onset of symptoms and were diagnosed approximately 3–4 years later on average than monolinguals. This was replicated with patients all diagnosed with AD. Note that the studies did not show that bilingualism directly prevents one from having AD, but rather enables functional cognition for a longer period of time, delaying symptom onset for those with the disease. For example, in one Alzheimer study, bilinguals actually had more pathology (signs of disease) and damage than the monolingual patients. This suggests that active use of the two languages protects against the symptoms of the disease. Areas of the brain that enable cognitive control may have benefited from the bilingual experience and so improve cognitive function in older age.

Several other studies replicated the finding that bilingualism contributes to cognitive reserve. For example, Abutalebi et al. (2015) used the flanker test to study unimodal bilinguals and monolinguals matched based on education level and SES. While this study had a relatively small sample size, the results did confirm previous research findings: the bilinguals outperformed the monolinguals on all experimental tasks, and the monolinguals' neural imaging showed more signs of age-related effects on performance of tasks and less gray matter density, which is associated with memory and function losses. Conversely, the bilinguals' neural imaging showed more gray matter density.

Age of acquisition

A debate within the linguistic community is whether the age of acquiring one's L2 has effects on the cognitive advantages. Bilingual children who acquire an L2 early may develop executive function earlier, around age 3, compared to monolingual hearing children who develop executive function around age 4 or 5. Native bilingual children who develop proficiency in L1 and L2 at the same time, perform better on executive function tasks such as the Attention Network Test, which measures attentional control, compared to their late bilingual and monolingual counterparts in studies controlled for age, verbal ability, and SES.

Language

Language use

Leopold, a pioneer of child language and bilingualism research, made many research findings by observing his daughter's, Hildegard, language use. In his studies, he observed that Hildegard had "loose connections" between the (phonetic) structure of words and their semantics (meaning) because of her frequent substitutions of English words with German words and vice versa. This was noted in her everyday speech and well-rehearsed songs or rhymes. He noted that she had a greater flexibility in the use of language that was unobserved in monolingual children of her age. Leopold considered that perhaps this loose connection between the meaning and form of a word could result in more abstract thinking or greater mental flexibility for bilingual children. Following this study, several others were formed to test similar things and find out more about the mental abilities of bilinguals with relation to their languages.

Semantic development

Anita Ianco-Worrall, author of Bilingualism and Cognitive Development, designed a study to test Leopold's observations and was able to replicate them. She tested two groups of monolingual and bilingual children at ages 4–6 and 6–9. These participants were given tasks to assess whether they showed a semantic or phonetic preference when categorizing words. An example of one task given in the study was to decide which of the two words, either can or hat, was more similar to the word cap. The semantic choice would be hat while the phonetic choice would be can. Other tasks were designed to provide a choice between semantic and phonetic interpretation of objects. For instance, in a hypothetical situation, could you call a cow a dog and if you did, would this dog bark?

The results of Ianco-Worrall's study showed that although both monolingual and bilingual children had no differences in the way they understood the words used, 54% of the younger bilingual children consistently showed a semantic preference in contrast to their monolingual peers. In monolingual children, semantic preference increased with age, suggesting that bilingual children reach a stage of semantic development 2–3 years earlier than their monolingual peers. This finding is in stark contrast to the early research and claims about bilingualism, which warned that bilingualism stunts children's linguistic development.

Language structure and awareness

In their book In Other Words, Ellen Bialystok and Kenji Hakuta, both professors studying bilingualism, examined the idea that "the knowledge of two languages is greater than the sum of its parts." They argued that there are linguistic benefits to being bilingual and that they are more than simply being able to speak two languages. A child learning two languages whose structures and rules are significantly different from each other requires the child to think in cognitively demanding ways. An example of this cognitive demand in action would be in the arbitrariness of labels for objects, or distinguishing between and using two different grammatical or syntactical structures. These areas are quite difficult for a child to learn, but with development through childhood, have been shown to increase the understanding the structure of language and introduce a greater awareness of meaning. Bilinguals develop what is referred to as metalinguistic awareness.

Reading ability

Bilingualism may also affect reading ability. Bialystok observed bilingual children's reading abilities and how the children made connections between the spoken and written language systems. Children were given a "Moving Word Task," where they needed to appropriately match a written word to an object on a card. Correctly matching the two after cards were rearranged suggested that the child could understand the written words as representations of specific words whose meanings cannot change. Comparisons to monolinguals suggested a literacy advantage, as the bilingual children achieved scores equal to monolinguals one year older. This may be attributed to greater connections between spoken and written languages. Other bilingual researchers found a relationship between phonological awareness and word recognition in native Spanish-speaking children learning to read English; phonological and word skills predicted how well a child could recognize English words. This suggested bilinguals could apply phonological awareness developed in L1 to the reading ability in L2.

