Hemispheric asymmetries in feature integration during visual word recognition

Although the definitive source of the left hemisphere's superiority for visual word recognition remains illusive, some argue that the left (LH) and right (RH) hemispheres engage different strategies during early perceptual processes involved in stimulus encoding. In particular, it is proposed that the LH treats a word as a unitary perceptual group whereas the RH processes the letters comprising a word as a series of individual perceptual units. The present study investigated support for this processing distinction by examining hemispheric strategies for temporal integration using Prinzmetal and Millis-Wright's (1984 Prinzmetal, W. and Millis-Wright, M. 1984. Cognitive and linguistic factors affect visual feature integration. Cognitive Psychology, 16: 305–340. [Crossref], [PubMed], [Web of Science ®] , [Google Scholar]) feature-binding paradigm. A total of 20 participants identified the colour and identity of a target letter, presented within a three-letter word (e.g., ART) or nonword (e.g., HRF), directed to their left or right visual field. Errors were classified on the basis of whether they involved substitution of a colour present within the stimulus but at a different location (ON error), or the substitution of a colour not present within the stimulus (OFF error). As anticipated, for word stimuli there was a higher proportion of OFF errors associated with trials directed to the RH, consistent with the notion that the LH treats words as single perceptual units and is hence biased toward miscombination of perceptual information present within the stimulus. The pattern of ON errors across stimulus type provided clear evidence of RH sequential encoding effects, with the number of errors increasing markedly across the ordinal position of the letters comprising the stimulus string. As such, these data provide new evidence that the LH's advantage for visual word recognition arises, at least in part, from the ability to encode verbal stimuli as single perceptual units.     

Hemispheric asymmetries in hierarchical stimulus processing are modulated by stimulus categories and their predictability

Hemispheric dominance has been behaviourally documented for the local (left hemisphere, LH) or global (right hemisphere, RH) processing of hierarchical letters. However, Fink et al. (1997) indicated that stimulus category modulates this hemispheric asymmetry. The purpose of this study was to investigate the influence of the category (letters versus objects) on hemispheric specialisation for global and local processing using a visual half-field presentation in a task where participants ignored whether the target appeared at the global or local level. In Experiment 1 we replicated the classic hemispheric asymmetry for global/local processing of hierarchical letters. In Experiment 2, which consisted of hierarchical object processing, a RH dominance for the local level was observed. In Experiment 3 a within-participant design was used where anticipation about the stimulus category was precluded, resulting in the classic RH and LH specialisations for global and local processing for both letter-based and object-based stimuli. Taken together, these results suggest that the highly demanding local processing stage engages one hemisphere more than the other, according to the lateralisation of cerebral networks specialised for stimulus category. In addition, the direction of lateralisation for the local level was also modulated by the predictability of the stimulus category.     

Hemispheric asymmetries in hierarchical stimulus processing are modulated by stimulus categories and their predictability

Hemispheric dominance has been behaviourally documented for the local (left hemisphere, LH) or global (right hemisphere, RH) processing of hierarchical letters. However, Fink et al. (1997) indicated that stimulus category modulates this hemispheric asymmetry. The purpose of this study was to investigate the influence of the category (letters versus objects) on hemispheric specialisation for global and local processing using a visual half-field presentation in a task where participants ignored whether the target appeared at the global or local level. In Experiment 1 we replicated the classic hemispheric asymmetry for global/local processing of hierarchical letters. In Experiment 2, which consisted of hierarchical object processing, a RH dominance for the local level was observed. In Experiment 3 a within-participant design was used where anticipation about the stimulus category was precluded, resulting in the classic RH and LH specialisations for global and local processing for both letter-based and object-based stimuli. Taken together, these results suggest that the highly demanding local processing stage engages one hemisphere more than the other, according to the lateralisation of cerebral networks specialised for stimulus category. In addition, the direction of lateralisation for the local level was also modulated by the predictability of the stimulus category.     

