U9: LATERALIZATION, LANGUAGE and the SPLIT BRAIN Mar/22/12 1. Cerebral Lateralization of The brain is actually 2 separate structures except for the cerebral Function – Intro commisures connecting them Lateralization of function talks about the differences in function and is studied through split brain patients – who had their L and R hemispheres separated by commissurotomy Left Hemisphere Damage to Aphrasia is an inability to produce or understand language Aphasia and Apraxia o Broca found that there was a relationship between the inferior prefrontal cortex of the left hemisphere known as Broca’s area Apraxia (Liepmann) is always associated with left-hemisphere damage even though the symptoms involve both sides of the body. These patients can’t perform movements when asked to perform them but when performing them naturally and without think about it, they could. These examples provided evidence for cerebral dominance where one hemisphere (usually the left) is the dominant hemisphere in the control of all complex behavioral and cognitive processes. Right hemisphere is the minor hemisphere 2. Anatomical Asymmetry Even though the 2 sides of the brain seem to look the same, there are a number of anatomical asymmetries between the two cerebral hemispheres 1. The RH is larger and heavier than the LH but the LH is more densely packed 2. In the temporal lobe, the planum temporale (location of Wernickes area) is larger on the left while the primary auditory cortex or Heschel’s gyrus is larger on the right because there are 2 on the right side o Comparing neurons in wernicke’s area with the same part of the right hemisphere, the areas of both hemispheres are organized into regularly spaced columns of interconnected neurons and the columns are connected by axons o The L side had longer axons and are farther apart. 3. In the thalamus, the lateral posterior nucleus is larger on the left, while the medial geniculate nucleus is larger on the right 4. Tend to have more temporal-parietal area on the right side because the Sylvian fissure extends further back 5. In Broca’s area there is more surface area on the right and more buried area on the left 6. There is asymmetry in the distribution of some neurotransmitters 7. The RH extends further forward and the LH extends further back Asymmetry and Handedness 8. There is also evidence that the cortical microcircuitry differs between comparable areas of the left and right hemispheres, suggesting that cerebral asymmetry is expressed down to the level of how neurons communicate with one another - - While everyone shows some brain asymmetry, a higher proportion of left handers show no asymmetry or a reversal of the direction of anatomical asymmetry o Ex. 60% of Dextrals (right) with L hemisphere lesions and 2% with R hemisphere lesions were aphasic while sinestrals (Left) were 30% and 24% respectively. Left handers have language functions distributed across hemispheres or in right hemisphere. Corpus callosum is larger in left-handers, and that might also contribute to the greater interaction between the two hemispheres Recovery from aphasia in left handers are more rapid and complete in left handers because language functions aren’t as lateralized and less sensitive to damage in one particular brain area. Relationship between Speech laterality and Handedness 1. Nearly all (about 95%) right-handed subjects are left-hemisphere dominant for Asymmetry and Gender 2. 3. 1. Q: What is the CC? 2. 3. speech Most left-handed or ambidextrous subjects (about 70%) are also lefthemisphere dominant for speech Early left-hemisphere damage can cause the right hemisphere to become dominant for speech and the left hand to be preferred. There are anatomical asymmetries that differ between males and females: Planum temporale: Left is larger than right, more often seen in males than in females Sylvian Fissure: Men have a longer horizontal component in the LH than females/ no difference in RH The posterior portion of the CC is significantly larger in females - The brains of males are more lateralized than the brains of females because male victims of L hemisphere strokes had more deficits on WAIS and 3x more likely to suffer from aphasia. Gender differences in Behavior 1. Women have greater verbal ability than men (fluency, analogy, comprehension) 2. Men have greater visuo-spatial abilities (mental rotation, geometry, map reading) 3. Men have greater mathematical ability (very controversial and has been linked to advantages in music composition and chess) 4. Men show greater aggression Explanation of Sex Differences 1. Hormonal effects on brain function Hormones have an organizing effect during development and men and women have different levels of hormones can lead to differences in brain structure and functioning. During menstrual cycle as estrogen levels rise, this was correlated with poorer spatial ability but enhanced articulatory and motor abilities. 