Vocabulary

Past research suggests that bilinguals have a smaller vocabulary size than their monolinguals counterparts. However, a 2022 meta-analysis casts doubt on this by showing that there are no linguistic costs in bilinguals who have acquired two languages early in life; rather the linguistic cost is mainly seen in L2 learners when tested in their second language. Given that bilinguals accumulate vocabulary from both their languages, when taking both languages into account, they have a much larger vocabulary than monolinguals. However, within each language bilinguals have a smaller vocabulary size and take longer to name pictures as seen in standardized vocabulary tests, such as the Peabody Picture Vocabulary Test and Boston Naming task. A possible explanation may be that the frequency of use of words is related to increased lexical accessibility, meaning that words that are used more frequently are accessed more quickly. Past research attributing proficiency differences between bilinguals and monolinguals from smaller vocabulary and language usage time may have too small of a focus within one language, and not broaden consideration to total knowledge in both languages. In addition, the need to select the appropriate language system makes ordinary linguistic processing more effortful. The simple act of retrieving a common word is more effortful for bilinguals than monolinguals due to the competition of the two languages.

Bialystok, Luk, Peets, and Yang's study from 2010 notes that certain vocabulary tests could yield artificially low scores for bilingual children according to the domain from which the test words are taken. For example, the research team found that monolingual and bilingual 6-year-olds in their study had similar scores on English words that were associated with schooling. However, when the children were tested on English words that were associated with the home, the scores were significantly lower for the bilingual (English-Spanish) children. The researchers interpret this result as reflecting an asymmetry in vocabulary domains and language exposure: monolingual and bilingual children were equally exposed to the school context in the same language (English), but English was not commonly used in the home environments of the bilingual children. Language context may play a greater role in domain proficiency than overall language proficiency.

Effects on L1 from prolonged exposure to L2

It has been suggested that prolonged naturalistic exposure to L2 affects how L2 is processed, but it may also affect how the L1 is processed. For example, in immersion contexts, the individual experiences reduced access to L1 and extensive contact with L2, which affects and facilitates processing of L2. However, this may also consequently affect processing of their L1, such as with increased difficulty in naming objects and phonology.

To test this hypothesis, Dussias & Sagarra (2007) investigated how individuals interpreted temporarily ambiguous phrases. For example, Alguien disparó al hijo de la actriz que estaba en el balcón = Someone shot the son of the actress who was on the balcony. When asked the question, ¿Quien estaba en el balcón? = Who was on the balcony?, monolingual Spanish speakers will typically answer el hijo = the son as they have a high attachment preference, meaning they attach the modifier to the "higher" verb phrase [shot the son]. This differs from monolingual English speakers who will typically answer the actress as they have a low attachment preference, meaning they attach the modifier to the "lower" verb phrase [the actress who was on the balcony]. The researchers found that Spanish-English bilinguals in a Spanish-speaking environment showed preference for the typical Spanish high-attachment strategy. However, Spanish-English bilinguals in an English-speaking environment showed preference for the typical English low-attachment strategy, even when reading the phrase in Spanish, their dominant language. This may be because they have more exposure to English constructions, making it more available to them. But altogether, this supports the idea that the L2, English in this case, is affecting the way the native Spanish speakers use their L1.

Executive functions

From Wikipedia, the free encyclopedia

In cognitive science and neuropsychology, executive functions (collectively referred to as executive function and cognitive control) are a set of cognitive processes that are necessary for the cognitive control of behavior: selecting and successfully monitoring behaviors that facilitate the attainment of chosen goals. 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 (e.g., reasoning and problem-solving).

Executive functions gradually develop and change across the lifespan of an individual and can be improved at any time over the course of a person's life. Similarly, these cognitive processes can be adversely affected by a variety of events which affect an individual. Both neuropsychological tests (e.g., the Stroop test) and rating scales (e.g., the Behavior Rating Inventory of Executive Function) are used to measure executive functions. They are usually performed as part of a more comprehensive assessment to diagnose neurological and psychiatric disorders.

Cognitive control and stimulus control, which is associated with operant and classical conditioning, represent opposite processes (internal vs external or environmental, respectively) that compete over the control of an individual's elicited behaviors; in particular, inhibitory control is necessary for overriding stimulus-driven behavioral responses (stimulus control of behavior). The prefrontal cortex is necessary but not solely sufficient for executive functions; for example, the caudate nucleus and subthalamic nucleus also have a role in mediating inhibitory control.

