Language processing in the brain

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syllables also correlated the hearing of each syllable with its own activation pattern in the pSTG. The involvement of the ADS in both speech perception and production has been further illuminated in several pioneering functional imaging studies that contrasted speech perception with overt or covert speech production. These studies demonstrated that the pSTS is active only during the perception of speech, whereas area Spt is active during both the perception and production of speech. The authors concluded that the pSTS projects to area Spt, which converts the auditory input into articulatory movements. Similar results have been obtained in a study in which participants' temporal and parietal lobes were electrically stimulated. This study reported that electrically stimulating the pSTG region interferes with sentence comprehension and that stimulation of the IPL interferes with the ability to vocalize the names of objects. The authors also reported that stimulation in area Spt and the inferior IPL induced interference during both object-naming and speech-comprehension tasks. The role of the ADS in speech repetition is also congruent with the results of the other functional imaging studies that have localized activation during speech repetition tasks to ADS regions. An intra-cortical recording study that recorded activity throughout most of the temporal, parietal and frontal lobes also reported activation in the pSTG, Spt, IPL and IFG when speech repetition is contrasted with speech perception. Neuropsychological studies have also found that individuals with speech repetition deficits but preserved auditory comprehension (i.e.,

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intermediate phase in the evolution of language. The roles of sound localization and integration of sound location with voices and auditory objects is interpreted as evidence that the origin of speech is the exchange of contact calls (calls used to report location in cases of separation) between mothers and offspring. The role of the ADS in the perception and production of intonations is interpreted as evidence that speech began by modifying the contact calls with intonations, possibly for distinguishing alarm contact calls from safe contact calls. The role of the ADS in encoding the names of objects (phonological long-term memory) is interpreted as evidence of gradual transition from modifying calls with intonations to complete vocal control. The role of the ADS in the integration of lip movements with phonemes and in speech repetition is interpreted as evidence that spoken words were learned by infants mimicking their parents' vocalizations, initially by imitating their lip movements. The role of the ADS in phonological working memory is interpreted as evidence that the words learned through mimicry remained active in the ADS even when not spoken. This resulted with individuals capable of rehearsing a list of vocalizations, which enabled the production of words with several syllables. Further developments in the ADS enabled the rehearsal of lists of words, which provided the infra-structure for communicating with sentences.

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discharges) in the vocal apparatus (mouth, tongue, vocal folds). This feedback marks the sound perceived during speech production as self-produced and can be used to adjust the vocal apparatus to increase the similarity between the perceived and emitted calls. Evidence for descending connections from the IFG to the pSTG has been offered by a study that electrically stimulated the IFG during surgical operations and reported the spread of activation to the pSTG-pSTS-Spt region A study that compared the ability of aphasic patients with frontal, parietal or temporal lobe damage to quickly and repeatedly articulate a string of syllables reported that damage to the frontal lobe interfered with the articulation of both identical syllabic strings ("Bababa") and non-identical syllabic strings ("Badaga"), whereas patients with temporal or parietal lobe damage only exhibited impairment when articulating non-identical syllabic strings. Because the patients with temporal and parietal lobe damage were capable of repeating the syllabic string in the first task, their speech perception and production appears to be relatively preserved, and their deficit in the second task is therefore due to impaired monitoring. Demonstrating the role of the descending ADS connections in monitoring emitted calls, an

497:. Cortical recording and functional imaging studies in macaque monkeys further elaborated on this processing stream by showing that acoustic information flows from the anterior auditory cortex to the temporal pole (TP) and then to the IFG. This pathway is commonly referred to as the auditory ventral stream (AVS; Figure 1, bottom left-red arrows). In contrast to the anterior auditory fields, tracing studies reported that the posterior auditory fields (areas CL-CM) project primarily to dorsolateral prefrontal and premotor cortices (although some projections do terminate in the IFG. Cortical recordings and anatomical tracing studies in monkeys further provided evidence that this processing stream flows from the posterior auditory fields to the frontal lobe via a relay station in the intra-parietal sulcus (IPS). This pathway is commonly referred to as the auditory dorsal stream (ADS; Figure 1, bottom left-blue arrows). Comparing the white matter pathways involved in communication in humans and monkeys with 785:, different phonological representations of the same word share the same semantic representation, this increase in density in the IPL verifies the existence of the phonological lexicon: the semantic lexicon of bilinguals is expected to be similar in size to the semantic lexicon of monolinguals, whereas their phonological lexicon should be twice the size. Consistent with this finding, cortical density in the IPL of monolinguals also correlates with vocabulary size. Notably, the functional dissociation of the AVS and ADS in object-naming tasks is supported by cumulative evidence from reading research showing that semantic errors are correlated with MTG impairment and phonemic errors with IPL impairment. Based on these associations, the semantic analysis of text has been linked to the inferior-temporal gyrus and MTG, and the phonological analysis of text has been linked to the pSTG-Spt- IPL 805:). Studies have also found that speech errors committed during reading are remarkably similar to speech errors made during the recall of recently learned, phonologically similar words from working memory. Patients with IPL damage have also been observed to exhibit both speech production errors and impaired working memory Finally, the view that verbal working memory is the result of temporarily activating phonological representations in the ADS is compatible with recent models describing working memory as the combination of maintaining representations in the mechanism of attention in parallel to temporarily activating representations in long-term memory. It has been argued that the role of the ADS in the rehearsal of lists of words is the reason this pathway is active during sentence comprehension For a review of the role of the ADS in working memory, see. 777:). Semantic paraphasia errors have also been reported in patients receiving intra-cortical electrical stimulation of the AVS (MTG), and phonemic paraphasia errors have been reported in patients whose ADS (pSTG, Spt, and IPL) received intra-cortical electrical stimulation. Further supporting the role of the ADS in object naming is an MEG study that localized activity in the IPL during the learning and during the recall of object names. A study that induced magnetic interference in participants' IPL while they answered questions about an object reported that the participants were capable of answering questions regarding the object's characteristics or perceptual attributes but were impaired when asked whether the word contained two or three syllables. An MEG study has also correlated recovery from 750:(in which hearing the syllable "ba" while seeing the viseme "ga" results in the perception of the syllable "da"). Another study has found that using magnetic stimulation to interfere with processing in this area further disrupts the McGurk illusion. The association of the pSTS with the audio-visual integration of speech has also been demonstrated in a study that presented participants with pictures of faces and spoken words of varying quality. The study reported that the pSTS selects for the combined increase of the clarity of faces and spoken words. Corroborating evidence has been provided by an 549:

demonstrated more selectivity for heard syllables in the anterior Heschl's gyrus (area hR) than posterior Heschl's gyrus (area hA1). In downstream associative auditory fields, studies from both monkeys and humans reported that the border between the anterior and posterior auditory fields (Figure 1-area PC in the monkey and mSTG in the human) processes pitch attributes that are necessary for the recognition of auditory objects. The anterior auditory fields of monkeys were also demonstrated with selectivity for con-specific vocalizations with intra-cortical recordings. and functional imaging One

