Broca's area

Brain: Broca's area
Approximate location of Broca's area highlighted in gray
Broca's area visible but not labeled.
Part of	Frontal lobe
Artery	Middle cerebral
Vein	Superior sagittal sinus
NeuroNames	ancil-251

Broca's area is a region of the brain responsible for speech production.

The importance of Broca’s area in producing language has been recognized since Paul Pierre Broca reported impairments in two patients he encountered. They had lost the ability to speak after injury to the posterior inferior frontal gyrus of the brain.^[1] Since then, the approximate region he identified has become known as Broca’s area, and the deficit in language production as Broca’s 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 areas 44 and 45.^[1] Studies of chronic aphasia have implicated an essential role of Broca’s area in various speech and language functions. Further, functional MRI studies have also identified activation patterns in Broca’s area associated with various language tasks.

Contents 1 Anatomy and connectivity 2 Broca's area revisited 2.1 Broca's patients 2.1.1 Leborgne 2.1.2 Lelong 2.2 MRI findings 3 Functions 3.1 Language comprehension 3.2 Action recognition and production 3.3 Speech-associated gestures 4 Aphasia 4.1 Broca's aphasia vs. other aphasias 5 Evolution of language 6 See also 7 References

Anatomy and connectivity

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 on to 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 brain in terms of shape, size, and position relative to sulcal and gyral structure, a resulting localization precision is limited.^[2]

Nevertheless, Broca’s area in the left hemisphere and its homologue in the right hemisphere are designations usually used to refer to pars triangularis (PTr) and pars opercularis (POp) of the inferior frontal gyrus. 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.^[3]

Area 45 receives more afferent connections from 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.^[3]

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, coressponding to areas 45 and 44, respectively, play different functional roles in the human with respect to language comprehension and action recognition/understanding.^[3]

Broca's area revisited

In a recent study, the preserved brains of both Leborgne and Lelong (patients of Paul Pierre 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.^[1]

Broca's patients

Leborgne

Leborgne was a patient of Paul Pierre Broca. He was unable to produce any words or phrases. The only word he could repetitively produce was 'tan'. After his death, a lesion was discovered on the surface of the left frontal lobe.

Lelong

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

MRI findings

Examination of the brains of Paul Pierre 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 interesting finding is that the lesion, 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 that language and cognition are far more complicated than once thought and involve various networks of brain regions.

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, recent evidence demonstrates 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.^[4] 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.^[5] Further, it has been recently found in functional magnetic resonance imaging (fMRI) experiments involving highly ambiguous sentences result in more activated inferior frontal gyrus.^[3] Therefore, activity level in 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.

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.^[6] An activation of BA 44 was also reported during execution of grasping and manipulation.^[7]

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.^[3]

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 goal and intention.^[8] 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.^[6],^[9] (see Evolution of Language below)

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.^[10]

Broca's aphasia vs. other aphasias

Patients with Broca's aphasia are individuals who know "what they want to say, they just cannot get it out." ^[10] They are typically able to understand what is being said to them, but unable to fluently speak. This is also known as non-fluent aphasia. Some of other symptoms may include problems with fluency, articulation, word-finding, repetition, and producing and comprehending complex grammatical sentences, both orally and in writing.^[1] These characteristics distinguish them from other individuals with other types of aphasia. Other aphasia types may have more difficulty understanding what is said to them. They may also struggle more with reading and writing than do individuals with Broca’s aphasia. While the individuals with Broca’s aphasia also have a good ability to self-monitor their language output, other types of aphasia may be more unaware of their language performance. Also, site of lesion (brain damaged area) differs between the different aphasias.

Major Characteristics of Different Types of Aphasia^[10]

Type of Aphasia	Repetition	Naming	Auditory Comprehension	Fluency
Broca's	Mod-severe	Mod-severe	Mild difficulty	Non-fluent, effortful, slow
Wernicke's	Mild-severe	Mild-severe	Defective	Fluent paraphasic
Conduction	Poor	Poor	Relatively good	Fluent
Mixed Transcortical	Moderate	Poor	Poor	Non-fluent
Transcortical Motor	Good	Mild-severe	Mild	Non-fluent
Transcortical Sensory	Good	Mod-severe	Poor	Fluent
Global	Poor	Poor	Poor	Non-fluent
Anomic	Mild	Mod-severe	Mild	Fluent

Evolution of language

Several models have been proposed to explain the origin of human language. Human language is thought to 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.”^[6] The recent finding that Broca’s area is involved during meaningful action observation supports this idea. It was hypothesized that Broca’s area precursor was involved in generating action meanings by interpreting motor sequences in terms of goal. It was further argued that this ability might have been generalized during the evolution that gave this area the capability to deal with meanings. The activated frontal language areas when observing meaningful hand shadows resembling moving animals provides evidence that the human language may have evolved from neural substrates already involved in gestural recognition. Therefore, the study has demonstrated human Broca’s area as the motor center for speech, assembling and decoding communicative 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.^[9]

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.^[11] 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.

See also

• Broca's aphasia
• Wernicke's area
• pars opercularis
• pars triangularis

References

v • d • e Brain: telencephalon (cerebrum · cerebral cortex · cerebral hemispheres) Primary four surface lobes Frontal lobe Precentral gyrus (Primary motor cortex, 4) Superior frontal gyrus/Frontal eye fields (6, 8, 9) · Middle frontal gyrus (46) · Inferior frontal gyrus/Broca's area (44-Pars opercularis, 45-Pars triangularis) Orbitofrontal cortex (10, 11, 12, 47) Prefrontal cortex · Premotor cortex Precentral sulcus · Superior frontal sulcus · Inferior frontal sulcus · Olfactory sulcus Parietal lobe Somatosensory cortex (Primary (1 · 2 · 3 · 43) · Secondary (5)) · Precuneus (7m) · Parietal operculum Parietal lobules (Superior (7l) · Inferior (40)) · Angular gyrus (39) Intraparietal sulcus · Marginal sulcus Occipital lobe Primary visual cortex (17) · (Cuneus · Lingual gyrus · Lateral occipital gyrus (18, 19)) Calcarine fissure Temporal lobe Primary auditory cortex (41, 42) · Superior temporal gyrus (38, 22/Wernicke's area) · Middle temporal gyrus (21) · Inferior temporal gyrus (20) Fusiform gyrus (37) Medial temporal lobe (Amygdala · Parahippocampal gyrus (27 · 28 · 34 · 35 · 36) Interlobar sulci/fissures lateral: Central (frontal+parietal) · Lateral (frontal+parietal+temporal) · Parietoöccipital medial: Medial longitudinal · Cingulate (frontal+cingulate) · Collateral (temporal+occipital) Cingulate cortex/gyrus Subgenual area (25) · Anterior cingulate (24, 32, 33) · Posterior cingulate (23, 31) · Retrosplenial cortex (26, 29, 30) Callosal sulcus White matter tracts Commissural fibers · Association fibers Internal capsule (Anterior limb · Genu · Posterior limb) · Corona radiata · External capsule · Lamina terminalis · Extreme capsule · Semioval center Olfactory tract · Terminal stria · Diagonal band of Broca Other Insular cortex gray: Olfactory bulb · Anterior olfactory nucleus · Basal optic nucleus of Meynert · Substantia innominata · Anterior perforated substance Corpus striatum · Limbic lobe Some categorizations are approximations, and some Brodmann areas span gyri.