Cognitive control is impaired in addiction, attention deficit hyperactivity disorder, autism, and a number of other central nervous system disorders. Stimulus-driven behavioral responses that are associated with a particular rewarding stimulus tend to dominate one's behavior in an addiction.

Neuroanatomy

Historically, the executive functions have been seen as regulated by the prefrontal regions of the frontal lobes, but it is still a matter of ongoing debate if that really is the case. Even though articles on prefrontal lobe lesions commonly refer to disturbances of executive functions and vice versa, a review found indications for the sensitivity but not for the specificity of executive function measures to frontal lobe functioning. This means that both frontal and non-frontal brain regions are necessary for intact executive functions. Probably the frontal lobes need to participate in basically all of the executive functions, but they are not the only brain structure involved.

Neuroimaging and lesion studies have identified the functions which are most often associated with the particular regions of the prefrontal cortex and associated areas.

  • The dorsolateral prefrontal cortex (DLPFC) is involved with "on-line" processing of information such as integrating different dimensions of cognition and behavior. As such, this area has been found to be associated with verbal and design fluency, ability to maintain and shift set, planning, response inhibition, working memory, organisational skills, reasoning, problem-solving, and abstract thinking.
Side view of the brain, illustrating dorsolateral prefrontal and orbitofrontal cortex
  • The anterior cingulate cortex (ACC) is involved in emotional drives, experience and integration. Associated cognitive functions include inhibition of inappropriate responses, decision making and motivated behaviors. Lesions in this area can lead to low drive states such as apathy, abulia or akinetic mutism and may also result in low drive states for such basic needs as food or drink and possibly decreased interest in social or vocational activities and sex.
  • The orbitofrontal cortex (OFC) plays a key role in impulse control, maintenance of set, monitoring ongoing behavior and socially appropriate behaviors. The orbitofrontal cortex also has roles in representing the value of rewards based on sensory stimuli and evaluating subjective emotional experiences. Lesions can cause disinhibition, impulsivity, aggressive outbursts, sexual promiscuity and antisocial behavior.

Furthermore, in their review, Alvarez and Emory state that:

The frontal lobes have multiple connections to cortical, subcortical and brain stem sites. The basis of "higher-level" cognitive functions such as inhibition, flexibility of thinking, problem solving, planning, impulse control, concept formation, abstract thinking, and creativity often arise from much simpler, "lower-level" forms of cognition and behavior. Thus, the concept of executive function must be broad enough to include anatomical structures that represent a diverse and diffuse portion of the central nervous system.

The cerebellum also appears to be involved in mediating certain executive functions, as do the ventral tegmental area and the substantia nigra.

Hypothesized role

The executive system is thought to be heavily involved in handling novel situations outside the domain of some of our 'automatic' psychological processes that could be explained by the reproduction of learned schemas or set behaviors. Psychologists Don Norman and Tim Shallice have outlined five types of situations in which routine activation of behavior would not be sufficient for optimal performance:

  1. Those that involve planning or decision-making
  2. Those that involve error correction or troubleshooting
  3. Situations where responses are not well-rehearsed or contain novel sequences of actions
  4. Dangerous or technically difficult situations
  5. Situations that require the overcoming of a strong habitual response or resisting temptation.

A prepotent response is a response for which immediate reinforcement (positive or negative) is available or has been previously associated with that response.

Executive functions are often invoked when it is necessary to override prepotent responses that might otherwise be automatically elicited by stimuli in the external environment. For example, on being presented with a potentially rewarding stimulus, such as a tasty piece of chocolate cake, a person might have the automatic response to take a bite. However, where such behavior conflicts with internal plans (such as having decided not to eat chocolate cake while on a diet), the executive functions might be engaged to inhibit that response.

Although suppression of these prepotent responses is ordinarily considered adaptive, problems for the development of the individual and the culture arise when feelings of right and wrong are overridden by cultural expectations or when creative impulses are overridden by executive inhibitions.

Historical perspective

Although research into the executive functions and their neural basis has increased markedly over recent years, the theoretical framework in which it is situated is not new. In the 1940s, the British psychologist Donald Broadbent drew a distinction between "automatic" and "controlled" processes (a distinction characterized more fully by Shiffrin and Schneider in 1977), and introduced the notion of selective attention, to which executive functions are closely allied. In 1975, the US psychologist Michael Posner used the term "cognitive control" in his book chapter entitled "Attention and cognitive control".

