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Lecture for IIIT Hyderabad Cognitive Neuroscience course 2011
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NEUROLINGUISTICSCompiled by Dr PS Deb
NEUROLINGUISTICS
Neurolinguistics is the study of the neural mechanisms in the human brain that control the comprehension, production, and acquisition of language.
Harry Whitaker, who founded the Journal of Neurolinguistics in 1985
PET, fMRI, EEG, MEG help to find the language related brain activity in normal individual
An interdisciplinary field, such as neuroscience, linguistics, cognitive science, neurobiology, communication disorders, neuropsychology, and computer science.
Subfield DescriptionResearch questions in
neurolinguistics
Phonetics the study of speech sounds
how the brain extracts speech sounds from an acoustic signal, how the brain separates speech sounds from background noise
Phonologythe study of how sounds are organized in a language
how the phonological system of a particular language is represented in the brain
Morphology and lexicology
the study of how words are structured and stored in the mental lexicon
how the brain stores and accesses words that a person knows
Syntaxthe study of how multiple-word utterances are constructed
how the brain combines words into constituents and sentences; how structural and semantic information is used in understanding sentencesSemantics
the study of how meaning is encoded in language
LOCALIZATIONS OF LANGUAGE PROCESSES (AREOLOGY)
Investigated the locations of specific language "modules" within the brain. What course language information follows through the brain as it is processed. Whether or not particular areas specialize in processing particular sorts of
information. How different brain regions interact with one another in language processing. How the locations of brain activation differs when a subject is producing or
perceiving a language other than his or her first language.
TIME COURSE OF LANGUAGE PROCESSES The use
of electrophysiological techniques to analyze the rapid processing of language in time.
The temporal ordering of specific peaks in brain activity may reflect discrete computational processes that the brain undergoes during language processing.
E.g. one neurolinguistic theory of sentence parsing proposes that three brain responses (the ELAN, N400, and P600) are products of three different steps in syntactic and semantic processing
LANGUAGE ACQUISITION The relationship between
brain structures and language acquisition
Infants from all linguistic environments go through similar and predictable stages (such as babbling),
To find correlations between stages of language development and stages of brain development.
The physical changes (known as neuroplasticity) that the brain undergoes during second language acquisition, when adults learn a new language.
NEUROLINGUISTICS METHODS
Lesion studies Neuroanatomy Stimulation studies Neuroimaging Electrophysiology Psycholiguistics Language Pathology
EXPERIMENTAL PARADIGMS
Mismatch paradigm Violation-based Priming Stimulation Subject tasks Lexical decision Grammaticality judgment, acceptability
judgment Probe verification Truth-value judgment Active distraction and double-task
HISTROY
Willium Wunddt: Founder of experimental psychology. Language as mechanism to transform thought into sentence
Gall: Phrenology Broca : First case of
expressive aphasia Wernikes: First case of
comprehension aphasia
LANGUAGE AREA IN PHRENOLOGY
BROCA’S APHASIA
Autopsy of a patient who could understand, with normal speech apparatus but could not speak or write a sentence.
Only articulate sound he could make was “tan”
After autopsying eight similar patient with lesion in the left frontal lobe
He made a famous statement that “we speak with the left hemisphere”1. language articulation lies the third frontal convolution of the inferior frontal gyrus; 2. there is left hemisphere dominance in language articulation;3. understanding language is a different cognitive task than producing it.
Paul Broca 1861
WERNICKE’S APHASIA
Ten years later, Carl Wernicke, a German neurologist,
discovered another part of the brain, this one involved in
understanding language, in the posterior portion of the left
temporal lobe.
People who had a lesion at this location could speak, but
their speech was often incoherent and made no sense.
Carl Wernicke 1871
The Aphasic Symptom complex (1874)
language processing is distributed in the brain, which is the central idea of most current cognitive models. He also made the correct prediction that patients with damage in the arcuate fasciculus would not be able to repeat speech sounds, a dysfunction later named as Conduction aphasia.
Wernicke’sBroca’s
LICHTHEIM’S HOUSE MODEL 1885WERNICKE-LICHTHEIM’S CONNECTIONISM
B
A M
THE WERNICKE-GESCHWIND MODEL OF LANGUAGE
B
A M V
HANDEDNESS AND LANGUAGE
THE BRAIN’S ANATOMICAL ASYMMETRY
LATERALIZATION OF LANGUAGE: WADA TEST
SPIT BRAIN EXPERIMENT
CHALLENGING CLASSICAL MODEL
BF Skiner: Behaviourist – Verbal behaviour 1957, Learning of language can be seen as mechanism
of reinforcement Noem Chomsky 1957 Syntactic structure
Ability to invent language is coded in gene and basis of understanding is similar across culture
There might be universal grammar Language is more widely represented than
older model Role of Motor cortex in comprehension Role of Right hemisphere more than realized
CORTICAL MAPPING OF THE LANGUAGE AREAS IN THE LEFT CEREBRAL CORTEX DURING NEUROSURGERY
(A) Location of the classical language areas.
