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Tuesday, October 22, 2019

Cerebral hemisphere

From Wikipedia, the free encyclopedia

Cerebral hemisphere
Blausen 0215 CerebralHemispheres.png
Human brain seen from front.
Cerebral hemisphere - animation.gif
  Right cerebral hemisphere
  Left cerebral hemisphere
Details
Identifiers
LatinHemisphaerium cerebri
NeuroNames241
NeuroLex IDbirnlex_1796
TAA14.1.09.002
FMA61817

The vertebrate cerebrum (brain) is formed by two cerebral hemispheres that are separated by a groove, the longitudinal fissure. The brain can thus be described as being divided into left and right cerebral hemispheres. Each of these hemispheres has an outer layer of grey matter, the cerebral cortex, that is supported by an inner layer of white matter. In eutherian (placental) mammals, the hemispheres are linked by the corpus callosum, a very large bundle of nerve fibers. Smaller commissures, including the anterior commissure, the posterior commissure and the fornix, also join the hemispheres and these are also present in other vertebrates. These commissures transfer information between the two hemispheres to coordinate localized functions.

There are three known poles of the cerebral hemispheres: the occipital pole, the frontal pole, and the temporal pole.

The central sulcus is a prominent fissure which separates the parietal lobe from the frontal lobe and the primary motor cortex from the primary somatosensory cortex.

Macroscopically the hemispheres are roughly mirror images of each other, with only subtle differences, such as the Yakovlevian torque seen in the human brain, which is a slight warping of the right side, bringing it just forward of the left side. On a microscopic level, the cytoarchitecture of the cerebral cortex, shows the functions of cells, quantities of neurotransmitter levels and receptor subtypes to be markedly asymmetrical between the hemispheres. However, while some of these hemispheric distribution differences are consistent across human beings, or even across some species, many observable distribution differences vary from individual to individual within a given species.

Structure

Each cerebral hemisphere has an outer layer of cerebral cortex which is of grey matter and in the interior of the cerebral hemispheres is an inner layer or core of white matter known as the centrum semiovale. The interior portion of the hemispheres of the cerebrum includes the lateral ventricles, the basal ganglia, and the white matter.

Poles

Poles of cerebral hemispheres
 
There are three poles of the cerebrum, the occipital pole, the frontal pole, and the temporal pole. The occipital pole is the posterior end of each occipital lobe in each hemisphere. It is more pointed than the rounder frontal pole. The frontal pole is at the frontmost part of the frontal lobe in each hemisphere, and is more rounded than the occipital pole. The temporal pole is located between the frontal and occipital poles, and sits in the anterior part of middle cranial fossa in each temporal lobe.

Composition

If the upper part of either hemisphere is removed, at a level about 1.25 cm above the corpus callosum, the central white matter will be exposed as an oval-shaped area, the centrum semiovale, surrounded by a narrow convoluted margin of gray substance, and studded with numerous minute red dots (puncta vasculosa), produced by the escape of blood from divided blood vessels.

If the remaining portions of the hemispheres be slightly drawn apart a broad band of white substance, the corpus callosum, will be observed, connecting them at the bottom of the longitudinal fissure; the margins of the hemispheres which overlap the corpus callosum are called the labia cerebri.

Each labium is part of the cingulate gyrus already described; and the groove between it and the upper surface of the corpus callosum is termed the callosal sulcus.

If the hemispheres are sliced off to a level with the upper surface of the corpus callosum, the white substance of that structure will be seen connecting the two hemispheres.

The large expanse of medullary matter now exposed, surrounded by the convoluted margin of gray substance, is called the centrum semiovale. The blood supply to the centrum semiovale is from the superficial middle cerebral artery. The cortical branches of this artery descend to provide blood to the centrum semiovale.

Development

The cerebral hemispheres are derived from the telencephalon. They arise five weeks after conception as bilateral invaginations of the walls. The hemispheres grow round in a C-shape and then back again, pulling all structures internal to the hemispheres (such as the ventricles) with them. The intraventricular foramina (also called the foramina of Monro) allows communication with the lateral ventricles. The choroid plexus is formed from ependymal cells and vascular mesenchyme.

Function

Hemisphere lateralization

Broad generalizations are often made in popular psychology about certain functions (e.g. logic, creativity) being lateralized, that is, located in the right or left side of the brain. These claims are often inaccurate, as most brain functions are actually distributed across both hemispheres. Most scientific evidence for asymmetry relates to low-level perceptual functions rather than the higher-level functions popularly discussed (e.g. subconscious processing of grammar, not "logical thinking" in general). In addition to this lateralization of some functions, the low-level representations also tend to represent the contralateral side of the body.

The best example of an established lateralization is that of Broca's and Wernicke's Areas (language) where both are often found exclusively on the left hemisphere. These areas frequently correspond to handedness however, meaning the localization of these areas is regularly found on the hemisphere opposite to the dominant hand. Function lateralization such as semantics, prosodic, intonation, accentuation, prosody, etc. has since been called into question and largely been found to have a neuronal basis in both hemispheres.

