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Impairment of binocular vision in the adult cat induces plastic changes in the callosal cortical map.

In the primary visual cortex of normally reared adult cat, neurons activated through the corpus callosum are almost entirely located at the 17/18 border. They display small receptive fields distributed along the central vertical meridian of the visual field and are orientation selective. Here we demonstrate that a few weeks of monocular deprivation or unilateral convergent strabismus produced in adulthood does not modify the cortical distribution of these neurons, but leads to an increase of their receptive field size mainly toward the ipsilateral hemifield and to a loss of their orientation selectivity. We conclude that manipulation of binocular vision in the adult modifies neither the location of the primary callosal cortical map nor its retinotopy. In contrast, it induces functional plastic changes in this map which lead to a significant widening of the area of visual space signalled through the corpus callosum. These plastic changes are interpreted as the result of the strengthening of normally hidden subthreshold synaptic inputs.

Animals↗

Nerve growth factor (NGF) prevents the shift in ocular dominance distribution of visual cortical neurons in monocularly deprived rats.

The hypothesis that NGF could play a role in the plasticity of the developing mammalian visual cortex was tested in monocularly deprived (MD) rats. In particular, we have asked whether an exogenous supply of NGF could prevent the changes in ocular dominance distribution induced by monocular deprivation. Hooded rats were monocularly deprived for 1 month, starting at postnatal day 14 (P14), immediately before eye opening, by means of eyelid suture. In eight rats, only monocular deprivation was performed; in eight rats, monocular deprivation was combined with intraventricular injections of beta-NGF, and in three rats, with intraventricular injections of cytochrome C. Injections (2 microliters) were given every other day for a period of 1 month. Single neuron activity was recorded in the primary visual cortex of MD rats, MD rats treated with NGF, and MD rats treated with cytochrome C at the end of the deprivation period, and in normal rats of the same age. We found that monocular deprivation caused a striking change in the ocular dominance distribution of untreated MD rats, reducing binocular cells by a factor of two and increasing by a factor of eight the number of cells dominated by the nondeprived eye. In MD NGF-treated rats, the ocular dominance distribution was indistinguishable from the normal. Cytochrome C treatment was completely ineffective in preventing the ocular dominance shift induced by monocular deprivation. To test whether NGF affected cortical physiology or interfered with transmission of visual information, we evaluated in NGF-treated rats the spontaneous discharge and the orientation selectivity. We found these functional properties to be in the normal range. We conclude that NGF is effective in preventing the effects of monocular deprivation in the rat visual cortex and suggest that NGF is a crucial factor in the competitive processes leading to the stabilization of functional geniculocortical connections during the critical period.

Animals↗

Intrinsic circuit organization of the major layers and sublayers of the dorsolateral prefrontal cortex in the rhesus monkey.

Intrinsic connections are likely to play important roles in cognitive information processing in the prefrontal association cortex. To gain insight into the organization of these circuits, intracortical connections of major laminar and sublaminar divisions were retrogradely labeled in Walker's area 9 and 46 in rhesus monkeys by using cholera toxin (B-subunit) conjugated to colloidal gold. Microinjections placed within particular cortical laminae produced unique patterns of retrograde labeling. Injections in layers II/III yielded labeling which was laterally widespread (2-7 mm) in supragranular layers, and more narrowly focused, i.e., conforming to a column, in layers IV-VI. In contrast, local circuits associated with layers IV and Vb displayed a regular, cylindrical organization, whereas intrinsic connections of layer Va were laterally extensive (3-5 mm) in layers III and Va. Finally, injections in layer VI gave rise to a narrow column of cell labeling traversing all layers, augmented by laterally extensive labeling (approximately 7 mm) in layer VI. The intrinsic connections of the prefrontal cortex were arrayed within mediolaterally elongated stripes which were often distributed asymmetrically in either the medial or lateral direction. In addition, labeled cells within these mediolaterally oriented fields were frequently grouped within discrete clusters or narrow bands. The intrinsic connections identified in this study differ from the local circuits of corresponding layers reported for primary visual cortex; the unique intrinsic wiring diagram of the prefrontal cortex may be related to its specialized cognitive and mnemonic functions.

Animals↗

Is humanlike cytoarchitectural asymmetry present in another species with complex social vocalization? A stereologic analysis of mustached bat auditory cortex.

