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Hemispheric Specialization

Colwyn Trevarthen

10.1002/cphy.cp010325

Source: Supplement 3: Handbook of Physiology, The Nervous System, Sensory Processes

Originally published: 1984

Published online: January 2011

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Hemispheric Differences in Normal Subjects
1.1 Perceptual Discrimination and Reaction Time
1.2 Handedness
2 Electrical Signs
2.1 Potentials Evoked by Stimuli
2.2 Potentials Related to Execution of Movements
2.3 Asymmetry in the Electroencephalogram
3 Anatomical Correlates
3.1 Left Hemisphere Sites and Speech Perception
3.2 Right Hemisphere Enlargements: Variations With Handedness
3.3 Anatomical Correlates of Left Hemisphere Deficits
4 Hemispherectomy and Cerebral Hemiatrophy
4.1 Hemispherectomy for Tumor
4.2 Hemispherectomy for Infantile Hemiplegia
5 Transitory Inactivation of one Hemisphere
5.1 Unilateral Carotid Injection of Amytal
5.2 Unilateral Electroconvulsion
6 Hemisphere Functions Dissociated by Commissurotomy
6.1 Cognitive and Perceptual Processes Within Both Hemispheres
6.2 Hemispheric Specialization for Language
6.3 Superiority of Right Hemisphere for Spatial Constructive Skills
6.4 Spontaneous Choice of Hemispheric Response Mode: Tests with Stimulus Chimeras
7 Effects of Left Cerebral Lesions
7.1 Classic Theories of the Hemispheres
7.2 Current Ideas on Left Hemisphere Specialization for Language
7.3 Disorders of Logical Inference
7.4 Left Hemisphere Systems for Voluntary Motor Coordination and for Perceptual Categorization
8 Disorders from Lesions of Right Hemisphere
8.1 Visuoconstructive Disorders
8.2 Unilateral Neglect of Extracorporeal Space or of Parts of Body
8.3 Perception of Orientation and Form of Objects
8.4 Color Perception
8.5 Loss of Face Recognition: Prosopagnosia
9 Development of Lateral Asymmetry of Brain Function
10 Sex Differences in Laterality
11 Asymmetries in Emotional Expression
12 Conclusions
Figure 1. Figure 1.

Projections to hemispheres of visual, auditory, and haptic sensory and motor functions divide left (L) and right (R) halves of extracorporeal space with inversion of left and right. Cognitive processes are asymmetrically represented in cortical areas remote from primary sensory fields of hand (H), audition (A) and vision (V) in both hemispheres.
Figure 2. Figure 2.

Left: Simple scheme to explain how hand reactions (LH, left hand; RH, right hand) are faster to stimuli in the same side (LVF, left visual field; RVF = right visual field). Crossed combinations involve a longer transcallosal route (CC). Right: More complex factors affecting reaction times. Contralateral hand control (C) is more rapid than ipsilateral (I) for one hemisphere. Temporary time differences in sensory‐motor processes result from orientational sets coupling brain stem to cortex (e.g., large black arrow activating right hemisphere), or from task‐specific cognitive sets that may be asymmetric (e.g., dotted arrows indicating frontal and temporoparietal processes favoring left hemisphere in, for example, a verbal task).
Figure 3. Figure 3.

Manual reaction times as a function of visual field of presentation (LVF, left visual field; RVF, right visual field). A: stimuli seen as easily integrated wholes. Faster identity matches (“SAME”) produce stronger left‐field superiorities than slower “DIFFERENT” discriminations. B: in more difficult task requiring separate feature analysis, difference discriminations typically are faster, but both responses induce right‐field superiorities. C: when stimuli do not comprise readily integrated wholes, or when memory load is high, there is field‐by‐judgment interaction.

Courtesy of J. Bradshaw
Figure 4. Figure 4.