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refer to the extent that a system's orthography represents morphemes as opposed to phonological segments. Systems that record larger morphosyntactic or phonological segments, such as logographic systems and syllabaries put greater demand on the memory of users. It would thus be expected that an opaque or deep writing system would put greater demand on areas of the brain used for lexical memory than would a system with transparent or shallow orthography.

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anterior primary auditory fields (areas R-RT) projecting to the anterior associative auditory fields (areas AL-RTL), and the posterior primary auditory field (area A1) projecting to the posterior associative auditory fields (areas CL-CM). Recently, evidence accumulated that indicates homology between the human and monkey auditory fields. In humans, histological staining studies revealed two separate auditory fields in the primary auditory region of

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healthy participants and patients with MTG-TP damage, concluded that the MTG-TP in both hemispheres participate in the automatic (rule based) stage of syntactic analysis (ELAN component), and that the left MTG-TP is also involved in a later controlled stage of syntax analysis (P600 component). Patients with damage to the MTG-TP region have also been reported with impaired sentence comprehension. See review for more information on this topic.

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aphasias, RHD signers were found to have a problem maintaining the spatial portion of their signs, confusing similar signs at different locations necessary to communicate with another properly. LHD signers, on the other hand, had similar results to those of hearing patients. Furthermore, other studies have emphasized that sign language is present bilaterally but will need to continue researching to reach a conclusion.

431:. Because almost all language input was thought to funnel via Wernicke's area and all language output to funnel via Broca's area, it became extremely difficult to identify the basic properties of each region. This lack of clear definition for the contribution of Wernicke's and Broca's regions to human language rendered it extremely difficult to identify their homologues in other primates. With the advent of the 1453: 1392: 234: 996:

children and adults suggest that spellers employ phonological rules in spelling regular words and nonwords, while lexical memory is accessed to spell irregular words and high-frequency words of all types. Similarly, lesion studies indicate that lexical memory is used to store irregular words and certain regular words, while phonological rules are used to spell nonwords.

485:), who was shown with reduced bilateral activation in areas hR and aSTG but with spared activation in the mSTG-pSTG. This connectivity pattern is also corroborated by a study that recorded activation from the lateral surface of the auditory cortex and reported of simultaneous non-overlapping activation clusters in the pSTG and mSTG-aSTG while listening to sounds. 742:

study in which participants were instructed to identify syllables also correlated the hearing of each syllable with its own activation pattern in the pSTG. Consistent with the role of the ADS in discriminating phonemes, studies have ascribed the integration of phonemes and their corresponding lip movements (i.e., visemes) to the pSTS of the ADS. For example, an

666:. In both humans and non-human primates, the auditory dorsal stream is responsible for sound localization, and is accordingly known as the auditory 'where' pathway. In humans, this pathway (especially in the left hemisphere) is also responsible for speech production, speech repetition, lip-reading, and phonological working memory and long-term memory. 618:

sentences has been demonstrated in functional imaging studies reporting stronger activation in the anterior MTG when proper sentences are contrasted with lists of words, sentences in a foreign or nonsense language, scrambled sentences, sentences with semantic or syntactic violations and sentence-like sequences of environmental sounds. One

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Single-route models posit that lexical memory is used to store all spellings of words for retrieval in a single process. Dual-route models posit that lexical memory is employed to process irregular and high-frequency regular words, while low-frequency regular words and nonwords are processed using a sub-lexical set of phonological rules.

781:(a disorder characterized by an impaired ability to name objects) with changes in IPL activation. Further supporting the role of the IPL in encoding the sounds of words are studies reporting that, compared to monolinguals, bilinguals have greater cortical density in the IPL but not the MTG. Because evidence shows that, in 939:

By resorting to lesion analyses and neuroimaging, neuroscientists have discovered that whether it be spoken or sign language, human brains process language in general, in a similar manner regarding which area of the brain is being used.Lesion analyses are used to examine the consequences of damage to

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damage was also shown with reduced activation in areas hR and aSTG of both hemispheres when hearing spoken words and environmental sounds. Recordings from the anterior auditory cortex of monkeys while maintaining learned sounds in working memory, and the debilitating effect of induced lesions to this

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studies further demonstrated functional dissociation between the left mSTG and aSTG, with the former processing short speech units (phonemes) and the latter processing longer units (e.g., words, environmental sounds). A study that recorded neural activity directly from the left pSTG and aSTG reported

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monkey study further demonstrated a role of the aSTG in the recognition of individual voices. The role of the human mSTG-aSTG in sound recognition was demonstrated via functional imaging studies that correlated activity in this region with isolation of auditory objects from background noise, and with

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and comparing it to the tonotopic organization of the monkey primary auditory fields, homology was established between the human anterior primary auditory field and monkey area R (denoted in humans as area hR) and the human posterior primary auditory field and the monkey area A1 (denoted in humans as

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was contrasted with closely matching sounds, and the studies were rated for the required level of attention, the authors concluded that attention to phonemes correlates with strong activation in the pSTG-pSTS region. An intra-cortical recording study in which participants were instructed to identify

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resulted in errors during object-naming tasks, and interference in the left IFG resulted in speech arrest. Magnetic interference in the pSTG and IFG of healthy participants also produced speech errors and speech arrest, respectively One study has also reported that electrical stimulation of the left

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Previous hypotheses have been made that damage to Broca's area or Wernicke’s area does not affect sign language being perceived; however, it is not the case. Studies have shown that damage to these areas are similar in results in spoken language where sign errors are present and/or repeated.In both

592:, which is a long-term memory repository of audio-visual representations that are interconnected on the basis of semantic relationships. (See also the reviews by discussing this topic). The primary evidence for this role of the MTG-TP is that patients with damage to this region (e.g., patients with 584:

The latter study further demonstrated that working memory in the AVS is for the acoustic properties of spoken words and that it is independent to working memory in the ADS, which mediates inner speech. Working memory studies in monkeys also suggest that in monkeys, in contrast to humans, the AVS is

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scanning studies, 3 auditory fields were identified in the primary auditory cortex, and 9 associative auditory fields were shown to surround them (Figure 1 top left). Anatomical tracing and lesion studies further indicated of a separation between the anterior and posterior auditory fields, with the

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A growing body of evidence indicates that humans, in addition to having a long-term store for word meanings located in the MTG-TP of the AVS (i.e., the semantic lexicon), also have a long-term store for the names of objects located in the Spt-IPL region of the ADS (i.e., the phonological lexicon).