Evaluating a split processing model of visual word recognition: effects of word length

A new theory of visual word recognition is based on the fact that the fovea is split in humans. When a reader fixates the center of a written word, the initial letters of the word that are to the left of fixation are projected first to the right cerebral hemisphere (RH) while the final letters are projected to the left cerebral hemisphere (LH). This paper explores the possibility that this has consequences for the early processing of the beginning and ends of centrally fixated words: specifically that lexical decision RTs are affected by the number of letters to the left of fixation but not by the number of letters to the right of fixation. For centrally presented five- and eight-letter words, we manipulated number of letters presented to the right or to the left of a fixation point (Experiment 1). We found that longer latencies to longer letter strings characterised the processing of the initial letters of words while LH word recognition features characterised the ends of words. Experiment 2 was a lateralized version of Experiment 1, and revealed the well established visual field and word length interaction. The results supported the split fovea theory.     

From Receptive to Perceptive Fields: Size-Dependent Asymmetries in Both Negative Afterimages and Subcortical On and Off Post-Stimulus Responses

Negative afterimages are perceptual phenomena that occur after physical stimuli disappear from sight. Their origin is linked to transient post-stimulus responses of visual neurons. The receptive fields (RFs) of these subcortical ON- and OFF-center neurons exhibit antagonistic interactions between central and surrounding visual space, resulting in selectivity for stimulus polarity and size. These two features are closely intertwined, yet their relationship to negative afterimage perception remains unknown. Here we tested whether size differentially affects the perception of bright and dark negative afterimages in humans of both sexes, and how this correlates with neural mechanisms in subcortical ON and OFF cells. Psychophysically, we found a size-dependent asymmetry whereby dark disks produce stronger and longer-lasting negative afterimages than bright disks of equal contrast at sizes >0.8°. Neurophysiological recordings from retinal and relay cells in female cat dorsal lateral geniculate nucleus showed that subcortical ON cells exhibited stronger sustained post-stimulus responses to dark disks, than OFF cells to bright disks, at sizes >1°. These sizes agree with the emergence of center-surround antagonism, revealing stronger suppression to opposite-polarity stimuli for OFF versus ON cells, particularly in dorsal lateral geniculate nucleus. Using a network-based retino-geniculate model, we confirmed stronger antagonism and temporal transience for OFF-cell post-stimulus rebound responses. A V1 population model demonstrated that both strength and duration asymmetries can be propagated to downstream cortical areas. Our results demonstrate how size-dependent antagonism impacts both the neuronal post-stimulus response and the resulting afterimage percepts, thereby supporting the idea of perceptual RFs reflecting the underlying neuronal RF organization of single cells.SIGNIFICANCE STATEMENT Visual illusions occur when sensory inputs and perceptual outcomes do not match, and provide a valuable tool to understand transformations from neural to perceptual responses. A classic example are negative afterimages that remain visible after a stimulus is removed from view. Such perceptions are linked to responses in early visual neurons, yet the details remain poorly understood. Combining human psychophysics, neurophysiological recordings in cats and retino-thalamo-cortical computational modeling, our study reveals how stimulus size and the receptive-field structure of subcortical ON and OFF cells contributes to the parallel asymmetries between neural and perceptual responses to bright versus dark afterimages. Thus, this work provides a deeper link from the underlying neural mechanisms to the resultant perceptual outcomes.     

Hemispheric Asymmetry for Word Naming: Effects of Frequency and Regularity of Pronunciation

Observers named visually presented words as quickly as possible when the words were presented to the centre of a viewing screen (Experiments 1, 5, and 6) or were flashed to either the left visual field/right hemisphere (LVF/RH) or right visual field/left hemisphere (RVF/LH) on each trial (Experiments, 2, 3, and 4). Words varied in frequency of occurrence in the language and in regularity of pronunciation. On lateralised trials, there was a RVF/LH advantage for both error rate and reaction time, with this hemispheric asymmetry for word naming being independent of both frequency and regularity. The RVF/LH advantage was also independent of whether the letters within a word were arranged horizontally or vertically. Error rates and reaction times were lower for high-than for lowfrequency words and lower for words with a regular pronunciation than for exception words. In addition, on centre trials the effects of regularity were larger for low-frequency words than for high-frequency words. However, when words were presented in the visual periphery, the effects of regularity were of the same magnitude for high-and low-frequency words. In view of the theoretical importance of the Frequency by Regularity interaction, this latter result suggests that word processing is qualitatively different in the visual periphery than in the centre of vision; indicating that perceptual asymmetries in a typical visual halffield experiment may be limited in what they can tell us about the relative contributions of the two hemispheres to processing words in the centre of vision.     