Upper: increase of late positive component of right parietal event‐related potential (ERP) during active touch by right hand in a right‐handed subject. At arrow, right index finger is dropped mechanically onto a circular ridge on a perspex rod. By palpation subject identifies orientation of a groove in the ridge (thick trace). Runs with smooth perspex surface show smaller N150 in left cortex (C) and smaller late positive potential (P400) in right cortex (E). A: vertical electrooculogram showing absence of eye movement artifacts. B: calibration. D, F: averages of EEG samples to check that nonresponse EEG background does not contribute to ERP wave forms. Lower: silent reading of seven‐word sentences presented one word at a time elicited ERPs in temporoparietal cortex with late positive component that was significantly larger on left side. Potentials shown are averages of ERPs to first six words

Upper: from Desmedt and Robertson 156, with permission of S. Karger AG, Basel; lower: from Kutas and Hillyard 355
Figure 5. Figure 5.

Upper surfaces of left and right temporal lobes (temporal planum) showing asymmetry posterior to Heschl's gyrus (HG). A: drawing published by von Economo and Horn in 1930. B: confirmation of asymmetry by Geschwind and Levitsky (brains sectioned along plane T in Fig. 9.) C: cytoarchitectonic areas of human auditory cortex in one brain. Black (KA), primary auditory receptive area, is equal on the two sides, but stippled (Tpt), temporoparietal cortex related to language function, shows clear asymmetry

A: from von Economo and Horn 624; B: from Galaburda et al. 202, copyright 1978 by the American Association for the Advancement of Science, and from Geschwind and Levitsky 225; C: adapted from a figure of Geschwind and Sanides in Galaburda, Geschwind, et al. 202
Figure 6. Figure 6.

Left (upper row) and right (lower row) hemispheres of orangutan (A), human fetus of 5 mo gestation (B), adult human (C), and endocranial casts of Neanderthal La Chapelle‐aux‐Saints skull (D). All the brains show Sylvian fissures (arrows) shorter and more upturned in right hemisphere

From LeMay 366 and from LeMay and Geschwind 369, with permission of S. Karger AG, Basel
Figure 7. Figure 7.

Top left: areas of left hemisphere most commonly damaged in Broca's nonfluent or motor aphasia (B), conduction aphasia (C), and Wernicke's fluent or receptive aphasia (W). Top right: right hemisphere showing parietooccipital area common to lesions in cases of visuoconstructive disorder. Center: neurohistological areas of cortex according to Brodmann. Primary receptive areas in black (areas 3, 17, 41, and olfactory lobe). Areas apparently unique to humans and absent in great apes are cross‐hatched (areas 37, 39, 40, 44–46). Bottom left: asymmetric areas in “association” cortex. Note clear asymmetry in termination of Sylvian sulcus with respect to line following upper surface of temporal lobe (T). Language areas of left hemisphere: Broca's (B), Wernicke's (W). Bottom right: right hemisphere. S, body image, visuospatial and visuoconstructive functions; F, major component for facial recognition

Top left: adapted from Kertesz et al. 316 and Mazzocchi and Vignolo 693; top right: adapted from Hécaen et al. 270; center: adapted from Brodmann 70
Figure 8. Figure 8.

Three sectional pictures of brains traced from images obtained by x‐ray computerized axial tomography (CT scan) showing asymmetries in frontal pole (b wider than a) and occipital pole (c wider than d). Ventricles also show consistent asymmetries; left horn projects farther posteriorly

CT scans courtesy of M. LeMay. From LeMay 366
Figure 9. Figure 9.

Schematic representation of written language systems available to left and right hemispheres, summarizing tests with three children between ages of 10 and 15 yr after complete removal of one hemisphere in infancy. MW, right‐hemispherectomy patient, male. SM (male) and CA (female), patients with left hemisphere removed

From Dennis et al. 135
Figure 10. Figure 10.

Development of myelin in different brain tracts. Late‐maturing components (LMC) connect with the slowest‐maturing areas of cortex; these include territories in which psychological processes are asymmetrically represented in most adults. Major developments in language skills occur as these late developments of the brain occur

Myelinization data from Lecours 362
Figure 11. Figure 11.

Lateralized functions revealed by psychological tests of commissurotomy patients

Adapted from Sperry 570
Figure 12. Figure 12.