Considerable evidence suggests that left hemispheric lateralization for language comprehension in humans is associated with cortical microstructural asymmetries. However, despite the fact that left hemispheric dominance for the analysis of species-specific social vocalizations has been reported in several other species, little is known concerning microstructural asymmetries in auditory cortex of nonhumans. To test whether such neuroanatomical lateralization characterizes another species with complex social vocalizations, we performed stereologic analyses of Nissl-stained cells in layer III of area DSCF in mustached bats (Pteronotus parnellii). Area DSCF was selected because it contains neurons which are sensitive to several temporal features of conspecific vocalizations. Primary visual cortex (V1) was also studied as a comparative reference. We measured neuron densities, glial densities, and neuronal volumes in both hemispheres of 10 adult male bats. Results indicate that these variables are not significantly lateralized in area DSCF or V1. Additionally, magnopyramidal cells (i.e., the largest 10% of neurons from both hemispheres) were not asymmetric in their frequency of distribution at the population level. Although several individual bats had asymmetric neuron distributions, consistent hemispheric bias was not evident. Absence of population-level microstructural asymmetry in area DSCF of mustached bats suggests alternative evolutionary scenarios including: (1) microstructural lateralization of auditory cortical circuitry may be a unique adaptation for human language, and (2) the specialized biosonar function of mustached bat auditory cortex may require symmetrical cytoarchitectural structure. Resolution of these alternatives will require further data on the microstructure of auditory cortex in species with lateralized perception of acoustic social communication.

Adaptation, Physiological↗

Precise placement of multiple electrodes into functionally predefined cortical locations.

Current research on topics such as effective connectivity, neuronal coding strategy or signal propagation in the central nervous system requires simultaneous recordings from multiple sites within functionally grouped but topologically distributed neuronal clusters. We have addressed this issue by characterization of the cortical functional architecture using optical imaging of intrinsic signals (OI) and subsequent placement of multiple, individually adjustable electrodes into pre-selected domains. In order to achieve maximum precision and flexibility for the positioning of electrodes, a plastic cylinder containing channels of an extremely high aspect ratio (density >20 channels/mm(2)) was fixed above the cortex and individual channel positions were superimposed onto the functional maps of orientation columns obtained previously with OI. Subsequently, channels corresponding to the desired locations in the functional map were used as guide tubes for electrode insertion. The spatial precision of this approach was in the range of 100 microm and experiments in cat primary visual cortex revealed a close correlation between the desired and the actually recorded orientation preferences of the targeted columns. The method is applicable to all cortical areas in which OI is feasible and offers a high degree of flexibility with respect to the number and geometry of applicable probes. It is, thus, an excellent tool for studying distributed codes and interactions between multiple predefined recording sites.

Animals↗

Corticostriatal and corticotectal projections from visual cortical areas 17, 18 and 18a in the pigmented rat.

Regions of the visual cortex and the deep layers of the superior colliculus (SC) have been suggested to be functionally linked via an 'indirect' pathway through the basal ganglia. The present report demonstrates projections from the striate (area 17) and extrastriate (areas 18 and 18a) visual cortex in Long-Evans hooded rats to the striatum (ST) and SC with anterograde (biocytin) and retrograde (WGA-HRP and fluorescent dyes) tracers. Biocytin injections into each cortical area produced markedly different patterns of labeling in ST and SC. Injections into area 17 resulted in a dense labeling in the superficial layers of SC, with little labeling present in the deep layers of SC or ST. In contrast, injections into area 18a, which produced marked labeling in the deep layers of SC with moderate in the superficial layers, resulted in dense labeling in the caudal two-thirds of the dorsal region of ST; injections into area 18 produced numerous terminals mainly in the deep layers of SC and in the rostral two-thirds of the dorsal aspect of ST. WGA-HRP injections into ST resulted in numerous retrogradely labeled pyramidal neurons in areas 18 and 18a, but only a few labeled neurons were observed in area 17. Corticostriatal neurons in extrastriate visual cortex were distributed predominantly in layer V, with smaller numbers in layers II and III, whereas corticotectal neurons were located only in layer V of both the striate and extrastriate visual cortex. Although corticostriatal and corticotectal neurons were intermingled in layer V, no double-labeled neurons were observed following injections of different fluorescent dyes into ST and SC. It appears, therefore, that: 1) the major source of visual input to both ST and the deep layers of SC in the rat arises not from the primary visual cortex but from extrastriate visual areas; and that 2) the projections from the extrastriate visual area to ST and SC originate from different populations of corticofugal neurons.