Hemispheric asymmetry for word naming: effects of frequency and regularity of pronunciation

Observers named visually presented words as quickly as possible when the words were presented to the centre of a viewing screen (Experiments 1, 5, and 6) or were flashed to either the left visual field/right hemisphere (LVF/RH) or right visual field/left hemisphere (RVF/LH) on each trial (Experiments, 2, 3, and 4). Words varied in frequency of occurrence in the language and in regularity of pronunciation. On lateralised trials, there was a RVF/LH advantage for both error rate and reaction time, with this hemispheric asymmetry for word naming being independent of both frequency and regularity. The RVF/LH advantage was also independent of whether the letters within a word were arranged horizontally or vertically. Error rates and reaction times were lower for high-than for low-frequency words and lower for words with a regular pronunciation than for exception words. In addition, on centre trials the effects of regularity were larger for low-frequency words than for high-frequency words. However, when words were presented in the visual periphery, the effects of regularity were of the same magnitude for high-and low-frequency words. In view of the theoretical importance of the Frequency by Regularity interaction, this latter result suggests that word processing is qualitatively different in the visual periphery than in the centre of vision; indicating that perceptual asymmetries in a typical visual half-field experiment may be limited in what they can tell us about the relative contributions of the two hemispheres to processing words in the centre of vision.     

A magnetic stimulation examination of orthographic neighborhood effects in visual word recognition

The split-fovea theory proposes that visual word recognition is mediated by the splitting of the foveal image, with letters to the left of fixation projected to the right hemisphere (RH) and letters to the right of fixation projected to the left hemisphere (LH). We applied repetitive transcranial magnetic stimulation (rTMS) over the left and right occipital cortex during a lexical decision task to investigate the extent to which word recognition processes could be accounted for according to the split-fovea theory. Unilateral rTMS significantly impaired lexical decision latencies to centrally presented words, supporting the suggestion that foveal representation of words is split between the cerebral hemispheres rather than bilateral. Behaviorally, we showed that words that have many orthographic neighbors sharing the same initial letters ("lead neighbors") facilitated lexical decision more than words with few lead neighbors. This effect did not apply to end neighbors (orthographic neighbors sharing the same final letters). Crucially, rTMS over the RH impaired lead-, but not end-neighborhood facilitation. The results support the split-fovea theory, where the RH has primacy in representing lead neighbors of a written word.     

Re-evaluating split-fovea processing in word recognition: Effects of word length

Several studies have claimed that, when fixating a word, letters to the left and right of fixation project to different hemispheres and are consequently subjected to different processes. In support of this claim, Lavidor M, et al. (2001; hereafter LES&B) report that lexical decisions were affected by increasing the number of letters to the left of fixation but not to the right, and that this indicates divided hemispheric access at the point of fixation to length-sensitive processes in the right hemisphere (RH). We re-evaluated these claims in Experiment 1 using Lavidor et al.'s original stimuli and procedure of merely instructing participants where to fixate. In contrast to the earlier study, increases in the number of letters to the left and right of the designated fixation location produced near-identical effects on reaction time, and increases to the left actually improved response accuracy and increases to the right impaired it. When larger stimuli were used to improve stimulus perceptibility and an eye-tracker monitored fixation accuracy (Experiment 2), left and right increases in the number of letters again produced near-identical effects on reaction time (and accuracy), but frequent and substantial fixation errors were revealed. When an eye-tracker ensured accurate fixations (Experiment 3), left and right increases in the number of letters again produced near-identical effects on reaction time and accuracy. Thus, the findings of all three experiments provide no support for the findings of LES&B (2001) and no evidence of split-fovea processing. The findings also indicate the dangers of assuming fixation of precisely-specified locations within words, both in experiments designed to reveal split-foveal processing and hemispheric asymmetry and in more normal circumstances of word perception.     