Methods for testing commissurotomy patients. Upper: standard tests of perception and learning in disconnected hemispheres. Visual stimuli back‐projected tachistoscopically while subject (at left) fixates a central point on screen. Auditory stimuli presented in headphones with dichotic conflict. Objects felt in hand or by foot, but out of sight. Lower: Zaidel's contact lens technique. Cap on contact lens in focal plane of reduced visual image occludes one‐half of visual field of one eye. This permits subject to freely scan test pictures while receiving visual input in one hemisphere only. Standard intelligence tests may be administered to one hemisphere at a time

Upper drawing: courtesy of R. W. Sperry; lower, courtesy of Zaidel 673
Figure 13. Figure 13.

Tests of visuospatial function and haptic stereognosis for commissurotomy patients. Upper: Nebes compared ability of left and right hands to recognize size of a circle from an arc fragment. A, somatosensory to visual; B, visual to somatosensory; C, somatosensory to somatosensory. Lower: in Levy's tests, shaped wooden blocks, such as the truncated pyramid with grooves shown on left, were felt out of sight in the hand; then a matching shape was chosen from three diagrams such as A, B, and C that represent objects of similar form, unfolded

Upper: from Nebes 694; in: Hemisphere Disconnections and Cerebral Function, edited by M. Kinsbourne and W. L. Smith, 1975; courtesy of Charles C Thomas, Publisher, Springfield, IL. Lower: courtesy of J. Levy
Figure 14. Figure 14.

Bilateral chimeric stimuli (bottom left) presented tachistoscopically with subject fixating a central point (cf. Fig. 15). This gave rise to separate perceptions in the two hemispheres (right). Naming led subject to describe stimulus perceived in left hemisphere. Matching by pointing with either hand in array shown top left led to choice of image perceived in right hemisphere

From Levy, Trevarthen, and Sperry 382. Reproduced by permission of Oxford University Press
Figure 15. Figure 15.

Tachistoscope (modified from Gerbrands) used to present chimeric stimuli. A: with chimeras made from drawings of familiar objects, responses were again segregated between the hemispheres as described in Fig. 14. B: with same stimuli as in A, a test of matching by phonological comparison—i.e., by rhyming in the head without any form of overt speech and then pointing to an appropriate picture—evoked activity strongly lateralized to left hemisphere (e.g., “bee” rhymes with “key”)

From Levy and Trevarthen 380
Figure 16. Figure 16.

Scheme to represent unequal representation of body scheme in left and right hemispheres. Neglect of left side is produced by right parietal lesions; left parietal lesions are not associated with contralateral neglect.
Figure 17. Figure 17.

Developmental compensatory changes in lateralization of word perception processes due to congenital deafness. Left: in tachistoscopic tests, normal adults (solid lines) perceive four‐letter words, presented unilaterally (U) or bilaterally (B), more accurately in right visual field (RVF). Deaf adults (dashed lines) show no such asymmetry. Right: event‐related potentials of normal subjects to words presented in the right visual field show large negative wave (N 410) in left temporal cortex. With deaf subjects, right‐field stimuli show smaller negative wave (N 330) in right hemisphere, possibly a result of greater use of visual processes in language by the deaf.

Courtesy of H. J. Neville


Figure 1.

Projections to hemispheres of visual, auditory, and haptic sensory and motor functions divide left (L) and right (R) halves of extracorporeal space with inversion of left and right. Cognitive processes are asymmetrically represented in cortical areas remote from primary sensory fields of hand (H), audition (A) and vision (V) in both hemispheres.



Figure 2.

Left: Simple scheme to explain how hand reactions (LH, left hand; RH, right hand) are faster to stimuli in the same side (LVF, left visual field; RVF = right visual field). Crossed combinations involve a longer transcallosal route (CC). Right: More complex factors affecting reaction times. Contralateral hand control (C) is more rapid than ipsilateral (I) for one hemisphere. Temporary time differences in sensory‐motor processes result from orientational sets coupling brain stem to cortex (e.g., large black arrow activating right hemisphere), or from task‐specific cognitive sets that may be asymmetric (e.g., dotted arrows indicating frontal and temporoparietal processes favoring left hemisphere in, for example, a verbal task).