The work of influential researchers such as Michael Posner, Joaquin Fuster, Tim Shallice, and their colleagues in the 1980s (and later Trevor Robbins, Bob Knight, Don Stuss, and others) laid much of the groundwork for recent research into executive functions. For example, Posner proposed that there is a separate "executive" branch of the attentional system, which is responsible for focusing attention on selected aspects of the environment. The British neuropsychologist Tim Shallice similarly suggested that attention is regulated by a "supervisory system", which can override automatic responses in favour of scheduling behaviour on the basis of plans or intentions. Throughout this period, a consensus emerged that this control system is housed in the most anterior portion of the brain, the prefrontal cortex (PFC).

Psychologist Alan Baddeley had proposed a similar system as part of his model of working memory and argued that there must be a component (which he named the "central executive") that allows information to be manipulated in short-term memory (for example, when doing mental arithmetic).

Development

The executive functions are among the last mental functions to reach maturity. This is due to the delayed maturation of the prefrontal cortex, which is not completely myelinated until well into a person's third decade of life. Development of executive functions tends to occur in spurts, when new skills, strategies, and forms of awareness emerge. These spurts are thought to reflect maturational events in the frontal areas of the brain. Attentional control appears to emerge in infancy and develop rapidly in early childhood. Cognitive flexibility, goal setting, and information processing usually develop rapidly during ages 7–9 and mature by age 12. Executive control typically emerges shortly after a transition period at the beginning of adolescence. It is not yet clear whether there is a single sequence of stages in which executive functions appear, or whether different environments and early life experiences can lead people to develop them in different sequences.

Early childhood

Inhibitory control and working memory act as basic executive functions that make it possible for more complex executive functions like problem-solving to develop. Inhibitory control and working memory are among the earliest executive functions to appear, with initial signs observed in infants, 7 to 12 months old. Then in the preschool years, children display a spurt in performance on tasks of inhibition and working memory, usually between the ages of 3 and 5 years. Also during this time, cognitive flexibility, goal-directed behavior, and planning begin to develop. Nevertheless, preschool children do not have fully mature executive functions and continue to make errors related to these emerging abilities – often not due to the absence of the abilities, but rather because they lack the awareness to know when and how to use particular strategies in particular contexts.

Preadolescence

Preadolescent children continue to exhibit certain growth spurts in executive functions, suggesting that this development does not necessarily occur in a linear manner, along with the preliminary maturing of particular functions as well. During preadolescence, children display major increases in verbal working memory; goal-directed behavior (with a potential spurt around 12 years of age); response inhibition and selective attention; and strategic planning and organizational skills. Additionally, between the ages of 8 and 10, cognitive flexibility in particular begins to match adult levels. However, similar to patterns in childhood development, executive functioning in preadolescents is limited because they do not reliably apply these executive functions across multiple contexts as a result of ongoing development of inhibitory control.

Adolescence

Many executive functions may begin in childhood and preadolescence, such as inhibitory control. Yet, it is during adolescence when the different brain systems become better integrated. At this time, youth implement executive functions, such as inhibitory control, more efficiently and effectively and improve throughout this time period. Just as inhibitory control emerges in childhood and improves over time, planning and goal-directed behavior also demonstrate an extended time course with ongoing growth over adolescence. Likewise, functions such as attentional control, with a potential spurt at age 15, along with working memory, continue developing at this stage.

Adulthood

The major change that occurs in the brain in adulthood is the constant myelination of neurons in the prefrontal cortex. At age 20–29, executive functioning skills are at their peak, which allows people of this age to participate in some of the most challenging mental tasks. These skills begin to decline in later adulthood. Working memory and spatial span are areas where decline is most readily noted. Cognitive flexibility, however, has a late onset of impairment and does not usually start declining until around age 70 in normally functioning adults. Impaired executive functioning has been found to be the best predictor of functional decline in the elderly.

Models

Top-down inhibitory control

Aside from facilitatory or amplificatory mechanisms of control, many authors have argued for inhibitory mechanisms in the domain of response control, memory, selective attention, theory of mind, emotion regulation, as well as social emotions such as empathy. A recent review on this topic argues that active inhibition is a valid concept in some domains of psychology/cognitive control.

Working memory model

One influential model is Baddeley's multicomponent model of working memory, which is composed of a central executive system that regulates three subsystems: the phonological loop, which maintains verbal information; the visuospatial sketchpad, which maintains visual and spatial information; and the more recently developed episodic buffer that integrates short-term and long-term memory, holding and manipulating a limited amount of information from multiple domains in temporal and spatially sequenced episodes.