Right‐hemispheric processing of non‐linguistic word features: Implications for mapping language recovery after stroke

Verbal stimuli often induce right‐hemispheric activation in patients with aphasia after left‐hemispheric stroke. This right‐hemispheric activation is commonly attributed to functional reorganization within the language system. Yet previous evidence suggests that functional activation in right‐hemispheric homologues of classic left‐hemispheric language areas may partly be due to processing nonlinguistic perceptual features of verbal stimuli. We used functional MRI (fMRI) to clarify the role of the right hemisphere in the perception of nonlinguistic word features in healthy individuals. Participants made perceptual, semantic, or phonological decisions on the same set of auditorily and visually presented word stimuli. Perceptual decisions required judgements about stimulus‐inherent changes in font size (visual modality) or fundamental frequency contour (auditory modality). The semantic judgement required subjects to decide whether a stimulus is natural or man‐made; the phonologic decision required a decision on whether a stimulus contains two or three syllables. Compared to phonologic or semantic decision, nonlinguistic perceptual decisions resulted in a stronger right‐hemispheric activation. Specifically, the right inferior frontal gyrus (IFG), an area previously suggested to support language recovery after left‐hemispheric stroke, displayed modality‐independent activation during perceptual processing of word stimuli. Our findings indicate that activation of the right hemisphere during language tasks may, in some instances, be driven by a “nonlinguistic perceptual processing” mode that focuses on nonlinguistic word features. This raises the possibility that stronger activation of right inferior frontal areas during language tasks in aphasic patients with left‐hemispheric stroke may at least partially reflect increased attentional focus on nonlinguistic perceptual aspects of language. Hum Brain Mapp, 2013. © 2012 Wiley Periodicals, Inc.     

The effect of stimulus features on working memory of categorical and coordinate spatial relations in patients with unilateral brain damage

Spatial relations are typically divided into categorical and coordinate spatial relations. Categorical relations are abstract and show a left hemisphere (LH) advantage, whereas coordinate relations are metric and related to a right hemisphere (RH) advantage. In the current study a working memory task was used to asses categorical and coordinate performance with two different stimulus sets. In this task, participants had to compare two sequentially presented stimuli, consisting of a dot and a cross. The cross size used in the stimuli was either large or small; a direct manipulation of the amount of information provided to determine a category, or to assess a distance. Patients with damage in the LH or the RH and highly comparable controls were tested. In control participants, categorical processing is faster with the use of a large cross, i.e., more visual information about category boundaries. In contrast, coordinate performance was more accurate with a small cross, i.e., presenting less unnecessary visual information. LH patients showed a specific defect in processing categorical stimuli with a small cross and coordinate stimuli with a large cross. The RH patients were impaired in all conditions except for the categorical small cross condition. We conclude that a larger amount of information present in stimuli increases categorical processing performance and decreases coordinate processing performance, while opposite effects are found for less stimulus information.     

Effects of form familiarity on perception of words,pseudowords, and nonwords in the two cerebral hemispheres

Previous investigations of hemispheric processes of word perception provide a mixed picture of the sensitivity of each hemisphere to the familiarity of the visual form of lateralized displays. We investigated this issue by presenting words, pseudowords, and nonwords briefly to either the left (LH) or right (RH) hemisphere in lowercase, uppercase, and a matched, unfamiliar mixed-case form, and used an eye tracker to ensure central fixation and the Reicher-Wheeler task to suppress influences of stimulus asymmetry. Familiarity of form exerted a substantial effect on perception. In particular, perception of LH and RH displays of words, pseudowords, and nonwords was least accurate for mixed case, intermediate for upper case, and most accurate for lowercase. However, form had no effect on the LH advantage observed for words, pseudowords, and nonwords, indicating that form affected processing in both hemispheres to a similar extent. Moreover, LH and RH displays both showed that mixed case disrupted performance most for words, and more for pseudowords than for nonwords, indicating the sensitivity to form shown by each hemisphere reflected more than a general perceptual process. Implications for the role of form familiarity in hemispheric processing of words are discussed.     