Figure 3.

Manual reaction times as a function of visual field of presentation (LVF, left visual field; RVF, right visual field). A: stimuli seen as easily integrated wholes. Faster identity matches (“SAME”) produce stronger left‐field superiorities than slower “DIFFERENT” discriminations. B: in more difficult task requiring separate feature analysis, difference discriminations typically are faster, but both responses induce right‐field superiorities. C: when stimuli do not comprise readily integrated wholes, or when memory load is high, there is field‐by‐judgment interaction.

Courtesy of J. Bradshaw


Figure 4.

Upper: increase of late positive component of right parietal event‐related potential (ERP) during active touch by right hand in a right‐handed subject. At arrow, right index finger is dropped mechanically onto a circular ridge on a perspex rod. By palpation subject identifies orientation of a groove in the ridge (thick trace). Runs with smooth perspex surface show smaller N150 in left cortex (C) and smaller late positive potential (P400) in right cortex (E). A: vertical electrooculogram showing absence of eye movement artifacts. B: calibration. D, F: averages of EEG samples to check that nonresponse EEG background does not contribute to ERP wave forms. Lower: silent reading of seven‐word sentences presented one word at a time elicited ERPs in temporoparietal cortex with late positive component that was significantly larger on left side. Potentials shown are averages of ERPs to first six words

Upper: from Desmedt and Robertson 156, with permission of S. Karger AG, Basel; lower: from Kutas and Hillyard 355


Figure 5.

Upper surfaces of left and right temporal lobes (temporal planum) showing asymmetry posterior to Heschl's gyrus (HG). A: drawing published by von Economo and Horn in 1930. B: confirmation of asymmetry by Geschwind and Levitsky (brains sectioned along plane T in Fig. 9.) C: cytoarchitectonic areas of human auditory cortex in one brain. Black (KA), primary auditory receptive area, is equal on the two sides, but stippled (Tpt), temporoparietal cortex related to language function, shows clear asymmetry

A: from von Economo and Horn 624; B: from Galaburda et al. 202, copyright 1978 by the American Association for the Advancement of Science, and from Geschwind and Levitsky 225; C: adapted from a figure of Geschwind and Sanides in Galaburda, Geschwind, et al. 202


Figure 6.

Left (upper row) and right (lower row) hemispheres of orangutan (A), human fetus of 5 mo gestation (B), adult human (C), and endocranial casts of Neanderthal La Chapelle‐aux‐Saints skull (D). All the brains show Sylvian fissures (arrows) shorter and more upturned in right hemisphere

From LeMay 366 and from LeMay and Geschwind 369, with permission of S. Karger AG, Basel


Figure 7.

Top left: areas of left hemisphere most commonly damaged in Broca's nonfluent or motor aphasia (B), conduction aphasia (C), and Wernicke's fluent or receptive aphasia (W). Top right: right hemisphere showing parietooccipital area common to lesions in cases of visuoconstructive disorder. Center: neurohistological areas of cortex according to Brodmann. Primary receptive areas in black (areas 3, 17, 41, and olfactory lobe). Areas apparently unique to humans and absent in great apes are cross‐hatched (areas 37, 39, 40, 44–46). Bottom left: asymmetric areas in “association” cortex. Note clear asymmetry in termination of Sylvian sulcus with respect to line following upper surface of temporal lobe (T). Language areas of left hemisphere: Broca's (B), Wernicke's (W). Bottom right: right hemisphere. S, body image, visuospatial and visuoconstructive functions; F, major component for facial recognition

Top left: adapted from Kertesz et al. 316 and Mazzocchi and Vignolo 693; top right: adapted from Hécaen et al. 270; center: adapted from Brodmann 70


Figure 8.

Three sectional pictures of brains traced from images obtained by x‐ray computerized axial tomography (CT scan) showing asymmetries in frontal pole (b wider than a) and occipital pole (c wider than d). Ventricles also show consistent asymmetries; left horn projects farther posteriorly

CT scans courtesy of M. LeMay. From LeMay 366


Figure 9.