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In addition to repeating and producing speech, the ADS appears to have a role in monitoring the quality of the speech output. Neuroanatomical evidence suggests that the ADS is equipped with descending connections from the IFG to the pSTG that relay information about motor activity (i.e., corollary

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caused patients to believe that they had spoken when they had not and that IFG stimulation caused patients to unconsciously move their lips. The contribution of the ADS to the process of articulating the names of objects could be dependent on the reception of afferents from the semantic lexicon of

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In addition to extracting meaning from sounds, the MTG-TP region of the AVS appears to have a role in sentence comprehension, possibly by merging concepts together (e.g., merging the concept 'blue' and 'shirt' to create the concept of a 'blue shirt'). The role of the MTG in extracting meaning from

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Top: The auditory cortex of the monkey (left) and human (right) is schematically depicted on the supratemporal plane and observed from above (with the parieto- frontal operculi removed). Bottom: The brain of the monkey (left) and human (right) is schematically depicted and displayed from the side.

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In terms of complexity, writing systems can be characterized as "transparent" or "opaque" and as "shallow" or "deep". A "transparent" system exhibits an obvious correspondence between grapheme and sound, while in an "opaque" system this relationship is less obvious. The terms "shallow" and "deep"

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studies in which the auditory perception of phonemes was contrasted with closely matching sounds, and the studies were rated for the required level of attention, the authors concluded that attention to phonemes correlates with strong activation in the pSTG-pSTS region. An intra-cortical recording

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study instructed participants to speak under normal conditions or when hearing a modified version of their own voice (delayed first formant) and reported that hearing a distorted version of one's own voice results in increased activation in the pSTG. Further demonstrating that the ADS facilitates

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In contradiction to the Wernicke–Lichtheim–Geschwind model that implicates sound recognition to occur solely in the left hemisphere, studies that examined the properties of the right or left hemisphere in isolation via unilateral hemispheric anesthesia (i.e., the WADA procedure) or intra-cortical

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There have been other hypotheses about the lateralization of the two hemispheres. Specifically, the right hemisphere was thought to contribute to the overall communication of a language globally whereas the left hemisphere would be dominant in generating the language locally. Through research in

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The auditory dorsal stream also has non-language related functions, such as sound localization and guidance of eye movements. Recent studies also indicate a role of the ADS in localization of family/tribe members, as a study that recorded from the cortex of an epileptic patient reported that the

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Throughout the 20th century, our knowledge of language processing in the brain was dominated by the Wernicke–Lichtheim–Geschwind model. The Wernicke–Lichtheim–Geschwind model is primarily based on research conducted on brain-damaged individuals who were reported to possess a variety of language

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The single-route model for reading has found support in computer modelling studies, which suggest that readers identify words by their orthographic similarities to phonologically alike words. However, cognitive and lesion studies lean towards the dual-route model. Cognitive spelling studies on

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study in which participants were instructed to read a story further correlated activity in the anterior MTG with the amount of semantic and syntactic content each sentence contained. An EEG study that contrasted cortical activity while reading sentences with and without syntactic violations in

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It is presently unknown why so many functions are ascribed to the human ADS. An attempt to unify these functions under a single framework was conducted in the 'From where to what' model of language evolution In accordance with this model, each function of the ADS indicates of a different

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is often treated as the temporary activation of the representations stored in long-term memory that are used for speech (phonological representations). This sharing of resources between working memory and speech is evident by the finding that speaking during rehearsal results in a significant

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patients) to match spoken words to written words presented to the right or left hemifields, reported vocabulary in the right hemisphere that almost matches in size with the left hemisphere (The right hemisphere vocabulary was equivalent to the vocabulary of a healthy 11-years old child). This

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An issue in the cognitive and neurological study of reading and spelling in English is whether a single-route or dual-route model best describes how literate speakers are able to read and write all three categories of English words according to accepted standards of orthographic correctness.

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Accumulative converging evidence indicates that the AVS is involved in recognizing auditory objects. At the level of the primary auditory cortex, recordings from monkeys showed higher percentage of neurons selective for learned melodic sequences in area R than area A1, and a study in humans

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motor feedback during mimicry is an intra-cortical recording study that contrasted speech perception and repetition. The authors reported that, in addition to activation in the IPL and IFG, speech repetition is characterized by stronger activation in the pSTG than during speech perception.

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deficit in patients after direct intra-cortical electrical stimulation to this same region. Insight into the purpose of speech repetition in the ADS is provided by longitudinal studies of children that correlated the learning of foreign vocabulary with the ability to repeat nonsense words.

516:), and from there to dorsolateral prefrontal and premotor cortices (Figure 1, bottom right-blue arrows), and the aSTG was shown to project to the anterior temporal lobe (middle temporal gyrus-temporal pole; MTG-TP) and from there to the IFG (Figure 1 bottom right-red arrows). 231:

Orange frames mark the region of the auditory cortex, which is displayed in the top sub-figures. Top and Bottom: Blue colors mark regions affiliated with the ADS, and red colors mark regions affiliated with the AVS (dark red and blue regions mark the primary auditory fields).