Animals↗

Flashed pattern-induced activity in the visual system: I. The short latency evoked response recorded from the cat visual cortex.

Previous methods for estimating visual acuity have used the visual evoked response conventionally the late visual evoked response components or the steady-state potential. The present experiments were undertaken to evaluate the possible use of short latency flashed pattern evoked responses in estimating pattern dependent activity in the cat visual system. Recordings were made from the skull bone and the dura above the primary visual cortex and intracortically. The visual evoked responses to patterned (checks) and non-patterned light flashes of high intensity and short duration were recorded. The visual evoked response activity recorded from the cortical surface had an onset latency of 14-15 ms. The initial positive-negative potential sequences of the responses were similar for patterned and non-patterned stimuli, however a difference was recorded from 35-40 ms after stimulus. The smallest check size which separated a pattern from a non-pattern VER was in the order of 10 min of arc. The results indicate that the short-latency cortical VER may be used to estimate visual resolution.

Animals↗

Structural covariance in the human cortex.

The morphology of the human cortex varies remarkably across individuals, regardless of overall brain size. It is currently unclear whether related cortical regions covary in gray matter density, as a result of mutually trophic influences or common experience-related plasticity. We acquired a structural magnetic resonance imaging scan from 172 subjects and extracted the regional gray matter densities from 12 readily identifiable regions of interest involved in sensorimotor or higher-order cognitive functions. We then used these values to predict regional densities in the remaining areas of the cortex, using voxel-based morphometry. This revealed patterns of positive and negative covariance that provide insight into the topographical organization of multiple cortical regions. We report that the gray matter density of a region is a good predictor of the density of the homotopic region in the contralateral hemisphere, with the striking exception of primary visual cortex. Whereas some regions express patterns of regional covariance that are mirror symmetrical relative to the interhemispheric fissure, other regions express asymmetric patterns of regional covariance. Finally, patterns of covariance are remarkably consistent between males and females, with the exception of the left amygdala, which is positively associated with the left and right anterior inferior temporal cortex in males and with the right angular gyrus in females. Our study establishes that the density of different cortical regions is coordinated within an individual. The coordinated variations we report are likely to be determined by both genetic and environmental factors and may be the basis for differences in individual behavior.

Adult↗

Brain activity in visual cortex predicts individual differences in reading performance.

The relationship between brain activity and reading performance was examined to test the hypothesis that dyslexia involves a deficit in a specific visual pathway known as the magnocellular (M) pathway. Functional magnetic resonance imaging was used to measure brain activity in dyslexic and control subjects in conditions designed to preferentially stimulate the M pathway. Dyslexics showed reduced activity compared with controls both in the primary visual cortex and in a secondary cortical visual area (MT+) that is believed to receive a strong M pathway input. Most importantly, significant correlations were found between individual differences in reading rate and brain activity. These results support the hypothesis for an M pathway abnormality in dyslexia and imply a strong relationship between the integrity of the M pathway and reading ability.

Dyslexia↗

Serotoninergic innervation of the ferret cerebral cortex. I. Adult pattern.

We have investigated the serotoninergic innervation of the adult ferret cerebral cortex with immunohistochemical techniques. Distribution pattern of serotoninergic fibers in the ferret neocortex is characterized by a decrease in the density of fibers as one moves from the pial surface towards the white matter. Throughout the entire cerebral cortex, the serotoninergic fibers are very dense within the supragranular layers, especially within layer 1. In contrast, granular and infragranular layers exhibit only a sparse innervation. Although this general pattern of innervation is roughly the same in all cortical areas, significant variations in the fiber density are apparent in different regions. Areas 17, 1, 6, and 8 (primary visual cortex, presumptive somatosensory cortex, presumptive motor cortex, and prefrontal cortex, respectively) are described in more detail to illustrate the diversity of the serotoninergic innervation patterns. The density of innervation is highest in areas 1 and 6, intermediate in area 8, and lowest in area 17. It is noteworthy that while areas 1, 6, and 8 show a marked decrease in fiber density at the boundary between layer 3 and 4, the less strongly innervated area 17 shows a change in density in the transition from layer 2 to layer 3. The types of fibers found within the ferret cortex are similar to those described in other mammalian species. The bulk of the innervation is made by very fine fusiform axons with small ovoid varicosities. In addition to this fiber type, axons with thick round varicosities and some smooth nonvaricose axons were found. The latter types occur in very small numbers within the supragranular layers and mostly in more anterior cortical regions. While the general innervation pattern and the fiber types are similar to those described in the cat cerebral cortex, the pericellular baskets found in the cat cortex (Mulligan and Törk, J Comp Neurol 270:86-110, 1988) are not seen in the ferret.