3. The Split Brain Cutting the Corpus Callosum Myers & Sperry 2. Genetic Sex-linkage Another theory proposes that a major factor in behavioural sex differences in spatial ability is genetic Recessive gene on X chromosome responsible for differences 3. Maturation Rate Girls speak sooner than boys so language areas of the brain are faster to mature in females 4. Environment Boys are expected to be more adventurous and explore their environment, an activity that improves spatial skills. 5. Preferred Cognitive Mode Men and women use different strategies when solving problems (women tend to solve them verbally) and this may cause observed sex differences in behaviour. Showed 2 functions of the Corpus Callosum: 1. Transfer learned information from one hemisphere to the other 2. When cut, each hemisphere can function independently - - Trained cats to perform visual discrimination of 2 panels – one with a circle on it and one with a square on it with the positions of these shapes varied from trial to trial The cats learned which symbol to press in order to get food. M&S thought that there are 2 ways for visual information to move from one eye to the opposite hemisphere: (1) corpus callosum (2) optic chiasm Wanted to study one hemisphere at a time so they cut both of these and covered one eye with a patch, restricting all incoming visual information to the hemisphere on the same side as the uncovered eye. Results - - Tests of Cerebral Lateralization Sodium Amytal Test All cats learned the discrimination with a patch on one eye even though cutting the optic chiasm created an area of blindness When the patch was transferred to the other eye for intact cats and control with either the OC or CC transected, they performed the task with close to 100% accuracy When transferred the patch to the experimental cats, the performance dropped for the hemisphere that hadn’t learned the task, but these cats learned the task as if they had never seen it before. 4. - - - Dichotic Listening Test - - Functional Brain Imaging Patients with Unilateral Damage Administered to patients prior to neurosurgery so that the surgeon knows the side of speech lateralization and can take care not to damage the language areas Sodium amytal is injected into one carotid artery and anaesthetizes the ipsilateral hemisphere and allows the abilities of the contralateral hemisphere to be assessed Inject on dominant speech hemisphere (left) muteness or speech errors Inject on nondominant speech hemisphere (right) no mutism and a few speech errors. A sequence of 3 pairs of digits is presented through earphones, the 2 digits of each pair are presented simultaneously, one digit to each ear When the subject is asked to report the six digits, there is a slight but consistent tendency to report more of the digits presented to the ear contralateral (opposite side) to the dominant language hemisphere. o Ex. Right-hemisphere specialization for language performed better with the L ear than the R. Although the sounds from each ear are projected to both hemispheres, contralateral connections are stronger and take precedence when 2 different sounds are simultaneously competing for access to the same cortical auditory centers. - PET or fMRI techniques have revealed that there is typically more activity in the left hemisphere than the right - If damage is localized to only one hemisphere, any deficits seen may be a result of lateralization of function o o Split Brain Patients Q: What does this mean: Commissurotomized patients have 2 independent streams of consciousness: however, unlike splitbrain laboratory animals, the two hemispheres of communisurized patients are not equal? 5. Differences in the L and R Hemispheres Comparing Functions of L and R Hemispheres Evidence of 2 Independent Streams of Consciousness Ex. Distinction has been made in the processing of whole objects versus the parts of those same objects . One part of the brain is involved in determine the overall shape of a triangle, another part processes the specific details (i.e. dots within triangle) R hemisphere seems responsible for global visual processing L hemisphere responsible for local visual processing - Have a cut in their corpus collosum to reduce the severity of convulsions 2 Hemispheres are disconnected from each other Typical experiment involves the subject seated in front of a computer monitor and an object or word would then appear briefly in either the right or left visual field of the monitor and the subject would report what they saw. All stimuli presented in the left visual field were transmitted to the right visual cortex and vice versa. Fine tactual and motor tasks were performed by each hand under a ledge. o Object in right visual field, the subject says comb because the L hemisphere is dominant for language production o Obj in left visual field, the subject can’t report anything because the R hemisphere is unable to speak and L hemisphere can’t see the stimulus. o The R hemisphere still processes the stimulus because if asked to identify that object by touch with their left hand, they are able to do the task. **Think of this as Ms. R Hemisphere who understands a few simple instructions but can’t speak, who receives sensory information from the left visual field