Cerebral hemispheres of a human embryo at 8 weeks.
 
Perceptual information is processed in both hemispheres, but is laterally partitioned: information from each side of the body is sent to the opposite hemisphere (visual information is partitioned somewhat differently, but still lateralized). Similarly, motor control signals sent out to the body also come from the hemisphere on the opposite side. Thus, hand preference (which hand someone prefers to use) is also related to hemisphere lateralization.

In some aspects, the hemispheres are asymmetrical; the right side is slightly bigger. There are higher levels of the neurotransmitter norepinephrine on the right and higher levels of dopamine on the left. There is more white matter (longer axons) on the right and more grey matter (cell bodies) on the left.

Linear reasoning functions of language such as grammar and word production are often lateralized to the left hemisphere of the brain. In contrast, holistic reasoning functions of language such as intonation and emphasis are often lateralized to the right hemisphere of the brain. Other integrative functions such as intuitive or heuristic arithmetic, binaural sound localization, etc. seem to be more bilaterally controlled.

Clinical significance

Infarcts of the centrum ovale can occur.

As a treatment for epilepsy the corpus callosum may be severed to cut the major connection between the hemispheres in a procedure known as a corpus callosotomy

A hemispherectomy is the removal or disabling of one of the hemispheres of the brain. This is a rare procedure used in some extreme cases of seizures which are unresponsive to other treatments.

Broca's area

From Wikipedia, the free encyclopedia
 
Broca's area
Broca’s area - BA44 and BA45.png
Broca's area is made up of Brodmann areas 44 (pars opercularis) and 45 (pars triangularis)
Broca's area - lateral view.png
Broca's area (shown in red)
Details
Part ofFrontal lobe
ArteryMiddle cerebral
VeinSuperior sagittal sinus
Identifiers
MeSHD065711
NeuroNames2062
FMA242176

Broca's area, or the Broca area , is a region in the frontal lobe of the dominant hemisphere, usually the left, of the brain with functions linked to speech production.

Language processing has been linked to Broca's area since Pierre Paul Broca reported impairments in two patients. They had lost the ability to speak after injury to the posterior inferior frontal gyrus (pars triangularis) (BA45) of the brain. Since then, the approximate region he identified has become known as Broca's area, and the deficit in language production as Broca's aphasia, also called expressive aphasia. Broca's area is now typically defined in terms of the pars opercularis and pars triangularis of the inferior frontal gyrus, represented in Brodmann's cytoarchitectonic map as Brodmann area 44 and Brodmann area 45 of the dominant hemisphere.

Functional magnetic resonance imaging has shown language processing to also involve the third part of the inferior frontal gyrus the pars orbitalis, as well as the ventral part of BA6 and these are now often included in a larger area called Broca's region.

Studies of chronic aphasia have implicated an essential role of Broca's area in various speech and language functions. Further, fMRI studies have also identified activation patterns in Broca's area associated with various language tasks. However, slow destruction of the Broca's area by brain tumors can leave speech relatively intact, suggesting its functions can shift to nearby areas in the brain.

Structure

Brodmann area 44

Brodmann area 45

Broca's area is often identified by visual inspection of the topography of the brain either by macrostructural landmarks such as sulci or by the specification of coordinates in a particular reference space. The currently used Talairach and Tournoux atlas projects Brodmann's cytoarchitectonic map onto a template brain. Because Brodmann's parcelation was based on subjective visual inspection of cytoarchitectonic borders and also Brodmann analyzed only one hemisphere of one brain, the result is imprecise. Further, because of considerable variability across brains in terms of shape, size, and position relative to sulcal and gyral structure, a resulting localization precision is limited.

Nevertheless, Broca's area in the left hemisphere and its homologue in the right hemisphere are designations usually used to refer to the triangular part of inferior frontal gyrus (PTr) and the opercular part of inferior frontal gyrus (POp). The PTr and POp are defined by structural landmarks that only probabilistically divide the inferior frontal gyrus into anterior and posterior cytoarchitectonic areas of 45 and 44, respectively, by Brodmann's classification scheme.

Area 45 receives more afferent connections from the prefrontal cortex, the superior temporal gyrus, and the superior temporal sulcus, compared to area 44, which tends to receive more afferent connections from motor, somatosensory, and inferior parietal regions.

The differences between area 45 and 44 in cytoarchitecture and in connectivity suggest that these areas might perform different functions. Indeed, recent neuroimaging studies have shown that the PTr and Pop, corresponding to areas 45 and 44, respectively, play different functional roles in the human with respect to language comprehension and action recognition/understanding.

Functions

Language comprehension

For a long time, it was assumed that the role of Broca's area was more devoted to language production than language comprehension. However, there is evidence to demonstrate that Broca's area also plays a significant role in language comprehension. Patients with lesions in Broca's area who exhibit agrammatical speech production also show inability to use syntactic information to determine the meaning of sentences. Also, a number of neuroimaging studies have implicated an involvement of Broca's area, particularly of the pars opercularis of the left inferior frontal gyrus, during the processing of complex sentences. Further, it has recently been found in functional magnetic resonance imaging (fMRI) experiments involving highly ambiguous sentences result in a more activated inferior frontal gyrus. Therefore, the activity level in the inferior frontal gyrus and the level of lexical ambiguity are directly proportional to each other, because of the increased retrieval demands associated with highly ambiguous content.