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Studies of present-day humans have demonstrated a role for the ADS in speech production, particularly in the vocal expression of the names of objects. For instance, in a series of studies in which sub-cortical fibers were directly stimulated interference in the left pSTG and

801:). The involvement of the phonological lexicon in working memory is also evidenced by the tendency of individuals to make more errors when recalling words from a recently learned list of phonologically similar words than from a list of phonologically dissimilar words (the 1619:

Anderson JM, Gilmore R, Roper S, Crosson B, Bauer RM, Nadeau S, Beversdorf DQ, Cibula J, Rogish M, Kortencamp S, Hughes JD, Gonzalez Rothi LJ, Heilman KM (October 1999). "Conduction aphasia and the arcuate fasciculus: A reexamination of the Wernicke–Geschwind model".

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and its application for lesion mappings, however, it was shown that this model is based on incorrect correlations between symptoms and lesions. The refutation of such an influential and dominant model opened the door to new models of language processing in the brain.

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further demonstrated similar patterns of connectivity to the auditory cortex of the monkey. Recording from the surface of the auditory cortex (supra-temporal plane) reported that the anterior Heschl's gyrus (area hR) projects primarily to the middle-anterior

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study that contrasted congruent audio-visual speech with incongruent speech (pictures of still faces) reported pSTS activation. For a review presenting additional converging evidence regarding the role of the pSTS and ADS in phoneme-viseme integration see.

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In the last two decades, significant advances occurred in our understanding of the neural processing of sounds in primates. Initially by recording of neural activity in the auditory cortices of monkeys and later elaborated via histological staining and

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literature also reported that the anterior MTG and TP were consistently active during semantic analysis of speech and text; and an intra-cortical recording study correlated neural discharge in the MTG with the comprehension of intelligible sentences.

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Matsumoto R, Imamura H, Inouchi M, Nakagawa T, Yokoyama Y, Matsuhashi M, Mikuni N, Miyamoto S, Fukuyama H, Takahashi R, Ikeda A (April 2011). "Left anterior temporal cortex actively engages in speech perception: A direct cortical stimulation study".

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Scheich H, Baumgart F, Gaschler-Markefski B, Tegeler C, Tempelmann C, Heinze HJ, Schindler F, Stiller D (February 1998). "Functional magnetic resonance imaging of a human auditory cortex area involved in foreground-background decomposition".

604:). Semantic paraphasias were also expressed by aphasic patients with left MTG-TP damage and were shown to occur in non-aphasic patients after electro-stimulation to this region. or the underlying white matter pathway Two meta-analyses of the 3584:

Roberts AC, Tomic DL, Parkinson CH, Roeling TA, Cutter DJ, Robbins TW, Everitt BJ (May 2007). "Forebrain connectivity of the prefrontal cortex in the marmoset monkey (Callithrix jacchus): an anterograde and retrograde tract-tracing study".

959:

There is a comparatively small body of research on the neurology of reading and writing. Most of the studies performed deal with reading rather than writing or spelling, and the majority of both kinds focus solely on the English language.

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For example, a study examining patients with damage to the AVS (MTG damage) or damage to the ADS (IPL damage) reported that MTG damage results in individuals incorrectly identifying objects (e.g., calling a "goat" a "sheep," an example of

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Cusick CG, Seltzer B, Cola M, Griggs E (September 1995). "Chemoarchitectonics and corticocortical terminations within the superior temporal sulcus of the rhesus monkey: evidence for subdivisions of superior temporal polysensory cortex".

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study of fetuses at their third trimester also demonstrated that area Spt is more selective to female speech than pure tones, and a sub-section of Spt is selective to the speech of their mother in contrast to unfamiliar female voices.

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scan of an auditory agnosia patient demonstrated bilateral reduced activation in the anterior auditory cortices, and bilateral electro-stimulation to these regions in both hemispheres resulted with impaired speech recognition.

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region on working memory recall, further implicate the AVS in maintaining the perceived auditory objects in working memory. In humans, area mSTG-aSTG was also reported active during rehearsal of heard syllables with MEG. and

1011:, but subsequently branches off into different routes depending upon whether or not access to lexical memory or semantic information is needed (which would be expected with irregular words under a dual-route model). A 2007 563:

that the aSTG, but not pSTG, was more active when the patient listened to speech in her native language than unfamiliar foreign language. Consistently, electro stimulation to the aSTG of this patient resulted in impaired

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Willness C (2016-01-08). "The Oxford handbook of organizational climate and culture By Benjamin Schneider & Karen M. Barbera (Eds.) New York, NY: Oxford University Press, 2014. ISBN 978-0-19-986071-5". Book Reviews.

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study that contrasted the perception of audio-visual speech with audio-visual non-speech (pictures and sounds of tools). This study reported the detection of speech-selective compartments in the pSTS. In addition, an

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In terms of spelling, English words can be divided into three categories – regular, irregular, and “novel words” or “nonwords.” Regular words are those in which there is a regular, one-to-one correspondence between

7719:

Zhang Y, Wang C, Zhao X, Chen H, Han Z, Wang Y (September 2010). "Diffusion tensor imaging depicting damage to the arcuate fasciculus in patients with conduction aphasia: a study of the Wernicke–Geschwind model".

356:. In accordance with the 'from where to what' model of language evolution, the reason the ADS is characterized with such a broad range of functions is that each indicates a different stage in language evolution. 488:

Downstream to the auditory cortex, anatomical tracing studies in monkeys delineated projections from the anterior associative auditory fields (areas AL-RTL) to ventral prefrontal and premotor cortices in the

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Vigneau M, Beaucousin V, Hervé PY, Duffau H, Crivello F, Houdé O, Mazoyer B, Tzourio-Mazoyer N (May 2006). "Meta-analyzing left hemisphere language areas: phonology, semantics, and sentence processing".

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Kotz SA, von Cramon DY, Friederici AD (October 2003). "Differentiation of syntactic processes in the left and right anterior temporal lobe: Event-related brain potential evidence from lesion patients".

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There are obvious patterns for utilizing and processing language. In sign language, Broca’s area is activated while processing sign language employs Wernicke’s area similar to that of spoken language.

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region when patients named objects in pictures Intra-cortical electrical stimulation studies also reported that electrical interference to the posterior MTG was correlated with impaired object naming

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Karbe H, Herholz K, Weber-Luxenburger G, Ghaemi M, Heiss WD (June 1998). "Cerebral networks and functional brain asymmetry: evidence from regional metabolic changes during word repetition".