and left hand and who controls the fine motor responses of the L hand. Mr. L hemisphere is verbally adept, receives sensory information from R visual field and R hand and controls motor responses of the R hand. Left Right - Better at language-related tasks Better at understanding spoken - Controls ipsilateral body words and grammar movements Better at spatial ability - Verbal memory Process emotional stimuli – - Episodic memory facial emotion recognition and - Acts as the Interpreter musical tasks :hypothetical neuronal Greater role in nonverbal mechanism that continuously memory assesses patterns of events and tries to make sense of them. Language, emotion, musical ability... are composed of dozens of different individual cognitive activities and there’s no reason to assume each of them is lateralized in a hemisphere. Thus, complex cognitive tasks – reading, judging space – should be broken down into their consitutent cognitive processes. - - - - When an object is presented to the left hemisphere either by touching with the right hand or viewing something in the right visual field, the split-brain patient could pick out the correct object with the right hand When object is presented to R hemisphere by touching with L hand or viewing something in the L visual field, the patient could pick out the correct object with the L hand but not the R hand o When an apple picture was flashed in the right visual field of a split-brain patient, the L hemisphere could do one of 2 things: 1. L could say it saw a picture of an apple 2. Patient can reach under the ledge with the right hand and feel out the apple. If the non-speaking R hemisphere were asked to indicate the object that had previously been presented to the left hemisphere, it can’t. This is communication between hemispheres via a nonneural route – in Cross Cuing Q: How does this work? Was shaking his head and frowning the cue to the left? - - If a split brain patient is presented with 2 objects at the same time, one on each visual field, they can reach into 2 different bags at the same time and pull out the two objects. - Occurs when the 2 hemispheres are presented with different information about the correct choice and then asked to reach out and pick up the correct object from a collection in full view Right hand pick out what left hemisphere saw but the right hemisphere seeing what it thinks is an error being made causes the left hand to grab the right and pull it to the other object. Learning 2 things at once Helping Hand Phenomenon Q: Why does the right hand act first? one test, a red or green light was flashed in the left visual field and the split-brain patient was then asked to name the color: red or green, most patients only get 50% right However one subject performed almost perfectly when he initially said the incorrect color using his left hemisphere, his head shook and the patient changed his guess to the correct color because the right hemisphere heard the left’s incorrect guess of the left and was signalled to the left hemisphere that it was wrong by shaking the person’s head. - - 6. Experimentation with Split Brain Patients Lateralization of Attention Luck, Hillyard, Mangun, Gazzaniga - - Used split-brain patients and used a visual search task in which the items were rectangles made from red and blue squares When the blue square was placed immediately above the red square, it was a distractor item and when the squares were reversed, the red square was a target item. Stimuli was presented unilaterally or bilaterally in sets of 2,4, and 8 The subjects had to decide whether the target item was in display and if so in which visual field Control – no significant differences between reaction time functions for the unilateral arrays as compared to the bilateral arrays Split brain patients – faster for bilateral arrays, each hemisphere was able to conduct an independent serial search. Guided Visual Task Split brain patient searched for a target black circle among distractor black squares, grey circles and grey squares, the reaction times presented in R visual field were faster than for L, suggesting that L hemisphere is specialized for processing stimuli with specific shared features. Faces Levy, Trevarthen, Sperry When disconnected, the 2 hemispheres may work differently depending on the type of orienting required. With reflexive/ exogenous orienting there may be independence between the left and right hemispheres and can work in parallel With endogenous (voluntary) orienting, the hemispheres may actually compete with each other Chimeric figures test: Visual images are presented to the subject that consist of faces and other patterns that have been split down the center and then recombined with other images Patients did not realize any Z Lens Q: What relationship does using a Z lens have to do with hearing speech in one ear? Information Shared between Split Hemipsheres Zaidel - - The Z lens limits visual input to one hemisphere of split-brain patients It is a contact lens that is opaque on one side and permits visual input to enter only one hemisphere irrespective