There is also specialisation for particular aspects of comprehension within Broca's area. Work by Devlin et al. (2003) showed in a repetitive transcranial magnetic stimulation (rTMS) study that there was an increase in reaction times when performing a semantic task under rTMS aimed at the pars triangularis (situated in the anterior part of Broca's area). The increase in reaction times is indicative that that particular area is responsible for processing that cognitive function. Disrupting these areas via TMS disrupts computations performed in the areas leading to an increase in time needed to perform the computations (reflected in reaction times). Later work by Nixon et al. (2004) showed that when the pars opercularis (situated in the posterior part of Broca's area) was stimulated under rTMS there was an increase in reaction times in a phonological task. Gough et al. (2005) performed an experiment combining elements of these previous works in which both phonological and semantic tasks were performed with rTMS stimulation directed at either the anterior or the posterior part of Broca's area. The results from this experiment conclusively distinguished anatomical specialisation within Broca's area for different components of language comprehension. Here the results showed that under rTMS stimulation:
  • Semantic tasks only showed a decrease in reaction times when stimulation was aimed at the anterior part of Broca's area (where a decrease of 10% (50ms) was seen compared to a no-TMS control group)
  • Phonological tasks showed a decrease in reaction times when stimulation was aimed at the posterior part of Broca's area (where a decrease of 6% (30ms) was seen compared to control)
To summarise, the work above shows anatomical specialisation in Broca's area for language comprehension, with the anterior part of Broca's area responsible for understanding the meaning of words (semantics) and the posterior part of Broca's area responsible for understanding how words sound (phonology).

Action recognition and production

Recent experiments have indicated that Broca's area is involved in various cognitive and perceptual tasks. One important contribution of Brodmann's area 44 is also found in the motor-related processes. Observation of meaningful hand shadows resembling moving animals activates frontal language area, demonstrating that Broca's area indeed plays a role in interpreting action of others. An activation of BA 44 was also reported during execution of grasping and manipulation.

Speech-associated gestures

It has been speculated that because speech-associated gestures could possibly reduce lexical or sentential ambiguity, comprehension should improve in the presence of speech-associated gestures. As a result of improved comprehension, the involvement of Broca's area should be reduced.

Many neuroimaging studies have also shown activation of Broca's area when representing meaningful arm gestures. A recent study has shown evidence that word and gesture are related at the level of translation of particular gesture aspects such as its motor goal and intention. This finding helps explain why, when this area is defective, those who use sign language also suffer from language deficits. This finding that aspects of gestures are translated in words within Broca's area also explains language development in terms of evolution. Indeed, many authors have proposed that speech evolved from a primitive communication that arose from gestures.

Speaking without Broca's area

Damage to Broca's area is commonly associated with telegraphic speech made up of content vocabulary. For example, a person with Broca's aphasia may say something like, "Drive, store. Mom." meaning to say, "My mom drove me to the store today." Therefore, the content of the information is correct, but the grammar and fluidity of the sentence is missing.

The essential role of the Broca's area in speech production has been questioned since it can be destroyed while leaving language nearly intact. In one case of a computer engineer, a slow-growing glioma tumor was removed. The tumor and the surgery destroyed the left inferior and middle frontal gyrus, the head of the caudate nucleus, the anterior limb of the internal capsule, and the anterior insula. However, there were minimal language problems three months after removal and the individual returned to his professional work. These minor problems include the inability to create syntactically complex sentences including more than two subjects, multiple causal conjunctions, or reported speech. These were explained by researchers as due to working memory problems. They also attributed his lack of problems to extensive compensatory mechanisms enabled by neural plasticity in the nearby cerebral cortex and a shift of some functions to the homologous area in the right hemisphere.

Clinical significance

Stuttering

A speech disorder known as stuttering is seen to be associated with underactivity in Broca's area.

Aphasia

Aphasia is an acquired language disorder affecting all modalities such as writing, reading, speaking, and listening and results from brain damage. It is often a chronic condition that creates changes in all areas of one's life.

Expressive aphasia vs. other aphasias

Patients with expressive aphasia, also known as Broca's aphasia, are individuals who know "what they want to say, they just cannot get it out". They are typically able to comprehend words, and sentences with a simple syntactic structure (see above), but are more or less unable to generate fluent speech. Other symptoms that may be present include problems with fluency, articulation, word-finding, word repetition, and producing and comprehending complex grammatical sentences, both orally and in writing.

This specific group of symptoms distinguishes those who have expressive aphasia from individuals with other types of aphasia. There are several distinct "types" of aphasia, and each type is characterized by a different set of language deficits. Although those who have expressive aphasia tend to retain good spoken language comprehension, other types of aphasia can render patients completely unable to understand any language at all, unable to understand any spoken language (auditory verbal agnosia), whereas still other types preserve language comprehension, but with deficits. People with expressive aphasia may struggle less with reading and writing than those with other types of aphasia. Although individuals with expressive aphasia tend to have a good ability to self-monitor their language output (they "hear what they say" and make corrections), other types of aphasics can seem entirely unaware of their language deficits. 