832:, argue that language as a social phenomenon is external to the brain. The individual receives the linguistic system from the outside, and the given language shapes the individual's brain. 980:

in spelling. Irregular words are those in which no such correspondence exists. Nonwords are those that exhibit the expected orthography of regular words but do not carry meaning, such as

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Unsworth N, Engle RW (January 2007). "The nature of individual differences in working memory capacity: active maintenance in primary memory and controlled search from secondary memory".

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in the brainstem which gives rise to the auditory ventral stream. The posterior branch enters the dorsal and posteroventral cochlear nucleus to give rise to the auditory dorsal stream.

1019:, an area used for phonological processing, while the spelling of irregular words produced greater activation of areas used for lexical memory and semantic processing, such as the left 10406:

Jardri R, Houfflin-Debarge V, Delion P, Pruvo JP, Thomas P, Pins D (April 2012). "Assessing fetal response to maternal speech using a noninvasive functional brain imaging technique".

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Stevenson RA, James TW (February 2009). "Audiovisual integration in human superior temporal sulcus: Inverse effectiveness and the neural processing of speech and object recognition".

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Benson RR, Whalen DH, Richardson M, Swainson B, Clark VP, Lai S, Liberman AM (September 2001). "Parametrically dissociating speech and nonspeech perception in the brain using fMRI".

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of the brain. Processing research has failed to find support for the inverse idea that syntactic structures reflect the brain's natural processing preferences cross-linguistically.

710:) suffer from circumscribed damage to the Spt-IPL area or damage to the projections that emanate from this area and target the frontal lobe Studies have also reported a transient 10312:

Mazzoni P, Bracewell RM, Barash S, Andersen RA (March 1996). "Spatially tuned auditory responses in area LIP of macaques performing delayed memory saccades to acoustic targets".

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Cornelissen K, Laine M, Renvall K, Saarinen T, Martin N, Salmelin R (June 2004). "Learning new names for new objects: cortical effects as measured by magnetoencephalography".

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bilateral recognition of sounds is also consistent with the finding that unilateral lesion to the auditory cortex rarely results in deficit to auditory comprehension (i.e.,

773:). Conversely, IPL damage results in individuals correctly identifying the object but incorrectly pronouncing its name (e.g., saying "gof" instead of "goat," an example of 9613: 6266:

Martin RC, Shelton JR, Yaffee LS (February 1994). "Language processing and working memory: Neuropsychological evidence for separate phonological and semantic capacities".

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Lewis JW, Van Essen DC (December 2000). "Corticocortical connections of visual, sensorimotor, and multimodal processing areas in the parietal lobe of the macaque monkey".

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Poremba A, Malloy M, Saunders RC, Carson RE, Herscovitch P, Mishkin M (January 2004). "Species-specific calls evoke asymmetric activity in the monkey's temporal poles".

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Mechelli A, Crinion JT, Noppeney U, O'Doherty J, Ashburner J, Frackowiak RS, Price CJ (October 2004). "Neurolinguistics: structural plasticity in the bilingual brain".

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Kaiser J, Ripper B, Birbaumer N, Lutzenberger W (October 2003). "Dynamics of gamma-band activity in human magnetoencephalogram during auditory pattern working memory".

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Although sound perception is primarily ascribed with the AVS, the ADS appears associated with several aspects of speech perception. For instance, in a meta-analysis of

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Although sound perception is primarily ascribed with the AVS, the ADS appears associated with several aspects of speech perception. For instance, in a meta-analysis of

4064:"Intracortical responses in human and monkey primary auditory cortex support a temporal processing mechanism for encoding of the voice onset time phonetic parameter" 5131:

Roux FE, Miskin K, Durand JB, Sacko O, Réhault E, Tanova R, Démonet JF (October 2015). "Electrostimulation mapping of comprehension of auditory and visual words".

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Seidenberg MS, Petitto LA (1987). "Communication, symbolic communication, and language: Comment on Savage-Rumbaugh, McDonald, Sevcik, Hopkins, and Rupert (1986)".

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techniques indicates of similar connections of the AVS and ADS in the two species (Monkey, Human). In humans, the pSTG was shown to project to the parietal lobe (

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Saur D, Kreher BW, Schnell S, Kümmerer D, Kellmeyer P, Vry MS, Umarova R, Musso M, Glauche V, Abel S, Huber W, Rijntjes M, Hennig J, Weiller C (November 2008).

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Koizumi, Masatoshi; Yasugi, Yoshiho; Tamaoka, Katsuo; Kiyama, Sachiko; Kim, Jungho; Ajsivinac Sian, Juan Esteban; García Mátzar, Pedro Oscar (September 2014).

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de la Mothe LA, Blumell S, Kajikawa Y, Hackett TA (May 2006). "Cortical connections of the auditory cortex in marmoset monkeys: core and medial belt regions".

894:

studies suggest that language processing is based on the interaction of syntax and semantics, and the research does not support innate grammatical structures.

540:. This pathway is responsible for sound recognition, and is accordingly known as the auditory 'what' pathway. The functions of the AVS include the following. 2969:"Resection of the medial temporal lobe disconnects the rostral superior temporal gyrus from some of its projection targets in the frontal lobe and thalamus" 567:(see also for similar results). Intra-cortical recordings from the right and left aSTG further demonstrated that speech is processed laterally to music. An 7002:

Hickok G, Buchsbaum B, Humphries C, Muftuler T (July 2003). "Auditory-motor interaction revealed by fMRI: speech, music, and working memory in area Spt".

7898:

Service E, Kohonen V (April 1995). "Is the relation between phonological memory and foreign language learning accounted for by vocabulary acquisition?".

5603:"Distribution of auditory and visual naming sites in nonlesional temporal lobe epilepsy patients and patients with space-occupying temporal lobe lesions" 2419:

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2282:

Morel A, Garraghty PE, Kaas JH (September 1993). "Tonotopic organization, architectonic fields, and connections of auditory cortex in macaque monkeys".

10631:"Are There Separate Neural Systems for Spelling? New Insights into the Role of Rules and Memory in Spelling from Functional Magnetic Resonance Imaging" 8874:

Lee H, Devlin JT, Shakeshaft C, Stewart LH, Brennan A, Glensman J, Pitcher K, Crinion J, Mechelli A, Frackowiak RS, Green DW, Price CJ (January 2007).

94: 87: 9244:

Waters GS, Rochon E, Caplan D (February 1992). "The role of high-level speech planning in rehearsal: Evidence from patients with apraxia of speech".