of eye movement. - When the Z lens was used to assess the behavioral reactions of the right hemispheres of split brain patients to various emotion-charged images: photos of relatives, pets, themselves... the patient’s behaviors were emotionally appropriate, meaning that R hemispheres are capable of emotional expression. Interestingly, the emotional content was reflected in patients’ speech and nonverbal behavior. A patient’s left hemisphere was able to react appropriately to an image presented on the R hemisphere. Task difficulty also contributes to the reduced lateralization of function, because complex tasks require both hemispheres to be used. Damage in the angular gyrus (border b/w the left temporal and parietal lobes, just anterior to the occipital lobe) is the cause of alexia (lose ability to read) and agraphia (lose ability to write and spell) Left angular gyrus is responsible for comprehending language-related visual input causing patients to lose ability o read and write. Has not stood up well to the challenge of testing and hasn’t been supported 7. Wernicke-Geschwind Model of Language Lateralization - Historical Antecedents of the Wernicke-Geschwind Model discordance and when asked to pick out the face they had seen, they always chose the one in their left visual field (RH) When Archimbaldo paintings were shown, a split-brain patient’s left brain could only see the objects that made up the face while the right brain could see the face. Visual completion is when people with scotomas can fill out the missing visual field - - Broca’s Aphasia: selective lesions of Broca’s area produce a syndrome of aphasia whose symptoms are expressive or disjointed Wernicke’s aphasia: Receptive deficits - Poor comprehension of both written and spoken language and speech that is meaningless but retains superficial structure, rhythm and intonation. Damage to the pathway connecting Broca’s and Wernicke’s areas is the arcuate fasciculus would produce a third type of aphasia, which he called conduction aphasia which is difficulty repeating words they just heard. There are 7 components of the WernickeGeschwind model - How it works When you are having a conversation, the auditory signals triggered by the speech of the other person are received by your primary auditory cortex and conducted to Wernicke’s area for understanding Wernicke’s area generates the neural representation of the throught underlying the reply and it is transmitted to Broca’s area via the left arcuate fasciculus. Once in Broca’s area, the signal activates the right neurons of primary motor cortex and muscles of speaking If reading aloud, the signal received by your primary visual cortex is transmitted to your left angular gyrus, which translates the visual form of - - Effects of Cortical damage on Language Abilities - - - Effects of Electrical Stimulation of the Cortex on Language Abilities the word into its auditory code and transmits it to Wernicke’s area for comprehension Wernicke’s area then triggers the appropriate responses in arcuate fasciculus, Broca’s area and motor cortex to elicit the motor responses. Surgery that destroys all of Broca’s area but little surrounding tissue typically has no lasting effects on speech. Some speech problems were observed after the removal of Broca’s area but their temporal course suggested that they were products of post surgical edema/ swelling rather than the excision of the area itself. Small lesions to Broca’s area seldom produced lasting language deficits and lesions restricted to Wernicke didn’t always produce lasting language deficits. Lesions in parietal or temporal areas were just as likely to produce articulation problems Refuting the Wernicke- Geschwind Model Findings using CT and MRI on aphasia are: 1. No aphasic patients have damage restricted to Broca’s area or Wernicke’s area and damage to the surrounding W-G areas have little lasting effect on the use of language. 2. Aphasic patietns almost always have significant damage to subcortical white matter 3. Large anterior lesions are more likely to produce expressive symptoms whereas large posterior lesions are more likely to produce receptive symptoms. Aphasia always involves both expressive and receptive symptoms. 4. Global aphasia – severe disruption of all language related abilities is usually related to massive lesions of anterior cortex, posterior cortex and underlying white matter. 5. Aphasic patients sometimes have brain damage that doesn’t encroach on the W-G areas - - - Electrical stimulation was done by assessing the responses of conscious patients who were under local anesthetic to stimulation applied to various points on the cortical surface Electrical stimulation is more localized than a brain lesion and found that sites at which stimulation blocked or disrupted speech in conscious neurosurgical patients were scattered throughout a large expanse of frontal, temporal and parietal cortex instead of being restricted to W-G areas. Right hemisphere stimulation almost never disrupted speech. Findings - Areas of cortex at which stimulation could disrupt language extended far beyond the boundaries of the W=G language areas Each of the language tests were disrupted by stimulation at widely scattered sites There were major differences among the subjects in the organization of language abilities. 