Animals↗

Loss of neurons in magnocellular and parvocellular layers of the lateral geniculate nucleus in glaucoma.

OBJECTIVES: To determine whether there is loss of lateral geniculate nucleus relay neurons, which convey visual information to the visual cortex, in experimental glaucoma in monkeys. METHOD: Four cynomolgus monkeys with experimentally induced glaucoma in the right eye (referred to as the glaucoma group) and 5 control monkeys were studied. In both groups, the same conditions of fixation, tissue processing, staining, and measurement were used. In each monkey, the left lateral geniculate nucleus target neurons in magnocellular layer 1 and parvocellular layers 4 and 6, connected to the right glaucomatous eye, were studied. Immunocytochemistry with antibody to parvalbumin was used to specifically label relay neurons connecting to the visual cortex. The number of parvalbumin-immunoreactive neurons was estimated using an unbiased 3-dimensional counting method. The t test was used to compare the experimental and control groups. RESULTS: The mean ( SD) number of neurons in mavnocellular layer 1 was significantly decreased in the glaucoma group compared with the control group (20 692 9567 vs 37 687 8017; P = .02). The mean ( SD) number of neurons in parvocellular layers 4 and 6 was significantly decreased in the glaucoma group compared with the control group (100 141 44 906 vs 174 090 39 136; P = .03). Data are given as the mean SD. CONCLUSION: Significant loss of lateral geniculate nucleus relay neurons terminating in the primary visual cortex occurs in the magnocellular and parvocellular layers in an experimental monkey model of glaucoma. CLINICAL RELEVANCE: Knowledge of the fate of neurons in the central visual system may lead to a better understanding of the nature and progression of visual loss in glaucomatous optic neuropathy.

Animals↗

Conserved patterns of cortico-cortical connections define areal hierarchy in rat visual cortex.

The prevalence of reciprocal connections in the cerebral cortex indicates that they play a fundamental role in the processing of sensory information. We have investigated the laminar termination patterns of such paired connections between different visual cortical areas of the rat, and have found two basic projection types: one which includes layer 4 and a second which includes layer 1 and avoids layer 4. The projections from primary visual cortex (area 17) to extrastriate visual cortical targets in the cytoarchitectonical areas 18a and 18b, and from 18a to a site in 18b, are of the first type. In contrast, the return projections from 18a and 18b to area 17 and from 18b to 18a, are of the second type. Thus each pair of connections has one element of each type, giving every circuit a nearly identical asymmetric structure. These laminar patterns resemble those of forward and feedback connections in primate cortex, indicating that corticocortical connectivity patterns are highly conserved through evolution, and that, as in monkeys, these connections define a hierarchical organization of areas in rat visual cortex.

Animals↗

Cortical and thalamic visual evoked potentials during sleep-wake states and spike-wave discharges in the rat.

Flash visual evoked potentials (VEP) were simultaneously recorded from the primary visual cortex and the dorsal lateral geniculate nucleus in freely-moving WAG/Rij rats, to investigate whether the thalamic VEP shows the same state-dependent alterations as the cortical VEP. VEPs obtained during active and passive wakefulness (AW and PW), slow-wave sleep (SWS), REM sleep and during the occurrence of spike-wave discharges (SWD), a specific trait of the genetically epileptic WAG/Rij rat, were compared. The general architecture of the thalamic VEP resembles the cortical VEP, although its polarity is reversed. This facilitated the interpretation of components in terms of underlying neuronal events. The primary excitation peak is differently modulated in cortex and thalamus. Whereas the thalamic component (P30) is not affected by brain-state, the cortical component (N1) shows a strong increase in latency during SWS and SWD. In contrast, the modulation of later components is highly similar for cortex and thalamus. VEPs obtained during AW and REM resemble each other. During SWS and SWD there is a considerable, and during PW a moderate, enlargement of primarily inhibitory components. After-discharges are enhanced during SWS, SWD and REM. No evidence is found for a major impairment of sensory transmission during SWD.