Researchers have found significant positive effects of biofeedback-enhanced relaxation on memory and inhibition in children. Biofeedback is a mind-body tool where people can learn to control and regulate their body to improve and control their executive functioning skills. To measure one's processes, researchers use their heart rate and or respiratory rates. Biofeedback-relaxation includes music therapy, art, and other mindfulness activities.

Executive functioning skills are important for many reasons, including children's academic success and social emotional development. According to the study "The Efficacy of Different Interventions to Foster Children's Executive Function Skills: A Series of Meta-Analyses", researchers found that it is possible to train executive functioning skills. Researchers conducted a meta-analytic study that looked at the combined effects of prior studies in order to find the overarching effectiveness of different interventions that promote the development of executive functioning skills in children. The interventions included computerized and non-computerized training, physical exercise, art, and mindfulness exercises. However, researchers could not conclude that art activities or physical activities could improve executive functioning skills.

Supervisory attentional system (SAS)

Another conceptual model is the supervisory attentional system (SAS). In this model, contention scheduling is the process where an individual's well-established schemas automatically respond to routine situations while executive functions are used when faced with novel situations. In these new situations, attentional control will be a crucial element to help generate new schema, implement these schema, and then assess their accuracy.

Self-regulatory model

Russell Barkley proposed a widely known model of executive functioning that is based on self-regulation. Primarily derived from work examining behavioral inhibition, it views executive functions as composed of four main abilities. One element is working memory that allows individuals to resist interfering information. A second component is the management of emotional responses in order to achieve goal-directed behaviors. Thirdly, internalization of self-directed speech is used to control and sustain rule-governed behavior and to generate plans for problem-solving. Lastly, information is analyzed and synthesized into new behavioral responses to meet one's goals. Changing one's behavioral response to meet a new goal or modify an objective is a higher level skill that requires a fusion of executive functions including self-regulation, and accessing prior knowledge and experiences.

According to this model, the executive system of the human brain provides for the cross-temporal organization of behavior towards goals and the future and coordinates actions and strategies for everyday goal-directed tasks. Essentially, this system permits humans to self-regulate their behavior so as to sustain action and problem-solving toward goals specifically and the future more generally. Thus, executive function deficits pose serious problems for a person's ability to engage in self-regulation over time to attain their goals and anticipate and prepare for the future.

Teaching children self-regulation strategies is a way to improve their inhibitory control and their cognitive flexibility. These skills allow children to manage their emotional responses. These interventions include teaching children executive function-related skills that provide the steps necessary to implement them during classroom activities and educating children on how to plan their actions before acting upon them. Executive functioning skills are how the brain plans and reacts to situations. Offering new self-regulation strategies allow children to improve their executive functioning skills by practicing something new. It is also concluded that mindfulness practices are shown to be a significantly effective intervention for children to self-regulate. This includes biofeedback-enhanced relaxation. These strategies support the growth of children's executive functioning skills.

Problem-solving model

Yet another model of executive functions is a problem-solving framework where executive functions are considered a macroconstruct composed of subfunctions working in different phases to (a) represent a problem, (b) plan for a solution by selecting and ordering strategies, (c) maintain the strategies in short-term memory in order to perform them by certain rules, and then (d) evaluate the results with error detection and error correction.

Lezak's conceptual model

One of the most widespread conceptual models on executive functions is Lezak's model. This framework proposes four broad domains of volition, planning, purposive action, and effective performance as working together to accomplish global executive functioning needs. While this model may broadly appeal to clinicians and researchers to help identify and assess certain executive functioning components, it lacks a distinct theoretical basis and relatively few attempts at validation.

Miller and Cohen's model

In 2001, Earl Miller and Jonathan Cohen published their article "An integrative theory of prefrontal cortex function", in which they argue that cognitive control is the primary function of the prefrontal cortex (PFC), and that control is implemented by increasing the gain of sensory or motor neurons that are engaged by task- or goal-relevant elements of the external environment. In a key paragraph, they argue:

We assume that the PFC serves a specific function in cognitive control: the active maintenance of patterns of activity that represent goals and the means to achieve them. They provide bias signals throughout much of the rest of the brain, affecting not only visual processes but also other sensory modalities, as well as systems responsible for response execution, memory retrieval, emotional evaluation, etc. The aggregate effect of these bias signals is to guide the flow of neural activity along pathways that establish the proper mappings between inputs, internal states, and outputs needed to perform a given task.