8. Evolutionary Perspective of Cerebral Lateralization and Language Analytic-Synthetic Theory - - There are 2 basic modes of thinking: 1. Analytic mode – L hemisphere operates in this logical, analytic manner 2. Synthetic mode – R hemisphere is a synthesizer, concerned with overall stimulus configuration. Problem with this theory is that it’s vague, it is not possible to specify the degree to which any task requires either analytic or synthetic processing. Motor Theory - Left Hemisphere isn’t specialized for control of speech specifically but - for the control of fine movements The problem with this theory is that it doesn’t suggest why motor function became lateralized in the first place. Linguistic Theory When did cerebral Lateralization evolve? - Primary role of the left hemisphere is language because when a study of deaf people who suffer unilateral brain damage, the L hemisphere damage can disrupt the use of sign language but not pantomime gestures. - Lateralization of function may have preceded human evolution. Right handed may have evolved from a preference for the right side of the body for feeding There are 2 main advantages of lateralization: 1. May be more efficient for the neurons performing a particular function to be concentrated in one hemisphere o Ex. It’s better to have one highly skilled hand rather than 2 moderately skilled hands 2. Two different kinds of cognitive processes may be more readily performed simultaneously if they’re lateralized to different hemispheres. o Ex. Motor theory suggests that language was lateralized to the L hemisphere because fine motor control was already lateralized there. - Evolution of Human Language - - Motor Theory of Speech Perception - - Gestural Language Pollick and deWaal 9. Cognitive neuroscience Approach to Language Premise 1 - - - - At 10 months, human infants can distinguish the sounds of all human languages but by 30 months, they can readily discriminate only those sounds that compose the languages to which they’ve been exposed. Even the most vocal nonhumans can produce a relatively few calls but they’re able to interpret a wide variety of other sounds in their environments – these nonhumans are limited by their inability to exert fine motor control over their voices, a talent only humans have. Proposes that the perception of speech depends on the words activating the same neural circuits in the motor system that would be activated if the listener said the words. Support comes from the discovery that just thinking about performing a particular action activates the same areas of the brain as performing the action – mirror neurons are activated Comparing the gestures and the vocalizations of chimpanzees, a highly nuanced vocabulary of hand gestures were used in many situations and in various combinations. The cognitive neuroscience approach to language is guided by 3 premises: Language can be broken down into constituent cognitive processes that are much simpler than the cognitive activities that Geschwind (and others) tried to localize to a single part of the brain These constitutent processes include phonological analysis (analysis of language sounds), grammatical analysis (analysis of language structure) and semantic analysis (of meaning) Premise 2 - Areas of the brain involved in language are not exclusively involved with that function. Areas involved with reading might also be involved with short term memory or pattern recognition. Premise 3 - Brain areas involved with language are small and widely distributed The Brain during Silent Reading Bavelier - Brain activity were recorded during the reading of sentences with control periods where the participants were presented with strings of consonants interposed between periods of silent reading. Used sensitive fMRI to study the activity of the brain and found: - 1. 2. PET Study of Naming Damasio - 10. Cognitive Neuroscience of Dyslexia - Developmental Dyslexia: Causes and Neural Mechanisms - - - Only small areas are activated at a given time in a single subject These areas vary from subject to subject and also within a single subject from trial to trial 3. The activity was spread over very large areas of the brain 4. The active areas were more often in the left hemisphere than the right 5. That the activity spread far beyond those areas predicted by Wernicke-Geschwind model. Also found right hemisphere damage rarely disrupts language activity PET activity was recorded from the left temporal lobes of healthy volunteers while they named images (famous faces, animals, tools) presented on a screen Naming objects activated areas of the left temporal lobe outside the classic Wernicke’s language area. Dyslexia is a pathological difficulty in reading, and there are 2 types: 1. Developmental Dyslexias: Which become apparent when a child is learning to read 2. Acquired dyslexias: caused by