In the classical sense, expressive aphasia is the result of injury to Broca's area; it is often the case that lesions in specific brain areas cause specific, dissociable symptoms, although case studies show there is not always a one-to-one mapping between lesion location and aphasic symptoms. The correlation between damage to certain specific brain areas (usually in the left hemisphere) and the development of specific types of aphasia makes it possible to deduce (albeit very roughly) the location of a suspected brain lesion based only on the presence (and severity) of a certain type of aphasia, though this is complicated by the possibility that a patient may have damage to a number of brain areas and may exhibit symptoms of more than one type of aphasia. The examination of lesion data in order to deduce which brain areas are essential in the normal functioning of certain aspects of cognition is called the deficit-lesion method; this method is especially important in the branch of neuroscience known as aphasiology. Cognitive science - to be specific, cognitive neuropsychology - are branches of neuroscience that also make extensive use of the deficit-lesion method.

Major characteristics of different types of acute aphasia
Type of aphasia Speech repetition Naming Auditory comprehension Fluency
Expressive aphasia Moderate–severe Moderate–severe Mild difficulty Non-fluent, effortful, slow
Receptive aphasia Mild–severe Mild–severe Defective Fluent paraphasic
Conduction aphasia Poor Poor Relatively good Fluent
Mixed transcortical aphasia Moderate Poor Poor Non-fluent
Transcortical motor aphasia Good Mild–severe Mild Non-fluent
Transcortical sensory aphasia Good Moderate–severe Poor Fluent
Global aphasia Poor Poor Poor Non-fluent
Anomic aphasia Mild Moderate–severe Mild Fluent

Newer implications related to lesions in Broca's area

Since studies carried out in the late 1970's it has been understood that the relationship between Broca's area and Broca's aphasia is not as consistent as once thought. Lesions to Broca's area alone don't result in a Broca's aphasia, nor do Broca's aphasic patients necessarily have lesions in Broca's area. Lesions to Broca's area alone are known to produce just a transient mutism that resolves inside 3–6 weeks. This discovery suggests that Broca's area may be included in some aspect of verbalization or articulation; however, it does not address its part in sentence comprehension. Still, Broca's area frequently emerges in functional imaging studies of sentence processing. However, it also becomes activated in word-level tasks. This suggests that Broca’s area is not dedicated to sentence processing but supports a function common to both. In fact, Broca's area can show activation in such non-linguistic tasks as imagery of motion.

Considering the hypothesis that Broca's area may be most involved in articulation, its activation in all of these tasks may be due to subjects' covert articulation while formulating a response. Despite this caveat, a consensus seems to be forming that whatever role Broca's area may play, it may relate to known working memory functions of the frontal areas. (There is a wide distribution of Talairach coordinates reported in the functional imaging literature that are referred to as part of Broca's area.) The processing of a passive voice sentence, for example, may require working memory to assist in the temporary retention of information while other relevant parts of the sentence are being manipulated (i.e. to resolve the assignment of thematic roles to arguments). Miyake, Carpenter, and Just have proposed that sentence processing relies on such general verbal working memory mechanisms while Caplan and Waters consider Broca’s area to be involved in working memory specifically for syntactic processing. Friederici (2002) breaks Broca's area into its component regions and suggests that Brodmann's area 44 is involved in working memory for both phonological and syntactic structure. This area becomes active first for phonology and later for syntax as the time course for the comprehension process unfolds. Brodmann's area 45 together with Brodmann's area 47 is viewed as being specifically involved in working memory for semantic features and thematic structure where processes of syntactic reanalysis and repair are required. These areas come online after Brodmann's area 44 has finished its processing role and where comprehension of complex sentences must rely on general memory resources. All of these theories indicate a move towards a view that syntactic comprehension problems arise from a computational rather than a conceptual deficit. Newer theories are taking a more dynamic view of how the brain integrates different linguistic and cognitive components and are examining the time course of these operations.

Neurocognitive studies have already implicated frontal areas adjacent to Broca's area as important for working memory in non-linguistic as well as linguistic tasks. Cabeza and Nyberg's analysis of imaging studies of working memory supports the view that BA45/47 is recruited for selecting or comparing information, while BA9/46 might be more involved in the manipulation of information in working memory. Since large lesions are typically required to produce a Broca's aphasia, it is likely that these regions may also become compromised in some patients and may contribute to their comprehension deficits for complex morphosyntactic structures.

Broca's area: A key center in the linking phonemic sequences

Broca's area has been previously associated with a variety of processes, including phonological segmentation, syntactic processing, and unification, all of which involve segmenting and linking different types of linguistic information. Although repeating and reading single words do not engage semantic and syntactic processing, they do require an operation linking phonemic sequences with motor gestures. Findings indicate that this linkage is coordinated by Broca's area through reciprocal interactions with temporal and frontal cortices responsible for phonemic and articulatory representations, respectively, including interactions with motor cortex before the actual act of speech. Based on these unique findings, it has been proposed that Broca's area is not the seat of articulation per se, but rather is a key node in manipulating and forwarding neural information across large-scale cortical networks responsible for key components of speech production.