Miller and Cohen draw explicitly upon an earlier theory of visual attention that conceptualises perception of visual scenes in terms of competition among multiple representations – such as colors, individuals, or objects. Selective visual attention acts to 'bias' this competition in favour of certain selected features or representations. For example, imagine that you are waiting at a busy train station for a friend who is wearing a red coat. You are able to selectively narrow the focus of your attention to search for red objects, in the hope of identifying your friend. Desimone and Duncan argue that the brain achieves this by selectively increasing the gain of neurons responsive to the color red, such that output from these neurons is more likely to reach a downstream processing stage, and, as a consequence, to guide behaviour. According to Miller and Cohen, this selective attention mechanism is in fact just a special case of cognitive control – one in which the biasing occurs in the sensory domain. According to Miller and Cohen's model, the PFC can exert control over input (sensory) or output (response) neurons, as well as over assemblies involved in memory, or emotion. Cognitive control is mediated by reciprocal PFC connectivity with the sensory and motor cortices, and with the limbic system. Within their approach, thus, the term "cognitive control" is applied to any situation where a biasing signal is used to promote task-appropriate responding, and control thus becomes a crucial component of a wide range of psychological constructs such as selective attention, error monitoring, decision-making, memory inhibition, and response inhibition.

Miyake and Friedman's model

Miyake and Friedman's theory of executive functions proposes that there are three aspects of executive functions: updating, inhibition, and shifting. A cornerstone of this theoretical framework is the understanding that individual differences in executive functions reflect both unity (i.e., common EF skills) and diversity of each component (e.g., shifting-specific). In other words, aspects of updating, inhibition, and shifting are related, yet each remains a distinct entity. First, updating is defined as the continuous monitoring and quick addition or deletion of contents within one's working memory. Second, inhibition is one's capacity to supersede responses that are prepotent in a given situation. Third, shifting is one's cognitive flexibility to switch between different tasks or mental states.

Miyake and Friedman also suggest that the current body of research in executive functions suggest four general conclusions about these skills. The first conclusion is the unity and diversity aspects of executive functions. Second, recent studies suggest that much of one's EF skills are inherited genetically, as demonstrated in twin studies. Third, clean measures of executive functions can differentiate between normal and clinical or regulatory behaviors, such as ADHD. Last, longitudinal studies demonstrate that EF skills are relatively stable throughout development.

Banich's "cascade of control" model

This model from 2009 integrates theories from other models, and involves a sequential cascade of brain regions involved in maintaining attentional sets in order to arrive at a goal. In sequence, the model assumes the involvement of the posterior dorsolateral prefrontal cortex (DLPFC), the mid-DLPFC, and the posterior and anterior dorsal anterior cingulate cortex (ACC).

The cognitive task used in the article is selecting a response in the Stroop task, among conflicting color and word responses, specifically a stimulus where the word "green" is printed in red ink. The posterior DLPFC creates an appropriate attentional set, or rules for the brain to accomplish the current goal. For the Stroop task, this involves activating the areas of the brain involved in color perception, and not those involved in word comprehension. It counteracts biases and irrelevant information, like the fact that the semantic perception of the word is more salient to most people than the color in which it is printed.

Next, the mid-DLPFC selects the representation that will fulfill the goal. The task-relevant information must be separated from other sources of information in the task. In the example, this means focusing on the ink color and not the word.

The posterior dorsal ACC is next in the cascade, and it is responsible for response selection. This is where the decision is made whether the Stroop task participant will say "green" (the written word and the incorrect answer) or "red" (the font color and correct answer).

Following the response, the anterior dorsal ACC is involved in response evaluation, deciding whether one's response were correct or incorrect. Activity in this region increases when the probability of an error is higher.

The activity of any of the areas involved in this model depends on the efficiency of the areas that came before it. If the DLPFC imposes a lot of control on the response, the ACC will require less activity.

Recent work using individual differences in cognitive style has shown exciting support for this model. Researchers had participants complete an auditory version of the Stroop task, in which either the location or semantic meaning of a directional word had to be attended to. Participants that either had a strong bias toward spatial or semantic information (different cognitive styles) were then recruited to participate in the task. As predicted, participants that had a strong bias toward spatial information had more difficulty paying attention to the semantic information and elicited increased electrophysiological activity from the ACC. A similar activity pattern was also found for participants that had a strong bias toward verbal information when they tried to attend to spatial information.

Assessment

Assessment of executive functions involves gathering data from several sources and synthesizing the information to look for trends and patterns across time and settings. Apart from standardized neuropsychological tests, other measures can and should be used, such as behaviour checklists, observations, interviews, and work samples. From these, conclusions may be drawn on the use of executive functions.