(B) Evidence for the variability of language representation among individuals.
The number in each circle indicates the percentage of the patients who showed interference with language in response to stimulation at that site.
Note also that many of the sites that elicited interference fall outside the classic language areas. (B after Ojemann et al., 1989.)
LANGUAGE RELATED AREA OF LEFT BRAIN PET
GESCHWIND’S TERRITORY
the inferior parietal lobule is connected by large bundles of nerve fibres to both Broca’s area and Wernicke’s area.
Information might therefore travel between these last two areas either directly, via the arcuate fasciculus, or by a second, parallel route that passes through the inferior parietal lobule.
The inferior parietal lobule is one of the last structures of the human brain to have developed in the course of evolution
AREA 24: INITIATION AND MAINTENANCE OF SPEECH
They are also important to attention and emotion and thus can influence many
higher functions. Damage to these areas does not cause an aphasia in the proper sense but impairs the initiation of movement (akinesia) and causes mutism, the complete absence of speech. Mutism is a rarity in aphasic patients and is seen only during the very early stages of the condition. Patients with akinesia and mutism fail to communicate by words, gestures, or facial expression. They have an impairment of the drive to communicate, rather than aphasia.
LANGUAGE-RELATED AREAS IN THE HUMAN BRAIN.
The implementation system is made up of several regions located around the left sylvian fissure. It includes the classical language areas (B = Broca's area; W = Wernicke's area) and the adjoining supramarginal gyrus (Sm), angular gyrus (AG), auditory cortex (A), motor cortex (M), and somatosensory cortex (Ss). The posterior and anterior components of the implementation system, respectively Wernicke's area and Broca's area, are interconnected by the arcuate fasciculusThe mediational system surrounds the implementation system like a belt (blue areas). The regions identified so far are located in the left temporal pole (TP), left inferotemporal cortex (It), and left prefrontal cortex (Pf). The left basal ganglia complex (not pictured) is an integral part of the language implementation system
THE BRAIN CORRELATES OF A GIVEN COGNITIVE REPRESENTATIONRepresentation R and process P implies answers to (at least) four critical
questions:
1. Where-question: Which brain parts, areas, and, eventually, neurons are
active during, and are critical for, process P and the representation(s) R P
relies on?
2. When-question: At which point in time in the usage or understanding of
language does process P occur; when is representation R activated and
processed?
3. How-question: Which neuronal circuit, which nerve cells linked in which
way, is the brain basis for representation R; which spatiotemporal pattern
of neuronal activation in this circuit does underpin the process P?
4. Why-question: For what reason are R and P located in these specific brain
parts and activated at these specific points in time, and why is R laid down
in this specific neuronal circuit, P being expressed by these specific
activation patterns?
SEARCH FOR MEANING CENTERS?
PET SPEAKING TASK (NAMING)
BRAIN ACTIVATION PATTERNS DURING PASSIVE WORD READING
SCHEMATIC ILLUSTRATION OF THE CORTICAL SYSTEMS FOR LANGUAGE AND ACTION
PHONOLOGICAL PROCESSING OF SPEECH SOUND
The phonemes [p] and [t] for example were mapped to adjacent areas in superiortemporal gyrus, anterior to primary auditory cortex and Heschl’s gyrus.
A similar phonological mapping was evident in the motor system, where the production of [p] and [t] activated different precentral areas in a soma-totopic fashion.
The articulatory mapping of phonemes to the motor system corresponded to the localization of the articulators mainly involved in the production of the respective speech sounds — the lips for [p] and the tongue for [t]
SYNTACTICAL PROCESSING
When comparing grammatical sentences to word strings with syntactic errors, the latter elicit stronger brain activation in left perisylvian cortex, especially in inferior-frontal and in superior-temporal cortex
When directly comparing sentences with different grammatical structure, for example active and passive, subject and object relative, and coordinated and subordinated sentences, the grammatically more demanding sentences tended to elicit stronger activation; again some of the activation differences were located in left perisylvian cortex
SYNTACTIC NETWORKS IN THE HUMAN BRAIN
The frontal operculum and BA 44 together with the anterior STG, dealing with the structure of phrases (noun phrase, prepositional phrase etc.) and
BA 44/45, together with the posterior STG, being responsible for thematic role assignment
Within the latter network, BA 44 seems to process hierarchical structures independent of semantics (i.e. in natural and semantic-free artificial grammars [44,45]), whereas
The posterior STG seems to support the integration of syntactic and semantic information to achieve understanding (that is to understand who is doing what to whom).
THE RAPID TIME COURSE OF LANGUAGE UNDERSTANDING Language understanding is a relatively late
process Semantic processing, along with lexical ones, were
assumed to be first indexed by the N400 component of the event-related brain potential and field.
Syntactic processing was assumed to be indexed by an even later component, called P600.