History

In a study published in 2007, the preserved brains of both Leborgne and Lelong (patients of Broca) were reinspected using high-resolution volumetric MRI. The purpose of this study was to scan the brains in three dimensions and to identify the extent of both cortical and subcortical lesions in more detail. The study also sought to locate the exact site of the lesion in the frontal lobe in relation to what is now called Broca's area with the extent of subcortical involvement.

Broca's patients

Louis Victor Leborgne (Tan)

Leborgne was a patient of Broca's. At 30 years old, he was almost completely unable to produce any words or phrases. He was able to repetitively produce only the word tan. After his death, a neurosyphilitic lesion was discovered on the surface of his left frontal lobe.

Lelong

Lelong was another patient of Broca's. He also exhibited reduced productive speech. He could only say five words, 'yes', 'no', 'three', 'always', and 'lelo' (a mispronunciation of his own name). A lesion within the lateral frontal lobe was discovered during Lelong's autopsy. Broca's previous patient, Leborgne, had this lesion in the same area of his frontal lobe. These two cases led Broca to believe that speech was localized to this particular area.

MRI findings

Examination of the brains of Broca's two historic patients with high-resolution MRI has produced several interesting findings. First, the MRI findings suggest that other areas besides Broca's area may also have contributed to the patients' reduced productive speech. This finding is significant because it has been found that, though lesions to Broca's area alone can possibly cause temporary speech disruption, they do not result in severe speech arrest. Therefore, there is a possibility that the aphasia denoted by Broca as an absence of productive speech also could have been influenced by the lesions in the other region. Another finding is that the region, which was once considered to be critical for speech by Broca, is not precisely the same region as what is now known as Broca's area. This study provides further evidence to support the claim that language and cognition are far more complicated than once thought and involve various networks of brain regions.

Evolution of language

The pursuit of a satisfying theory that addresses the origin of language in humans has led to the consideration of a number of evolutionary "models". These models attempt to show how modern language might have evolved, and a common feature of many of these theories is the idea that vocal communication was initially used to complement a far more dominant mode of communication through gesture. Human language might have evolved as the "evolutionary refinement of an implicit communication system already present in lower primates, based on a set of hand/mouth goal-directed action representations."

"Hand/mouth goal-directed action representations" is another way of saying "gestural communication", "gestural language", or "communication through body language". The recent finding that Broca's area is active when people are observing others engaged in meaningful action is evidence in support of this idea. It was hypothesized that a precursor to the modern Broca's area was involved in translating gestures into abstract ideas by interpreting the movements of others as meaningful action with an intelligent purpose. It is argued that over time the ability to predict the intended outcome and purpose of a set of movements eventually gave this area the capability to deal with truly abstract ideas, and therefore (eventually) became capable of associating sounds (words) with abstract meanings. The observation that frontal language areas are activated when people observe Hand Shadows is further evidence that human language may have evolved from existing neural substrates that evolved for the purpose of gesture recognition. The study, therefore, claims that Broca's area is the "motor center for speech", which assembles and decodes speech sounds in the same way it interprets body language and gestures. Consistent with this idea is that the neural substrate that regulated motor control in the common ancestor of apes and humans was most likely modified to enhance cognitive and linguistic ability. Studies of speakers of American Sign Language and English suggest that the human brain recruited systems that had evolved to perform more basic functions much earlier; these various brain circuits, according to the authors, were tapped to work together in creating language.

Another recent finding has showed significant areas of activation in subcortical and neocortical areas during the production of communicative manual gestures and vocal signals in chimpanzees. Further, the data indicating that chimpanzees intentionally produce manual gestures as well as vocal signals to communicate with humans suggests that the precursors to human language are present at both the behavioral and neuronanatomical levels. More recently, the neocortical distribution of activity-dependent gene expression in marmosets provided direct evidence that the ventrolateral prefrontal cortex, which comprises Broca's area in humans and has been associated with auditory processing of species-specific vocalizations and orofacial control in macaques, is engaged during vocal output in a New World monkey. These findings putatively set the origin of vocalization-related neocortical circuits to at least 35 million years ago, when the Old and New World monkey lineages split.

Wernicke's area

From Wikipedia, the free encyclopedia
 
Wernicke's area
BrocasAreaSmall.png
Wernicke's area is located in the temporal lobe, shown here in grey
Details
LocationTemporal lobe of the dominant cerebral hemisphere.
ArteryBranches from the middle cerebral artery
Identifiers
MeSHD065813
NeuroNames1233
NeuroLex IDnlx_144087
FMA242178

Wernicke's area (/ˈvɛərnɪkə/ or /ˈvɛərnɪki/; German: [ˈvɛʁnɪkə]), also called Wernicke's speech area, is one of the two parts of the cerebral cortex that are linked to speech (the other is Broca's area). It is involved in the comprehension of written and spoken language (in contrast to Broca's area that is involved in the production of language). It is traditionally thought to be in Brodmann area 39,40, which is located in the superior temporal lobe in the dominant cerebral hemisphere (which is the left hemisphere in about 95% of right handed individuals and 60% of left handed individuals).