There are several different kinds of instruments (e.g., performance based, self-report) that measure executive functions across development. These assessments can serve a diagnostic purpose for a number of clinical populations.

Experimental evidence

The executive system has been traditionally quite hard to define, mainly due to what psychologist Paul W. Burgess calls a lack of "process-behaviour correspondence". That is, there is no single behavior that can in itself be tied to executive function, or indeed executive dysfunction. For example, it is quite obvious what reading-impaired patients cannot do, but it is not so obvious what exactly executive-impaired patients might be incapable of.

This is largely due to the nature of the executive system itself. It is mainly concerned with the dynamic, "online" co-ordination of cognitive resources, and, hence, its effect can be observed only by measuring other cognitive processes. In similar manner, it does not always fully engage outside of real-world situations. As neurologist Antonio Damasio has reported, a patient with severe day-to-day executive problems may still pass paper-and-pencil or lab-based tests of executive function.

Theories of the executive system were largely driven by observations of patients with frontal lobe damage. They exhibited disorganized actions and strategies for everyday tasks (a group of behaviors now known as dysexecutive syndrome) although they seemed to perform normally when clinical or lab-based tests were used to assess more fundamental cognitive functions such as memory, learning, language, and reasoning. It was hypothesized that, to explain this unusual behaviour, there must be an overarching system that co-ordinates other cognitive resources.

Much of the experimental evidence for the neural structures involved in executive functions comes from laboratory tasks such as the Stroop task or the Wisconsin Card Sorting Task (WCST). In the Stroop task, for example, human subjects are asked to name the color that color words are printed in when the ink color and word meaning often conflict (for example, the word "RED" in green ink). Executive functions are needed to perform this task, as the relatively overlearned and automatic behaviour (word reading) has to be inhibited in favour of a less practiced task – naming the ink color. Recent functional neuroimaging studies have shown that two parts of the PFC, the anterior cingulate cortex (ACC) and the dorsolateral prefrontal cortex (DLPFC), are thought to be particularly important for performing this task.

Context-sensitivity of PFC neurons

Other evidence for the involvement of the PFC in executive functions comes from single-cell electrophysiology studies in non-human primates, such as the macaque monkey, which have shown that (in contrast to cells in the posterior brain) many PFC neurons are sensitive to a conjunction of a stimulus and a context. For example, PFC cells might respond to a green cue in a condition where that cue signals that a leftwards fast movement of the eyes and the head should be made, but not to a green cue in another experimental context. This is important, because the optimal deployment of executive functions is invariably context-dependent.

One example from Miller & Cohen involves a pedestrian crossing the street. In the United States, where cars drive on the right side of the road, an American learns to look left when crossing the street. However, if that American visits a country where cars drive on the left, such as the United Kingdom, then the opposite behavior would be required (looking to the right). In this case, the automatic response needs to be suppressed (or augmented) and executive functions must make the American look to the right while in the UK.

Neurologically, this behavioural repertoire clearly requires a neural system that is able to integrate the stimulus (the road) with a context (US or UK) to cue a behaviour (look left or look right). Current evidence suggests that neurons in the PFC appear to represent precisely this sort of information. Other evidence from single-cell electrophysiology in monkeys implicates ventrolateral PFC (inferior prefrontal convexity) in the control of motor responses. For example, cells that increase their firing rate to NoGo signals as well as a signal that says "don't look there!" have been identified.

Attentional biasing in sensory regions

Electrophysiology and functional neuroimaging studies involving human subjects have been used to describe the neural mechanisms underlying attentional biasing. Most studies have looked for activation at the 'sites' of biasing, such as in the visual or auditory cortices. Early studies employed event-related potentials to reveal that electrical brain responses recorded over left and right visual cortex are enhanced when the subject is instructed to attend to the appropriate (contralateral) side of space.

The advent of bloodflow-based neuroimaging techniques such as functional magnetic resonance imaging (fMRI) and positron emission tomography (PET) has more recently permitted the demonstration that neural activity in a number of sensory regions, including color-, motion-, and face-responsive regions of visual cortex, is enhanced when subjects are directed to attend to that dimension of a stimulus, suggestive of gain control in sensory neocortex. For example, in a typical study, Liu and coworkers presented subjects with arrays of dots moving to the left or right, presented in either red or green. Preceding each stimulus, an instruction cue indicated whether subjects should respond on the basis of the colour or the direction of the dots. Even though colour and motion were present in all stimulus arrays, fMRI activity in colour-sensitive regions (V4) was enhanced when subjects were instructed to attend to the colour, and activity in motion-sensitive regions was increased when subjects were cued to attend to the direction of motion. Several studies have also reported evidence for the biasing signal prior to stimulus onset, with the observation that regions of the frontal cortex tend to come active prior to the onset of an expected stimulus.