Early near-simultaneous brain responses (latency <250 ms) index different facets of the comprehension process, including word form analysis, semantic access along with syntactic and semantic context integration, suggesting near-simultaneity (or short-delay seriality) in psycholinguistic information access.
The short delays are potentially accountable in terms of cortical conduction times
BRAIN-BASED MODELS OF CIRCUITS, THEIR ACTIVATIONS AND DELAYS
The areas shown on the brain diagram at the top and implemented in the model of the language cortex (MLC) at the bottom are:
Primary auditory cortex (A1), auditory belt (AB), auditory parabelt (PB), inferior prefrontal (PF), premotor (PM) and primary motor (M1) cortex. AB and PB together are sometimes called the ‘auditory language area’ or ‘Wernicke’s region’ and AB and PB the ‘motor language area’ or ‘Broca’s region’.
PROPOSED NEW MODEL - The ventral stream helps in
auditory “what” processing The dorsal auditory stream is
critical for auditory–motor integration
Early cortical stages of speech perception involve auditory-responsive fields in the superior temporal gyrus (STG) bilaterally.
This cortical processing system then diverges into two processing streams, a ventral stream, which is involved in mapping sound onto meaning, and
A dorsal stream, which is involved in mapping sound onto articulatory-based representations.
OF THE RELATION BETWEEN SYSTEMS SUPPORTING SUB-LEXICAL SEGMENTATION ABILITY AND AUDITORY COMPREHENSION ABILITY
Observed dissociations between these abilities arise when damage or functional imaging studies affect portions of the system after the divergence point, whereas we would predict some degree of correlation between these abilities if damage or functional imaging targets earlier shared components of the system.
G. Hickok, D. Poeppel / Cognition 92 (2004) 67–99 77
THE DUAL-STREAM MODEL OF THE FUNCTIONAL ANATOMY OF LANGUAGE
Hickok, Gregory & David Poeppel. 2007. The cortical organization of speech processing.Nature Reviews Neuroscience 8, 393–402.
FROM BRAIN MECHANISMS TO EXPLANATION
The cortex is not, a tabula rasa learning structure. It is equipped with the structure and microstructure of areas, connections between areas and even the microstructure of neurons and their biochemical properties.
Fiber tracts in human and non-human primates
WHAT IS GAINED FROM RECENT NEUROSCIENCE RESEARCH? At the semantic level, language is
‘woven into action’ Motor and sensory systems activation
demonstrates semantic categories along brain dimensions
In the vicinity of sensorimotor domains, may play a role in abstract semantic processing and in general meaning access.
RECENT RESEARCH CONT…
Phonetic distinctive features have their correlates in local cortical activation in the auditory and motor systems
Neuronal ensemble theory along with empirical neurophysiological evidence supports the existence of discrete cortical representations and mechanistic underpinnings for rules of grammar.
It takes about half a second to understand a word or sentence — counted from the point in time when the last word critical for sentence understanding is first unambiguously present in the input — might imply a substantial delay in the comprehension process
RECENT RESEARCH CONT…
Supportive of rapid, almost instantaneous understanding comes from recent neurophysiological studies suggesting latencies of <250 ms of the earliest brain correlates of semantic word and sentence understanding and syntactic parsing.
These neurophysiological results support rapid and parallel psycholinguistic models and argue against slow-serial or -cascaded theories assuming sequential steps from phonological to syntactic and semantic modules of hundreds of milliseconds
HEMISPHERIC FUNCTION FOR LANGUAGE
LANGUAGE AQUISITION
THE DEVELOPMENT OF LANGUAGE: A CRITICAL PERIOD IN HUMANS
A critical period for learning language is shown by the decline in language ability (fluency) of non-native speakers of English as a function of their age upon arrival in the United States
FIBER TRACTS IN ADULTS AND CHILDREN
.Differences in fractional anisotropy between adults and 7-year-old children: (a) more medial and (b) more lateral sagital slices of the left hemisphere.
J. Brauer, PhD thesis, Leipzig University, 2008.
SIGN LANGUAGE
Signing deficits in congenitally deaf individuals who had learned sign language from birth and later suffered lesions of the language areas in the left hemisphereLeft hemisphere damage produced signing problems in these patients analogous to the aphasias seen after comparable lesions in hearing, speaking patients
BRAIN ACTIVATION IN CONGENITALLY BLIND SUBJECTS DURING A VERBAL-MEMORY TASK
IN PEOPLE BORN BLIND, BRAIN REGIONS THAT USUALLY PROCESS VISION CAN TACKLE LANGUAGE.
FUTURE
A neuro-mechanistic explanations detailing why specific brain areas are necessary for, or light up and index, specific facets of language processing,
how neuronal ensembles and distributed areas become activated with precisely timed milli-second delays, and which precise neuronal wirings can potentially account for neurometabolic activation of specific cortical clusters in semantic understanding
THANK YOU