Damage caused to Wernicke's area results in receptive, fluent aphasia. This means that the person with aphasia will be able to fluently connect words, but the phrases will lack meaning. This is unlike non-fluent aphasia, in which the person will use meaningful words, but in a non-fluent, telegraphic manner.

Structure

Wernicke's area is classically located in the posterior section of the superior temporal gyrus (STG) in the (most commonly) left cerebral hemisphere. This area encircles the auditory cortex on the lateral sulcus (the part of the brain where the temporal lobe and parietal lobe meet). This area is neuroanatomically described as the posterior part of Brodmann area 22.

However, there is an absence of consistent definitions as to the location. Some identify it with the unimodal auditory association in the superior temporal gyrus anterior to the primary auditory cortex (the anterior part of BA 22). This is the site most consistently implicated in auditory word recognition by functional brain imaging experiments. Others include also adjacent parts of the heteromodal cortex in BA 39 and BA40 in the parietal lobe.

While previously thought to connect Wernicke's area and Broca's area, new research demonstrates that the arcuate fasciculus instead connects to posterior receptive areas with premotor/motor areas, and not to Broca's area. Consistent with the word recognition site identified in brain imaging, the uncinate fasciculus connects anterior superior temporal regions with Broca's area.

Function

Right homologous area

Research using Transcranial magnetic stimulation suggests that the area corresponding to the Wernicke’s area in the non-dominant cerebral hemisphere has a role in processing and resolution of subordinate meanings of ambiguous words—such as ‘‘river’’ when given the ambiguous word "bank." In contrast, the Wernicke's area in the dominant hemisphere processes dominant word meanings (‘‘teller’’ given ‘‘bank’’).

Modern views

Neuroimaging suggests the functions earlier attributed to Wernicke's area occur more broadly in the temporal lobe and indeed happen also in Broca's area.


Support for a broad range of speech processing areas was furthered by a recent study caried out at the University of Rochester in which American Sign Language native speakers were subject to MRI while interpreting sentences that identified a relationship using either syntax (relationship is determined by the word order) or inflection (relationship is determined by physical motion of "moving hands through space or signing on one side of the body"). Distinct areas of the brain were activated with the frontal cortex (associated with ability to put information into sequences) being more active in the syntax condition and the temporal lobes (associated with dividing information into its constituent parts) being more active in the inflection condition. However, these areas are not mutually exclusive and show a large amount of overlap. These findings imply that while speech processing is a very complex process, the brain may be using fairly basic, preexisting computational methods.

Clinical significance

Human brain with Wernicke's area highlighted in red

Aphasia

Wernicke's area is named after Carl Wernicke, a German neurologist and psychiatrist who, in 1874, hypothesized a link between the left posterior section of the superior temporal gyrus and the reflexive mimicking of words and their syllables that associated the sensory and motor images of spoken words. He did this on the basis of the location of brain injuries that caused aphasia. Receptive aphasia in which such abilities are preserved is also known as Wernicke's aphasia. In this condition there is a major impairment of language comprehension, while speech retains a natural-sounding rhythm and a relatively normal syntax. Language as a result is largely meaningless (a condition sometimes called fluent or jargon aphasia).

While neuroimaging and lesion evidence generally support the idea that malfunction of or damage to Wernicke's area is common in people with receptive aphasia, this is not always so. Some people may use the right hemisphere for language, and isolated damage of Wernicke's area cortex (sparing white matter and other areas) may not cause severe receptive aphasia. Even when patients with Wernicke's area lesions have comprehension deficits, these are usually not restricted to language processing alone. For example, one study found that patients with posterior lesions also had trouble understanding nonverbal sounds like animal and machine noises. In fact, for Wernicke's area, the impairments in nonverbal sounds were statistically stronger than for verbal sounds.

Monday, October 21, 2019

Lateralization of brain function

From Wikipedia, the free encyclopedia

Diagram of the human brain.
The human brain is divided into two hemispheres–left and right. Scientists continue to explore how some cognitive functions tend to be dominated by one side or the other; that is, how they are lateralized.
 
The lateralization of brain function is the tendency for some neural functions or cognitive processes to be specialized to one side of the brain or the other. The medial longitudinal fissure separates the human brain into two distinct cerebral hemispheres, connected by the corpus callosum. Although the macrostructure of the two hemispheres appears to be almost identical, different composition of neuronal networks allows for specialized function that is different in each hemisphere. Lateralization of brain structures is based on general trends expressed in healthy patients; however, there are numerous counterexamples to each generalization. Each human's brain develops differently leading to unique lateralization in individuals. This is different from specialization as lateralization refers only to the function of one structure divided between two hemispheres. Specialization is much easier to observe as a trend since it has a stronger anthropological history. The best example of an established lateralization is that of Broca's and Wernicke's areas where both are often found exclusively on the left hemisphere. These areas frequently correspond to handedness, however, meaning that the localization of these areas is regularly found on the hemisphere corresponding to the dominant hand (anatomically on the opposite side). Function lateralization, such as semantics, intonation, accentuation, and prosody, has since been called into question and largely been found to have a neuronal basis in both hemispheres. Another example is that each hemisphere in the brain tends to represent one side of the body. In the cerebellum this is the same bodyside, but in the forebrain this is predominantly the contralateral side.