Connectivity between the PFC and sensory regions

Despite the growing currency of the 'biasing' model of executive functions, direct evidence for functional connectivity between the PFC and sensory regions when executive functions are used, is to date rather sparse. Indeed, the only direct evidence comes from studies in which a portion of frontal cortex is damaged, and a corresponding effect is observed far from the lesion site, in the responses of sensory neurons. However, few studies have explored whether this effect is specific to situations where executive functions are required. Other methods for measuring connectivity between distant brain regions, such as correlation in the fMRI response, have yielded indirect evidence that the frontal cortex and sensory regions communicate during a variety of processes thought to engage executive functions, such as working memory, but more research is required to establish how information flows between the PFC and the rest of the brain when executive functions are used. As an early step in this direction, an fMRI study on the flow of information processing during visuospatial reasoning has provided evidence for causal associations (inferred from the temporal order of activity) between sensory-related activity in occipital and parietal cortices and activity in posterior and anterior PFC. Such approaches can further elucidate the distribution of processing between executive functions in PFC and the rest of the brain.

Bilingualism and executive functions

A growing body of research demonstrates that bilinguals might show advantages in executive functions, specifically inhibitory control and task switching. A possible explanation for this is that speaking two languages requires controlling one's attention and choosing the correct language to speak. Across development, bilingual infants, children, and elderly show a bilingual advantage when it comes to executive functioning. The advantage does not seem to manifest in younger adults. Bimodal bilinguals, or people who speak one oral language and one sign language, do not demonstrate this bilingual advantage in executive functioning tasks. This may be because one is not required to actively inhibit one language in order to speak the other. Bilingual individuals also seem to have an advantage in an area known as conflict processing, which occurs when there are multiple representations of one particular response (for example, a word in one language and its translation in the individual's other language). Specifically, the lateral prefrontal cortex has been shown to be involved with conflict processing. However, there are still some doubts. In a meta-analytic review, researchers concluded that bilingualism did not enhance executive functioning in adults.

In disease or disorder

The study of executive function in Parkinson's disease suggests subcortical areas such as the amygdala, hippocampus and basal ganglia are important in these processes. Dopamine modulation of the prefrontal cortex is responsible for the efficacy of dopaminergic drugs on executive function, and gives rise to the Yerkes–Dodson Curve. The inverted U represents decreased executive functioning with excessive arousal (or increased catecholamine release during stress), and decreased executive functioning with insufficient arousal. The low activity polymorphism of catechol-O-methyltransferase is associated with slight increase in performance on executive function tasks in healthy persons. Executive functions are impaired in multiple disorders including anxiety disorder, major depressive disorder, bipolar disorder, attention deficit hyperactivity disorder, schizophrenia and autism. Lesions to the prefrontal cortex, such as in the case of Phineas Gage, may also result in deficits of executive function. Damage to these areas may also manifest in deficits of other areas of function, such as motivation, and social functioning.

Future directions

Other important evidence for executive functions processes in the prefrontal cortex have been described. One widely cited review article emphasizes the role of the medial part of the PFC in situations where executive functions are likely to be engaged – for example, where it is important to detect errors, identify situations where stimulus conflict may arise, make decisions under uncertainty, or when a reduced probability of obtaining favourable performance outcomes is detected. This review, like many others, highlights interactions between medial and lateral PFC, whereby posterior medial frontal cortex signals the need for increased executive functions and sends this signal on to areas in dorsolateral prefrontal cortex that actually implement control. Yet there has been no compelling evidence at all that this view is correct, and, indeed, one article showed that patients with lateral PFC damage had reduced ERNs (a putative sign of dorsomedial monitoring/error-feedback) – suggesting, if anything, that the direction of flow of the control could be in the reverse direction. Another prominent theory emphasises that interactions along the perpendicular axis of the frontal cortex, arguing that a 'cascade' of interactions between anterior PFC, dorsolateral PFC, and premotor cortex guides behaviour in accordance with past context, present context, and current sensorimotor associations, respectively.

Advances in neuroimaging techniques have allowed studies of genetic links to executive functions, with the goal of using the imaging techniques as potential endophenotypes for discovering the genetic causes of executive function.

More research is required to develop interventions that can improve executive functions and help people generalize those skills to daily activities and settings

Human extinction

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