Priming by natural category membership in the left and right cerebral hemispheres

The cerebral representation of category information was examined in a single word priming paradigm, during which the N400 component of the event-related potential (ERP) was recorded. The visual half-field paradigm was employed in order to selectively stimulate the two hemispheres. To investigate which aspects of category membership are relevant in producing priming, two types of related stimuli were presented. In one condition pairs of exemplars had a higher amount of feature overlap (e.g., MOSQUITO-FLEA) than in the other (e.g., SOFA–VASE). Significant priming was obtained only for stimuli in the high feature overlap condition and then only when these were presented to the left visual field (LVF)/right hemisphere (RH). This finding was interpreted within our recent model of semantic memory wherein the right hemisphere represents items on the basis of distributed individual features, whereas the left hemisphere (LH) represents semantic information locally, within a spreading activation system, where priming occurs exclusively through associative links. It was concluded that knowledge regarding category membership is maintained in the RH, on the basis of feature coding.     

Faster thalamocortical processing for dark than light visual targets

ON and OFF visual pathways originate in the retina at the synapse between photoreceptor and bipolar cells. OFF bipolar cells are shorter in length and use receptors with faster kinetics than ON bipolar cells and, therefore, process information faster. Here, we demonstrate that this temporal advantage is maintained through thalamocortical processing, with OFF visual responses reaching cortex ~3-6 ms before ON visual responses. Faster OFF visual responses could be demonstrated in recordings from large populations of cat thalamic neurons representing the center of vision (both X and Y) and from subpopulations making connection with the same cortical orientation column. While the OFF temporal advantage diminished as visual responses reached their peak, the integral of the impulse response was greater in OFF than ON neurons. Given the stimulus preferences from OFF and ON channels, our results indicate that darks are processed faster than lights in the thalamocortical pathway.     

The centre is not in the middle: Evidence from line and word bisection

English and German readers have been shown to mark a position to the left of the true centre as the subjective midpoint in word bisection. This effect resembles a well-known phenomenon observed with the bisection of solid lines (pseudoneglect), although this behavioural similarity does not imply a common origin. The purpose of the present study was twofold: on the one hand, to investigate the perceptual and lexical features that influence the bisection of Italian orthographic strings and, on the other hand, to investigate whether identical or partially independent processing mediate bisection of line and orthographic stimuli. Five experiments were carried out to explore to what extent stimulus type (lines, words, pseudowords, consonant strings, symbols), stimulus length (from 3 to 13 characters), list context (pure and mixed), and written word frequency (high and low) affected the bisection performance. The results showed that list context modulated the processing similarities across different materials and that word frequency failed to influence the magnitude of the bisection bias.More critically, across all five experiments, the results showed different effects for solid lines versus orthographic material. Lines were always bisected to the left, independent of length and list context. By contrast, a crossover effect emerged with orthographic material; for long stimuli (above five letters) the bias was consistently to the left, while short stimuli showed a consistent rightward bias. The results indicate that manual bisection involved partly different cognitive mechanisms during word and line perception and that this may depend on the characteristics of the stimuli (words/discrete vs. lines/continuous).     

Semantically convergent and semantically divergent priming in the cerebral hemispheres: lexical decision and semantic judgment

The effects of semantically divergent and semantically convergent priming on the processing of lexical ambiguity by the two cerebral hemispheres were examined in two visual hemifield experiments. The experiments investigated the ability of the right hemisphere (RH) and the left hemisphere (LH) to summate activation from two single word primes followed by a laterally presented ambiguous target word. In a lexical decision task (experiment 1), the two priming words were either both related to the dominant meaning of the target (new, fresh-NOVEL), or to the subordinate meaning (story, book-NOVEL), or to one dominant and one subordinate meaning (new, story-NOVEL). Results indicated that the LH benefited most from semantically convergent primes that converged onto the dominant meaning of the ambiguous target word, whereas the RH benefited most from semantically mixed (divergent) primes, that diverge on alternate meanings of the ambiguous target word. We used the same stimuli in a semantic relatedness judgment task (experiment 2), and found that the facilitation in the RH was significantly larger when the primes were mixed than when both primes converged on a single (i.e. either dominant or subordinate) meaning of the to-be-presented target word. In contrast, the only facilitation found in the LH was when the two primes were associated with a single meaning (either dominant or subordinate) of the to-be-presented target word. When the primes were mixed, there was no facilitation in the LH. These results support previous findings indicating that during word recognition, the RH activates a broader range of related meanings than the LH, including alternate meanings of ambiguous words. Thus, by summating activation for seemingly incongruous elements, the RH may be critically involved in at least one important component of verbal creativity.     

Lateral asymmetry: Hard or simple-minded?