Lateralized functions

Language

Language functions such as grammar, vocabulary and literal meaning are typically lateralized to the left hemisphere, especially in right-handed individuals. While language production is left-lateralized in up to 90% of right-handers, it is more bilateral, or even right-lateralized, in approximately 50% of left-handers.

Broca's area and Wernicke's area areas associated with the production of speech and comprehension of speech, respectively, are located in the left cerebral hemisphere for about 95% of right-handers, but about 70% of left-handers.

Sensory processing

The processing of basic sensory information is lateralized by being divided into left and right sides of the body or the space around the body.

In vision, about half the neurons of the optic nerve from each eye cross to project to the opposite hemisphere and about half do not cross to project to the hemisphere on the same side. This means that the left side of the visual field is processed largely by the visual cortex of the right hemisphere and vice versa for the right side of the visual field. 

In audition, about 90% of the neurons of the auditory nerve from one ear cross to project to the auditory cortex of the opposite hemisphere.

In the sense of touch, most of the neurons from the skin cross to project to the somatosensory cortex of the opposite hemisphere. 

Because of this functional division of the left and right sides of the body and of the space that surrounds it, the processing of information in the sensory cortices is essentially identical. That is, the processing of visual and auditory stimuli, spatial manipulation, facial perception, and artistic ability are represented bilaterally. Numerical estimation, comparison and online calculation depend on bilateral parietal regions while exact calculation and fact retrieval are associated with left parietal regions, perhaps due to their ties to linguistic processing.

Value systems

Rather than just being a series of places where different brain modules occur, there are running similarities in the kind of function seen in each side, for instance how right-side impairment of drawing ability making patients draw the parts of the subject matter with wholly incoherent relationships, or where the kind of left-side damage seen in language impairment not damaging the patient's ability to catch the significance of intonation in speech. This has led Iain McGilchrist to say that the two hemispheres as having different value systems, where the left hemisphere tends to reduce complex matters such as ethics to rules and measures, where the right hemisphere is disposed to the holistic and metaphorical.

Clinical significance

Depression is linked with a hyperactive right hemisphere, with evidence of selective involvement in "processing negative emotions, pessimistic thoughts and unconstructive thinking styles", as well as vigilance, arousal and self-reflection, and a relatively hypoactive left hemisphere, "specifically involved in processing pleasurable experiences" and "relatively more involved in decision-making processes". Additionally, "left hemisphere lesions result in an omissive response bias or error pattern whereas right hemisphere lesions result in a commissive response bias or error pattern." The delusional misidentification syndromes, reduplicative paramnesia and Capgras delusion are also often the result of right hemisphere lesions.

Hemisphere damage

Damage to either the right or left hemisphere, and its resulting deficits provide insight into the function of the damaged area. Left hemisphere damage has many effects on language production and perception. Damage or lesions to the right hemisphere can result in a lack of emotional prosody or intonation when speaking. Right hemisphere damage also has grave effects on understanding discourse. People with damage to the right hemisphere have a reduced ability to generate inferences, comprehend and produce main concepts, and a reduced ability to manage alternative meanings. Furthermore, people with right hemisphere damage often exhibit discourse that is abrupt and perfunctory or verbose and excessive. They can also have pragmatic deficits in situations of turn taking, topic maintenance and shared knowledge.

Lateral brain damage can also affect visual perceptual spatial resolution. People with left hemisphere damage may have impaired perception of high resolution, or detailed, aspects of an image. People with right hemisphere damage may have impaired perception of low resolution, or big picture, aspects of an image.

Plasticity

If a specific region of the brain, or even an entire hemisphere, is injured or destroyed, its functions can sometimes be assumed by a neighboring region in the same hemisphere or the corresponding region in the other hemisphere, depending upon the area damaged and the patient's age. When injury interferes with pathways from one area to another, alternative (indirect) connections may develop to communicate information with detached areas, despite the inefficiencies.

Broca's aphasia

Broca's aphasia is a specific type of expressive aphasia and is so named due to the aphasia that results from damage or lesions to the Broca's area of the brain, that exists most commonly in the left inferior frontal hemisphere. Thus, the aphasia that develops from the lack of functioning of the Broca's area is an expressive and non-fluent aphasia. It is called 'non-fluent' due the issues that arise because Broca's area is critical for language pronunciation and production. The area controls some motor aspects of speech production and articulation of thoughts to words and as such lesions to the area result in the specific non-fluent aphasia.