7488:

Baldo JV, Klostermann EC, Dronkers NF (May 2008). "It's either a cook or a baker: patients with conduction aphasia get the gist but lose the trace".

45: 8972:

Jobard G, Crivello F, Tzourio-Mazoyer N (October 2003). "Evaluation of the dual route theory of reading: a metanalysis of 35 neuroimaging studies".

6672:

Desmurget M, Reilly KT, Richard N, Szathmari A, Mottolese C, Sirigu A (May 2009). "Movement intention after parietal cortex stimulation in humans".

600:) are reported with an impaired ability to describe visual and auditory objects and a tendency to commit semantic errors when naming objects (i.e., 10269:

Linden JF, Grunewald A, Andersen RA (July 1999). "Responses to auditory stimuli in macaque lateral intraparietal area. II. Behavioral modulation".

9965:"On the (non)universality of the preference for subject-object word order in sentence comprehension: A sentence-processing study in Kaqchikel Maya" 7863:

Quigg M, Geldmacher DS, Elias WJ (May 2006). "Conduction aphasia as a function of the dominant posterior perisylvian cortex. Report of two cases".

5509:

Patterson K, Nestor PJ, Rogers TT (December 2007). "Where do you know what you know? The representation of semantic knowledge in the human brain".

4899:

Shultz S, Vouloumanos A, Pelphrey K (May 2012). "The superior temporal sulcus differentiates communicative and noncommunicative auditory signals".

5655:

Duffau H (March 2008). "The anatomo-functional connectivity of language revisited. New insights provided by electrostimulation and tractography".

1760:

Dronkers NF, Wilkins DP, Van Valin RD, Redfern BB, Jaeger JJ (May 2004). "Lesion analysis of the brain areas involved in language comprehension".

7441:"Brain Regions Underlying Repetition and Auditory-Verbal Short-term Memory Deficits in Aphasia: Evidence from Voxel-based Lesion Symptom Mapping" 5897:

Humphries C, Willard K, Buchsbaum B, Hickok G (June 2001). "Role of anterior temporal cortex in auditory sentence comprehension: an fMRI study".

3019:

Romanski LM, Bates JF, Goldman-Rakic PS (January 1999). "Auditory belt and parabelt projections to the prefrontal cortex in the rhesus monkey".

825:. The primary theoretical question is whether linguistic structures follow from the brain structures or vice versa. Externalist models, such as 6530:

Ulrich G (May 1978). "Interhemispheric functional relationships in auditory agnosia. An analysis of the preconditions and a conceptual model".

2870:"Functional Mapping of the Human Auditory Cortex: fMRI Investigation of a Patient with Auditory Agnosia from Trauma to the Inferior Colliculus" 2765: 137: 9271:

Cohen L, Bachoud-Levi AC (September 1995). "The role of the output phonological buffer in the control of speech timing: a single case study".

5274:

Strominger NL, Oesterreich RE, Neff WD (June 1980). "Sequential auditory and visual discriminations after temporal lobe ablation in monkeys".

6034:

Xu J, Kemeny S, Park G, Frattali C, Braun A (2005). "Language in context: emergent features of word, sentence, and narrative comprehension".

2462:

Wallace MN, Johnston PW, Palmer AR (April 2002). "Histochemical identification of cortical areas in the auditory region of the human brain".

109: 3259:

Romanski LM, Averbeck BB, Diltz M (February 2005). "Neural representation of vocalizations in the primate ventrolateral prefrontal cortex".

846: 968:. Another difficulty is that some studies focus on spelling words of English and omit the few logographic characters found in the script. 944:

types of languages, they are affected by damage to the left hemisphere of the brain rather than the right -usually dealing with the arts.

1573:

Aboitiz F, García VR (December 1997). "The evolutionary origin of the language areas in the human brain. A neuroanatomical perspective".

3032: 116: 5985:"The role of left inferior frontal and superior temporal cortex in sentence comprehension: localizing syntactic and semantic processes" 4015:"Early stages of melody processing: stimulus-sequence and task-dependent neuronal activity in monkey auditory cortical fields A1 and R" 2397: 936:. There are over 135 discrete sign languages around the world- making use of different accents formed by separate areas of a country. 7166:

Giraud AL, Price CJ (August 2001). "The constraints functional neuroimaging places on classical models of auditory word processing".

5745:

Creutzfeldt O, Ojemann G, Lettich E (October 1989). "Neuronal activity in the human lateral temporal lobe. I. Responses to speech".

7582:"Conduction aphasia, sensory-motor integration, and phonological short-term memory - an aggregate analysis of lesion and fMRI data" 473:(mSTG-aSTG) and the posterior Heschl's gyrus (area hA1) projects primarily to the posterior superior temporal gyrus (pSTG) and the 8209:"fMRI-Guided transcranial magnetic stimulation reveals that the superior temporal sulcus is a cortical locus of the McGurk effect" 7037:

Warren JE, Wise RJ, Warren JD (December 2005). "Sounds do-able: auditory-motor transformations and the posterior temporal plane".

123: 10075:

Tian B, Reser D, Durham A, Kustov A, Rauschecker JP (April 2001). "Functional specialization in rhesus monkey auditory cortex".

9922:

Wei, Xuehu; Adamson, Helyne; Schwendemann, Matthias; Goucha, Tómas; Friederici, Angela D.; Anwander, Alfred (19 February 2023).

9632: 9201:

Caplan D, Rochon E, Waters GS (August 1992). "Articulatory and phonological determinants of word length effects in span tasks".

1015:

study found that subjects asked to produce regular words in a spelling task exhibited greater activation in the left posterior

940:

specific brain regions involved in language while neuroimaging explore regions that are engaged in the processing of language.

645:), whereas a second lesion to the remaining hemisphere (which could occur years later) does. Finally, as mentioned earlier, an 3554: 10704: 6295:"Damage to left anterior temporal cortex predicts impairment of complex syntactic processing: a lesion-symptom mapping study" 3162:

Petkov CI, Kayser C, Steudel T, Whittingstall K, Augath M, Logothetis NK (March 2008). "A voice region in the monkey brain".