A letter or a three-dimensional shape was presented in the center of the visual field. Following the off-set of this stimulus either a comparison letter or a three-dimensional shape was flashed briefly in either the right or left visual field. The subject's task was to respond SAME, or DIFFERENT. The stimuli could be in the same plane, rotated in two dimensions (letters) or in three dimensions (three-dimensional shapes). The left visual field presentations (right hemisphere) of same-pair matches for letters only produced faster reaction times and fewer errors. In all other conditions reaction time measures showed no hemisphere effects. By contrast, error score data indicated that the left hemisphere was overwhelmingly more accurate.     

GABAa and GABAc receptor-mediated modulation of responses to color stimuli: electroretinographic study in the turtle Emys orbicularis

GABAergic transmission is involved in color coding in the retina. The specific contribution of different GABA receptors to spectral sensitivity of the retinal responses is not well characterized. We studied GABAa and GABAc receptor-mediated effects on the intensity-response functions of the electroretinographic ON (b-wave) and OFF (d-wave) responses to color stimuli. For this purpose, we compared the effects of GABAa receptor blockade by bicuculline with the effects of GABAa + GABAc receptor blockade by picrotoxin. The blockade of both GABAa and GABAc receptors caused an amplitude increase of the electroretinographic responses, but the effects of the two blockades depended in a specific manner on stimulus intensity and wavelength. The effects of GABAa receptor blockade showed distinct color ON/OFF asymmetry. The absolute and relative sensitivities of the ON responses to blue stimuli and OFF responses to red stimuli were increased to the greatest degree while the sensitivity of the ON responses to red stimuli and OFF responses to blue stimuli was least increased. In contrast, color ON/OFF asymmetry was not typical of the effects of GABAc receptor blockade. The most prominent GABAc effect was the sensitivity increase of the ON and OFF responses to blue stimuli and, to some lesser extent, to green stimuli. The results of this study indicate a specific role of GABAa and GABAc receptor-mediated influences in processing of chromatic information in the distal retina.     

Taxonomic and ad hoc categorization within the two cerebral hemispheres

A typicality effect refers to categorization which is performed more quickly or more accurately for typical than for atypical members of a given category. Previous studies reported a typicality effect for category members presented in the left visual field/right hemisphere (RH), suggesting that the RH applies a similarity-based categorization strategy. However, findings regarding the typicality effect within the left hemisphere (LH) are less conclusive. The current study tested the pattern of typicality effects within each hemisphere for both taxonomic and ad hoc categories, using words presented to the left or right visual fields. Experiment 1 tested typical and atypical members of taxonomic categories as well as non-members, and Experiment 2 tested typical and atypical members of ad hoc categories as well as non-members. The results revealed a typicality effect in both hemispheres and in both types of categories. Furthermore, the RH categorized atypical stimuli more accurately than did the LH. Our findings suggest that both hemispheres rely on a similarity-based categorization strategy, but the coarse semantic coding of the RH seems to facilitate the categorization of atypical members.     

Expectation-based attentional modulation of visual extinction in spatial neglect

Visual extinction, the failure of patients with unilateral focal brain damage to report the contralesional of two simultaneously presented stimuli, may be modulated by characteristics of the display such as similarity, collinearity, or connectedness. Since these factors affect the perceptual configuration of stimuli, the modulation of extinction is believed to reflect low-level perceptual grouping. In the present study, patient AG did not show any modulation of contralesional detection when the ipsilesional and contralesional stimulus grouped by colour, by form, or both (Experiment 1). In contrast, identification of the contralesional stimulus was facilitated when the stimuli grouped (Experiment 2), suggesting a modulation of extinction by specific task demands. Experiment 3 used a cueing procedure to demonstrate modulation of extinction by expectation biases. Prior to stimulus presentation, AG was cued to attend to a particular feature (e.g. colour). After stimulus exposure he was prompted to identify the expected feature on valid trials and the unexpected feature on invalid trials. AG showed a significant validity effect for contralesional stimuli i.e. he identified the expected feature (e.g. colour) significantly better than the unexpected feature (e.g. form). These results suggest that competition for selection between visual stimuli may not only be influenced by perceptual characteristics of the display, but also by high-level factors such as the response criterion or expectation biases.