Wernicke's aphasia

Wernicke's aphasia is the result of damage to the area of the brain that is commonly in the left hemisphere above the sylvian fissure. Damage to this area causes primarily a deficit in language comprehension. While the ability to speak fluently with normal melodic intonation is spared, the language produced by a person with Wernicke's aphasia is riddled with semantic errors, and may sound nonsensical to the listener. Wernicke's aphasia is characterized by phonemic paraphasias, neologism or jargon. Another characteristic of a person with Wernicke's aphasia is that they are unconcerned by the mistakes that they are making.

Society and culture

Misapplication

Terence Hines states that the research on brain lateralization is valid as a research program, though commercial promoters have applied it to promote subjects and products far outside the implications of the research. For example, the implications of the research have no bearing on psychological interventions such as EMDR and neurolinguistic programming, brain-training equipment, or management training.

Pop psychology

The oversimplification of lateralization in pop psychology. This belief was widely held even in the scientific community for some years.
 
Some popularizations oversimplify the science about lateralization, by presenting the functional differences between hemispheres as being more absolute than is actually the case.

Sex differences

In the 19th century and to a lesser extent the 20th, it was thought that each side of the brain was associated with a specific gender: the left corresponding with masculinity and the right with femininity and each half could function independently. The right side of the brain was seen as the inferior and thought to be prominent in women, savages, children, criminals, and the insane. A prime example of this in fictional literature can be seen in Robert Louis Stevenson's Strange Case of Dr. Jekyll and Mr. Hyde.

Evolutionary advantage

The widespread lateralization of many vertebrate animals indicates an evolutionary advantage associated with the specialization of each hemisphere.

History

Broca

One of the first indications of brain function lateralization resulted from the research of French physician Pierre Paul Broca, in 1861. His research involved the male patient nicknamed "Tan", who suffered a speech deficit (aphasia); "tan" was one of the few words he could articulate, hence his nickname. In Tan's autopsy, Broca determined he had a syphilitic lesion in the left cerebral hemisphere. This left frontal lobe brain area (Broca's area) is an important speech production region. The motor aspects of speech production deficits caused by damage to Broca's area are known as expressive aphasia. In clinical assessment of this aphasia, it is noted that the patient cannot clearly articulate the language being employed.

Wernicke

German physician Karl Wernicke continued in the vein of Broca's research by studying language deficits unlike expressive aphasia. Wernicke noted that not every deficit was in speech production; some were linguistic. He found that damage to the left posterior, superior temporal gyrus (Wernicke's area) caused language comprehension deficits rather than speech production deficits, a syndrome known as receptive aphasia.

Imaging

These seminal works on hemispheric specialization were done on patients or postmortem brains, raising questions about the potential impact of pathology on the research findings. New methods permit the in vivo comparison of the hemispheres in healthy subjects. Particularly, magnetic resonance imaging (MRI) and positron emission tomography (PET) are important because of their high spatial resolution and ability to image subcortical brain structures.

Movement and sensation

In the 1940s, neurosurgeon Wilder Penfield and his neurologist colleague Herbert Jasper developed a technique of brain mapping to help reduce side effects caused by surgery to treat epilepsy. They stimulated motor and somatosensory cortices of the brain with small electrical currents to activate discrete brain regions. They found that stimulation of one hemisphere's motor cortex produces muscle contraction on the opposite side of the body. Furthermore, the functional map of the motor and sensory cortices is fairly consistent from person to person; Penfield and Jasper's famous pictures of the motor and sensory homunculi were the result.

Split-brain patients

Research by Michael Gazzaniga and Roger Wolcott Sperry in the 1960s on split-brain patients led to an even greater understanding of functional laterality. Split-brain patients are patients who have undergone corpus callosotomy (usually as a treatment for severe epilepsy), a severing of a large part of the corpus callosum. The corpus callosum connects the two hemispheres of the brain and allows them to communicate. When these connections are cut, the two halves of the brain have a reduced capacity to communicate with each other. This led to many interesting behavioral phenomena that allowed Gazzaniga and Sperry to study the contributions of each hemisphere to various cognitive and perceptual processes. One of their main findings was that the right hemisphere was capable of rudimentary language processing, but often has no lexical or grammatical abilities. Eran Zaidel also studied such patients and found some evidence for the right hemisphere having at least some syntactic ability. 

Language is primarily localized in the left hemisphere. One of the experiments carried out by Gazzaniga involved a split-brain male patient sitting in front of a computer screen while having words and images presented on either side of the screen and the visual stimuli would go to either the right or left visual field, and thus the left or right brain, respectively. It was observed that if the patient was presented with an image to his left visual field (right brain), he would report not seeing anything. If he was able to feel around for certain objects, he could accurately pick out the correct object, despite not having the ability to verbalize what he saw. This led to confirmation that the left brain is localized for language whereas the right brain does not have this capability, and when the corpus callosum is cut, the two hemispheres cannot communicate in order for situation-pertinent speech to be produced.

Cryogenics

From Wikipedia, the free encyclopedia https://en.wikipedia.org/wiki/Cryogenics...