Action Potentials↗

Axo-axonic synapses formed by somatostatin-expressing GABAergic neurons in rat and monkey visual cortex.

In cerebral cortex of rat and monkey, the neuropeptide somatostatin (SOM) marks a population of nonpyramidal cells (McDonald et al. [1982] J. Neurocytol. 11:809-824; Hendry et al. [1984] J. Neurosci. 4:2497:2517; Laemle and Feldman [1985] J. Comp. Neurol. 233:452-462; Meineke and Peters [1986] J. Neurocytol. 15:121-136; DeLima and Morrison [1989] J. Comp. Neurol. 283:212-227) that represent a distinct type of gamma-aminobutyric acid (GABA) -ergic neuron (Gonchar and Burkhalter [1997] Cereb. Cortex 7:347-358; Kawaguchi and Kubota [1997] Cereb. Cortex 7:476-486) whose synaptic connections are incompletely understood. The organization of inhibitory inputs to the axon initial segment are of particular interest because of their role in the suppression of action potentials (Miles et al. [1996] Neuron 16:815:823). Synapses on axon initial segments are morphologically heterogeneous (Peters and Harriman [1990] J. Neurocytol. 19:154-174), and some terminals lack parvalbumin (PV) and contain calbindin (Del Rio and DeFelipe [1997] J. Comp. Neurol. 342:389-408), that is also expressed by many SOM-immunoreactive neurons (Kubota et al. [1994] Brain Res. 649:159-173; Gonchar and Burkhalter [1997] Cereb. Cortex 7:347-358). We studied the innervation of pyramidal neurons by SOM neurons in rat and monkey visual cortex and examined putative contacts by confocal microscopy and determined synaptic connections in the electron microscope. Through the confocal microscope, SOM-positive boutons were observed to form close appositions with somata, dendrites, and spines of intracortically projecting pyramidal neurons of rat area 17 and pyramidal cells in monkey striate cortex. In addition, in rat and monkey, SOM boutons were found to be associated with axon initial segments of pyramidal neurons. SOM axon terminals that were apposed to axon initial segments of pyramidal neurons lacked PV, which was shown previously to label axo-axonic terminals provided by chandelier cells (DeFelipe et al. [1989] Proc. Natl. Acad. Sci. USA 86:2093-2097; Gonchar and Burkhalter [1999a] J. Comp. Neurol. 406:346:360). Electron microscopic examination directly demonstrated that SOM axon terminals form symmetric synapses with the initial segments of pyramidal cells in supragranular layers of rat and monkey primary visual cortex. These SOM synapses differed ultrastructurally from the more numerous unlabeled symmetric synapses found on initial segments. Postembedding immunostaining revealed that all SOM axon terminals contained GABA. Unlike PV-expressing chandelier cell axons that innervate exclusively initial segments of pyramidal cell axons, SOM-immunoreactive neurons innervate somata, dendrites, spines, and initial segments, that are just one of their targets. Thus, SOM neurons may influence synaptic excitation of pyramidal neurons at the level of synaptic inputs to dendrites as well as at the initiation site of action potential output.

Animals↗

Activation of color-selective areas of the visual cortex in a blind synesthete.

Many areas of the visual cortex are activated when blind people are stimulated naturally through other sensory modalities (e.g., haptically; Sadato et al., 1996). While this extraneous activation of visual areas via other senses in normal blind people might have functional value (Kauffman et al., 2002; Lessard et al., 1998), it does not lead to conscious visual experiences. On the other hand, electrical stimulation of the primary visual cortex in the blind does produce illusory visual phosphenes (Brindley and Lewin, 1968). Here we provide the first evidence that high-level visual areas not only retain their specificity for particular visual characteristics in people who have been blind for long periods, but that activation of these areas can lead to visual sensations. We used fMRI to demonstrate activity in visual cortical areas specifically related to illusory colored and spatially located visual percepts in a synesthetic man who has been completely blind for 10 years. No such differential activations were seen in late-blind or sighted non-synesthetic controls; neither were these areas activated during color-imagery in the late-blind synesthete, implying that this subject's synesthesia is truly a perceptual experience.

Association↗

The organization of serotonergic projections to cerebral cortex in primates: regional distribution of axon terminals.