brain damage in individuals who were already capable of reading. There is a large genetic component to this Dyslexia is hard to identify because there are so many changes in the brains of individuals that no single kind of brain pathology has been found. Reading or absence of reading can also induce major changes in the brain. Researchers have tried to attribute developmental dyslexia to attentional and other sensorimotor deficits caused by damage to neural circuits linked to magnocellular layers of the lateral geniculate nuclei (receive information from rods for perception of movement) However even when visual, auditory, or motor deficits are present in dyslexic patients, they don’t account for all aspects of the disorder. It’s now widely agreed upon dyslexia comes from a disturbance of phonological processing (representation and comprehension of speech sounds) and not a disturbance of sensorimotor functioning. Ramus Theory on Developmental dyslexia Ramus argues that the first stage in the development of dyslexia is the occurrence of developmental errors in auditory areas around the lateral fissure A gene associated with dyslexia controls neuronal migration However, this theory doesn’t explain why dyslexia is often associated with cerebellar damage. Developmental Dyslexia and Culture - - Cognitive Neuroscience of Deep and Surface Dyslexia 1. - Dyslexia could be a cultural disease because twice as many English speakers as Italians are diagnosed as dyslexic English has more phonemes spelled in 1120 different ways, whereas Italian is less than that Italian readers displayed more activity in the left superior temporal lobe whereas English readers displayed more activity in the left inferior temporal and frontal lobes. However both language readers display the same pattern of abnormal PET activity – less than normal reading-related activity in the posterior regions of the temporal lobe. Reading aloud can be accomplished in 2 different ways: Lexical Procedure Based on specific stored information that has been acquired about written words Reader looks at word and recognizes it and says it. 2. - - - - Phonetic Procedure The reader looks at the word, recognizes the letters, sounds them out and says the word This is used for unfamiliar words. Knowing how reading aloud has proven useful in understanding the symptoms of 2 kinds of dyslexia resulting from brain damage: 1. Surface dyslexia Patients have lost their ability to pronounce words based on their specific memories of the words but can still apply rules of pronunciation in their reading. o Ex. Can pronounce words and nonwords with pronunciation that is consistent with common rules (e.g. fish, river, glass, spleemer and twipple) o Can’t pronounce words that don’t apply to the rules (e.g. have is pronounced like cave) 2. Deep Dyslexia Patients have lost their ability to apply rules of pronunciation in their reading but can pronounce familiar words based on their specific memories of them Completely incapable of pronouncing nonwords and have difficulty pronouncing uncommon words o Ex. Respond by responding to the overall look, meaning or derivation of the word o Quill might be said for quail based on overall look, hen for chicken (meaning) or wise for wisdom (derivation of the word) CH 16: CASES 1. The Split Brain The Case of Peter, Split Brain Patient Tormented by Conflict - 2. Evolutionary Perspective The Case of W.L The Man who Experienced Aphasia for Sign Language - 3. Dyslexia The case of N.I the woman who Read with her R Hemisphere - - - - In most split brain patients, the L hemisphere seems to control most everyday activities but when in a few patient cases, the R hemisphere is more active, there can be serious conflicts Peter received a commissurotomy and was left-hemisphere dominant for language He couldn’t respond with the L side of his body to verbal input because his L side would not control the left side via ipsilateral fibers Peter’s R hemisphere would cause his left side to do things he didn’t want to do. His L hemisphere would swear at the L hand and make the Right hand force the L hand to do what it wanted. W.L was a deaf man who got a stroke on the left frontotemporoparietal side. Compared to a recording of him before the stroke, W.L suffered a loss in his ability to use and understand sign language. He could produce and understand pantomime gestures suggested that his sign-language aphasia wasn’t the result of motor or sensory deficits nor was it resulting from cognitive deficits. N.I experienced periods of aphasia at 13 and suffered from convulsions. In an attempt to relieve these symptoms, a left hemispherectomy was performed – her left hemisphere was totally removed and her seizures stopped She recognizes letters but is incapable of translating them into sounds and she can read concrete familiar words, she can’t pronounce even simple nonsense words and her reading errors indicate she’s reading on the basis of the meaning and appearance of words. Experienced the same symptoms as patients with deep dyslexics.
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