Schematic representation of written language systems available to left and right hemispheres, summarizing tests with three children between ages of 10 and 15 yr after complete removal of one hemisphere in infancy. MW, right‐hemispherectomy patient, male. SM (male) and CA (female), patients with left hemisphere removed

From Dennis et al. 135


Figure 10.

Development of myelin in different brain tracts. Late‐maturing components (LMC) connect with the slowest‐maturing areas of cortex; these include territories in which psychological processes are asymmetrically represented in most adults. Major developments in language skills occur as these late developments of the brain occur

Myelinization data from Lecours 362


Figure 11.

Lateralized functions revealed by psychological tests of commissurotomy patients

Adapted from Sperry 570


Figure 12.

Methods for testing commissurotomy patients. Upper: standard tests of perception and learning in disconnected hemispheres. Visual stimuli back‐projected tachistoscopically while subject (at left) fixates a central point on screen. Auditory stimuli presented in headphones with dichotic conflict. Objects felt in hand or by foot, but out of sight. Lower: Zaidel's contact lens technique. Cap on contact lens in focal plane of reduced visual image occludes one‐half of visual field of one eye. This permits subject to freely scan test pictures while receiving visual input in one hemisphere only. Standard intelligence tests may be administered to one hemisphere at a time

Upper drawing: courtesy of R. W. Sperry; lower, courtesy of Zaidel 673


Figure 13.

Tests of visuospatial function and haptic stereognosis for commissurotomy patients. Upper: Nebes compared ability of left and right hands to recognize size of a circle from an arc fragment. A, somatosensory to visual; B, visual to somatosensory; C, somatosensory to somatosensory. Lower: in Levy's tests, shaped wooden blocks, such as the truncated pyramid with grooves shown on left, were felt out of sight in the hand; then a matching shape was chosen from three diagrams such as A, B, and C that represent objects of similar form, unfolded

Upper: from Nebes 694; in: Hemisphere Disconnections and Cerebral Function, edited by M. Kinsbourne and W. L. Smith, 1975; courtesy of Charles C Thomas, Publisher, Springfield, IL. Lower: courtesy of J. Levy


Figure 14.

Bilateral chimeric stimuli (bottom left) presented tachistoscopically with subject fixating a central point (cf. Fig. 15). This gave rise to separate perceptions in the two hemispheres (right). Naming led subject to describe stimulus perceived in left hemisphere. Matching by pointing with either hand in array shown top left led to choice of image perceived in right hemisphere

From Levy, Trevarthen, and Sperry 382. Reproduced by permission of Oxford University Press


Figure 15.

Tachistoscope (modified from Gerbrands) used to present chimeric stimuli. A: with chimeras made from drawings of familiar objects, responses were again segregated between the hemispheres as described in Fig. 14. B: with same stimuli as in A, a test of matching by phonological comparison—i.e., by rhyming in the head without any form of overt speech and then pointing to an appropriate picture—evoked activity strongly lateralized to left hemisphere (e.g., “bee” rhymes with “key”)

From Levy and Trevarthen 380


Figure 16.

Scheme to represent unequal representation of body scheme in left and right hemispheres. Neglect of left side is produced by right parietal lesions; left parietal lesions are not associated with contralateral neglect.



Figure 17.

Developmental compensatory changes in lateralization of word perception processes due to congenital deafness. Left: in tachistoscopic tests, normal adults (solid lines) perceive four‐letter words, presented unilaterally (U) or bilaterally (B), more accurately in right visual field (RVF). Deaf adults (dashed lines) show no such asymmetry. Right: event‐related potentials of normal subjects to words presented in the right visual field show large negative wave (N 410) in left temporal cortex. With deaf subjects, right‐field stimuli show smaller negative wave (N 330) in right hemisphere, possibly a result of greater use of visual processes in language by the deaf.

Courtesy of H. J. Neville
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Article Title: Hemispheric Specialization
 

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Colwyn Trevarthen. Hemispheric Specialization. Compr Physiol 2011, Supplement 3: Handbook of Physiology, The Nervous System, Sensory Processes: 1129-1190. First published in print 1984. doi: 10.1002/cphy.cp010325