Serotonergic axons are widely distributed in the primate forebrain and represent the most abundant ascending projection from the reticular formation. Immunocytochemical methods have been utilized to examine the density, laminar distribution and morphology of serotonergic axons in both primary projection (motor, somatosensory) and association areas (dorsolateral prefrontal, area 5) as well as in the hippocampus and in cingulate cortex of rhesus and cynomolgus macaques. Serotonergic axons are present in all areas of cortex examined, and all cortical layers receive serotonergic afferents. However, the intracortical distribution of serotonergic axon terminals is not uniform; rather, different regions of cortex exhibit dissimilarities in both the density and laminar distribution of serotonergic axons. Thus, there are local patterns of serotonin innervation that are characteristic of each cortical area. Highly diverse patterns of serotonin innervation are found in heterotypical areas of cortex; more subtle variations are present among homotypical areas. Two morphologic types of serotonergic axon terminals, fine and beaded, are present in all cortical areas, and they typically exhibit different laminar distributions: in most areas of neocortex, beaded axons predominate in layer I while fine axons predominate in layers II-VI. However, exceptions to this pattern were observed in primary visual cortex and in the hippocampal formation. The distinctive local patterns of serotonin innervation observed in this study indicate that raphe-cortical projections are likely to have differential influences on particular cytoarchitectonic areas of cerebral cortex in the primate. Moreover, the discrete laminar distribution of serotonin axons suggests that serotonergic projections selectively innervate particular neuronal elements in cerebral cortex. The present findings suggest that the two classes of serotonergic axons, fine and beaded, which have different patterns of termination, affect different sets of cortical neurons. In addition, these two serotonergic projections may be associated with different sets of serotonergic receptors and thus produce selective effects on cortical function.

Afferent Pathways↗

Functional and diffusion-weighted magnetic resonance imaging for visualization of the postthalamic visual fiber tracts and the visual cortex.

Diffusion-weighted magnetic resonance imaging (MRI) offers the possibility to study the course of the cerebral white matter tracts whereas functional MRI (fMRI) provides information about the specific functions of cortical areas. We evaluated the combination of fMRI and diffusion-weighted MRI to detect cortical visual areas with their corresponding visual fiber tracts in 15 healthy controls (age: 23 - 53 years, male : female = 8 : 7). We demonstrated activation within the primary visual cortex and white matter bundles connecting the lateral geniculate body and the striate cortex in all subjects investigated. Additional activation could be appreciated in some subjects within the lateral geniculate bodies (n = 2) and the motion-sensitive area V5 (n = 3). The combination of diffusion-weighted and functional imaging allows visualization of the origin, direction and functionality of large white matter tracts. This will prove helpful for imaging structural connectivity within the brain during functional imaging. Moreover, this technique might provide important information for neurosurgical patients presenting with space-occupying lesions close to the cortical and subcortical visual system since this technique can -- in contrast to diffusion tensor imaging -- easily be adopted into a neuronavigation system and can be performed on all MR scanners capable of diffusion-weighted imaging without specific post-processing programs.

Adult↗

The physiology of coloured hearing. A PET activation study of colour-word synaesthesia.

In a small proportion of the normal population, stimulation in one modality can lead to perceptual experience in another, a phenomenon known as synaesthesia. In the most common form of synaesthesia, hearing a word can result in the experience of colour. We have used the technique of PET, which detects brain activity as changes of regional cerebral blood flow (rCBF), to study the physiology of colour-word synaesthesia in a group of six synaesthete women. During rCBF measurements synaesthetes and six controls were blindfolded and were presented with spoken words or pure tones. Auditory word, but not tone, stimulation triggered synaesthesia in synaesthetes. In both groups word stimulation compared with tone stimulation activated the classical language areas of the perisylvian regions. In synaesthetes, a number of additional visual associative areas, including the posterior inferior temporal cortex and the parieto-occipital junctions, were activated. The former has been implicated in the integration of colour with shape and in verbal tasks which require attention to visual features of objects to which words refer. Synaesthetes also showed activations in the right prefrontal cortex, insula and superior temporal gyrus. By contrast, no significant activity was detected in relatively lower visual areas, including areas V1, V2 and V4. These results suggest that colour-word synaesthesia may result from the activity of brain areas concerned with language and visual feature integration. In the case of colour-word synaesthesia, conscious visual experience appears to occur without activation of the primary visual cortex.

Adult↗