1557: 1487: 1012: 1004: 916: 756: 751: 743: 738: 725: 698: 646: 619: 605: 581: 568: 559: 550: 478: 460: 451: 432: 105: 51: 6079:"Selective attention to semantic and syntactic features modulates sentence processing networks in anterior temporal cortex" 1042:

Far less information exists on the cognition and neurology of non-alphabetic and non-English scripts. Every language has a

3905:"The neural architecture of the language comprehension network: converging evidence from lesion and connectivity analyses" 6293:

Magnusdottir S, Fillmore P, den Ouden DB, Hjaltason H, Rorden C, Kjartansson O, Bonilha L, Fridriksson J (October 2013).

3062:

Tanaka D (June 1976). "Thalamic projections of the dorsomedial prefrontal cortex in the rhesus monkey (Macaca mulatta)".

2384:

Rauschecker JP, Tian B, Pons T, Mishkin M (May 1997). "Serial and parallel processing in rhesus monkey auditory cortex".

2026:

Rauschecker JP, Tian B, Hauser M (April 1995). "Processing of complex sounds in the macaque nonprimary auditory cortex".

1039:

and left SMG that are also important in reading, suggesting that a similar pathway is used for both reading and writing.

685:

the AVS, as an intra-cortical recording study reported of activation in the posterior MTG prior to activation in the Spt-

5798:

Mazoyer BM, Tzourio N, Frak V, Syrota A, Murayama N, Levrier O, Salamon G, Dehaene S, Cohen L, Mehler J (October 1993).

5470:"Temporal lobe lesions and semantic impairment: a comparison of herpes simplex virus encephalitis and semantic dementia" 10226:

Barrett DJ, Hall DA (August 2006). "Response preferences for "what" and "where" in human non-primary auditory cortex".

7310:

Selnes OA, Knopman DS, Niccum N, Rubens AB (June 1985). "The critical role of Wernicke's area in sentence repetition".

4756:

Lewis JW, Phinney RE, Brefczynski-Lewis JA, DeYoe EA (August 2006). "Lefties get it "right" when hearing tool sounds".

2816:"Evidence of functional connectivity between auditory cortical areas revealed by amplitude modulation sound processing" 5362:"Human dorsal and ventral auditory streams subserve rehearsal-based and echoic processes during verbal working memory" 2869: 9725: 9665: 9626: 2766:"Temporal envelope processing in the human auditory cortex: response and interconnections of auditory cortical areas" 1470:

Pickles JO (2015). "Chapter 1: Auditory pathways: anatomy and physiology". In Aminoff MJ, Boller F, Swaab DF (eds.).

210: 156: 59: 8497:"Anterior temporal involvement in semantic word retrieval: voxel-based lesion-symptom mapping evidence from aphasia" 7353:

Axer H, von Keyserlingk AG, Berks G, von Keyserlingk DG (March 2001). "Supra- and infrasylvian conduction aphasia".

5554:"Anterior temporal involvement in semantic word retrieval: voxel-based lesion-symptom mapping evidence from aphasia" 3633:"Association fibre pathways of the brain: parallel observations from diffusion spectrum imaging and autoradiography" 265:

and understood. Language processing is considered to be a uniquely human ability that is not produced with the same

2918:

Chang EF, Edwards E, Nagarajan SS, Fogelson N, Dalal SS, Canolty RT, Kirsch HE, Barbaro NM, Knight RT (June 2011).

10561: 8544:

Ojemann GA (June 1983). "Brain organization for language from the perspective of electrical stimulation mapping".

4482:

Obleser J, Boecker H, Drzezga A, Haslinger B, Hennenlotter A, Roettinger M, Eulitz C, Rauschecker JP (July 2006).

3864:

Menjot de Champfleur N, Lima Maldonado I, Moritz-Gasser S, Machi P, Le Bars E, Bonafé A, Duffau H (January 2013).

3441:"Selectivity for the spatial and nonspatial attributes of auditory stimuli in the ventrolateral prefrontal cortex" 9314:

Shallice T, Rumiati RI, Zadini A (September 2000). "The selective impairment of the phonological output buffer".

8350:"Visual phonetic processing localized using speech and nonspeech face gestures in video and point-light displays" 4153: 6922:"Role of left posterior superior temporal gyrus in phonological processing for speech perception and production" 3866:"Middle longitudinal fasciculus delineation within language pathways: a diffusion tensor imaging study in human" 554:

the recognition of spoken words, voices, melodies, environmental sounds, and non-speech communicative sounds. A

5799: 3672:

Seltzer B, Pandya DN (July 1984). "Further observations on parieto-temporal connections in the rhesus monkey".

1110: 802: 415:

related disorders. In accordance with this model, words are perceived via a specialized word reception center (

10187:"Automatic and intrinsic auditory "what" and "where" processing in humans revealed by electrical neuroimaging" 7672:"Language dysfunction after stroke and damage to white matter tracts evaluated using diffusion tensor imaging" 4709:"Cortical representation of natural complex sounds: effects of acoustic features and auditory object category" 3490:

Deacon TW (February 1992). "Cortical connections of the inferior arcuate sulcus cortex in the macaque brain".

477:(area PT; Figure 1 top right). Consistent with connections from area hR to the aSTG and hA1 to the pSTG is an 130: 10746: 4850: 1031:. Spelling nonwords was found to access members of both pathways, such as the left STG and bilateral MTG and 10502:"From Mimicry to Language: A Neuroanatomically Based Evolutionary Model of the Emergence of Vocal Language" 4851:"Human auditory belt areas specialized in sound recognition: a functional magnetic resonance imaging study" 1411:"From Mimicry to Language: A Neuroanatomically Based Evolutionary Model of the Emergence of Vocal Language" 1090: 1000: 328:

pathway is responsible for sound recognition, and is accordingly known as the auditory 'what' pathway. The

297: 277: 9566:

Buchsbaum BR, D'Esposito M (May 2008). "The search for the phonological store: from loop to convolution".

4302:"Where is the semantic system? A critical review and meta-analysis of 120 functional neuroimaging studies" 636:

is processed bilaterally. Moreover, a study that instructed patients with disconnected hemispheres (i.e.,

10018:"Populations of auditory cortical neurons can accurately encode acoustic space across stimulus intensity" 7531:

Fridriksson J, Kjartansson O, Morgan PS, Hjaltason H, Magnusdottir S, Bonilha L, Rorden C (August 2010).

6393:"Bilateral capacity for speech sound processing in auditory comprehension: evidence from Wada procedures" 5850:"Response of anterior temporal cortex to syntactic and prosodic manipulations during sentence processing" 2607:"Mapping the tonotopic organization in human auditory cortex with minimally salient acoustic stimulation" 20: 1066:, are phonographic. Most systems combine the two and have both logographic and phonographic characters. 10756: 10451:"From where to what: a neuroanatomically based evolutionary model of the emergence of speech in humans" 1350:"From where to what: a neuroanatomically based evolutionary model of the emergence of speech in humans" 933: 895: 850: 336:, and is accordingly known as the auditory 'where' pathway. In humans, this pathway (especially in the 8925:"Contrasting effects of vocabulary knowledge on temporal and parietal brain structure across lifespan" 5468:

Noppeney U, Patterson K, Tyler LK, Moss H, Stamatakis EA, Bright P, Mummery C, Price CJ (April 2007).

4621:

Belin P, Zatorre RJ (November 2003). "Adaptation to speaker's voice in right anterior temporal lobe".

459:, and by mapping the tonotopic organization of the human primary auditory fields with high resolution 10751: 8160:"A neural basis for interindividual differences in the McGurk effect, a multisensory speech illusion" 7211:"The left posterior superior temporal gyrus participates specifically in accessing lexical phonology" 5235:"The effect of temporal lobe and hippocampal lesions on auditory and visual recent memory in monkeys" 4799:

Maeder PP, Meuli RA, Adriani M, Bellmann A, Fornari E, Thiran JP, Pittet A, Clarke S (October 2001).

10726: 5940:

Vandenberghe R, Nobre AC, Price CJ (May 2002). "The response of left temporal cortex to sentences".

3766:"Dissociating the human language pathways with high angular resolution diffusion fiber tractography" 2656:"Extensive cochleotopic mapping of human auditory cortical fields obtained with phase-encoding fMRI" 1301:"The cortical organization of lexical knowledge: a dual lexicon model of spoken language processing" 7263:"ECoG gamma activity during a language task: differentiating expressive and receptive speech areas" 4800: 1043: 798: 686: 681: 676: 597: 513: 509: 505: 420: 9017:"Cross-cultural effect on the brain revisited: universal structures plus writing system variation" 7261:

Towle VL, Yoon HA, Castelle M, Edgar JC, Biassou NM, Frim DM, Spire JP, Kohrman MH (August 2008).

3817:"Delineation of the middle longitudinal fascicle in humans: a quantitative, in vivo, DT-MRI study" 883:, advocate a two-way model arguing that the brain shapes language, and language shapes the brain. 8799:"Exploring cross-linguistic vocabulary effects on brain structures using voxel-based morphometry" 8495:

Schwartz MF, Kimberg DY, Walker GM, Faseyitan O, Brecher A, Dell GS, Coslett HB (December 2009).

7978:"Functional connectivity in the human language system: a cortico-cortical evoked potential study" 7631:"MR tractography depicting damage to the arcuate fasciculus in a patient with conduction aphasia" 5552:

Schwartz MF, Kimberg DY, Walker GM, Faseyitan O, Brecher A, Dell GS, Coslett HB (December 2009).

1815:"The Wernicke conundrum and the anatomy of language comprehension in primary progressive aphasia" 1100: 1032: 1016: 891: 880: 470: 83: 10347:

Lachaux JP, Jerbi K, Bertrand O, Minotti L, Hoffmann D, Schoendorff B, Kahane P (October 2007).

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Oberauer K (2002). "Access to information in working memory: Exploring the focus of attention".

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Lachaux JP, Jerbi K, Bertrand O, Minotti L, Hoffmann D, Schoendorff B, Kahane P (October 2007).

3631:

Schmahmann JD, Pandya DN, Wang R, Dai G, D'Arceuil HE, de Crespigny AJ, Wedeen VJ (March 2007).

2125:"Cortical connections of auditory cortex in marmoset monkeys: lateral belt and parabelt regions" 10761: 9681:

Lakoff, George (1990). "Invariance hypothesis: is abstract reasoning based on image-schemas?".

7580:

Buchsbaum BR, Baldo J, Okada K, Berman KF, Dronkers N, D'Esposito M, Hickok G (December 2011).

1020: 858: 829: 812:

The 'from where to what' model of language evolution hypotheses 7 stages of language evolution.

663: 537: 490: 428: 313: 301: 9827: 6723:

Edwards E, Nagarajan SS, Dalal SS, Canolty RT, Kirsch HE, Barbaro NM, Knight RT (March 2010).

9867: 9609: 9359:"The selective impairment of the phonological output buffer: evidence from a Chinese patient" 8017:

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7976:

Matsumoto R, Nair DR, LaPresto E, Najm I, Bingaman W, Shibasaki H, Lüders HO (October 2004).

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Throughout the 20th century the dominant model for language processing in the brain was the

228:

Dual stream connectivity between the auditory cortex and frontal lobe of monkeys and humans.

10360: 10139: 10084: 10029: 9081: 8745: 8643: 8273: 7765:"Sensory-to-motor integration during auditory repetition: a combined fMRI and lesion study" 7763:

Jones OP, Prejawa S, Hope TM, Oberhuber M, Seghier ML, Leff AP, Green DW, Price CJ (2014).

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Hickok G, Okada K, Barr W, Pa J, Rogalsky C, Donnelly K, Barde L, Grant A (December 2008).

6183:"Syntactic structure building in the anterior temporal lobe during natural story listening" 5668: 5422: 5187: 5084: 5021: 4955: 3967: 3217: 2667: 2338: 2185: 2035: 876: 316:

has been developed. In accordance with this model, there are two pathways that connect the

285: 8766: 7816:"Conduction aphasia elicited by stimulation of the left posterior superior temporal gyrus" 6011: 5767: 2814:

Guéguin M, Le Bouquin-Jeannès R, Faucon G, Chauvel P, Liégeois-Chauvel C (February 2007).

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6725:"Spatiotemporal imaging of cortical activation during verb generation and picture naming" 4849:

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Please help update this article to reflect recent events or newly available information.

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Narain C, Scott SK, Wise RJ, Rosen S, Leff A, Iversen SD, Matthews PM (December 2003).

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Poliva O, Bestelmeyer PE, Hall M, Bultitude JH, Koller K, Rafal RD (September 2015).

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