Hemispheric specialization for categorical and coordinate spatial representations: A reappraisal

The purpose of the present study was to examine Kosslyn's (1987) claim that the left hemisphere (LH) is specialized for the computation of categorical spatial representations and that the right hemisphere (RH) is specialized for the computation of coordinate spatial representations. Categorical representations involve making judgements about the relative position of the components of a visual stimulus (e.g., whether one component is above/below another). Coordinate representations involve calibrating absolute distances between the components of a visual stimulus (e.g., whether one component is within 5 mm of another). Thirty-two male and 32 female undergraduates were administered two versions of a categorical or a coordinate task over three blocks of 36 trials. Within each block, items were presented to the right visual field-left hemisphere (RVF-LH), the left visual field-right hemisphere (LVF-RH), or a centralized position. Overall, results were more supportive of Kosslyn's assertions concerning the role played by the RH in the computation of spatial representations. Specifically, subjects displayed an LVF-RH advantage when performing both versions of the coordinate task. The LVF-RH advantage on the coordinate task, however, was confirmed to the first block of trials. Finally, it was found that males were more likely than females to display faster reaction times (RTs) on coordinate tasks, slower RTs on categorical tasks, and an LVF-RH advantage in computing coordinate tasks.     

Form-specific visual priming in the right cerebral hemisphere.

Results of 4 experiments indicate that both within-modality and case-specific visual priming for words are greater when test stimuli are presented initially to the right cerebral hemisphere (RH). In contrast, neither within-modality nor case-specific explicit memory for words is greater when stimuli are presented initially to the RH. Priming is measured using word-stem completion, and explicit memory is measured using word-stem cued recall. In both cases, Ss first rate how much they like words, and then word stems are presented briefly to the RH (in the left visual field) or to the left hemisphere (in the right visual field). Results suggest that at least 2 separate systems encode the visual representations that produce priming. The system that is more effective in the RH is better at representing form-specific information, whereas another system that is not more effective in the RH does not distinguish among distinct instances of word forms.     

Language, perception, and the schematic representation of spatial relations

Schemas are abstract nonverbal representations that parsimoniously depict spatial relations. Despite their ubiquitous use in maps and diagrams, little is known about their neural instantiation. We sought to determine the extent to which schematic representations are neurally distinguished from language on the one hand, and from rich perceptual representations on the other. In patients with either left hemisphere damage or right hemisphere damage, a battery of matching tasks depicting categorical spatial relations was used to probe for the comprehension of basic spatial concepts across distinct representational formats (words, pictures, and schemas). Left hemisphere patients underperformed right hemisphere patients across all tasks. However, focused residual analyses using voxel-based lesion-symptom mapping (VLSM) suggest that (1) left hemisphere deficits in the representation of categorical spatial relations are difficult to distinguish from deficits in naming these relations and (2) the right hemisphere plays a special role in extracting schematic representations from richly textured pictures.     

视觉表象产生的大脑半球专门化效应

采用Kosslyn单侧视野速示技术, 以英文字母图片为学习材料, 通过三个实验考察了视觉表象产生的大脑半球专门化效应。实验一提出在两种类型的视觉表象产生任务中, 有两种截然不同的加工起作用, 但却不能直接证实这两种不同加工机制的存在。实验二和实验三则进一步证实了两种表象产生任务具有不同的认知加工机制, 并表现出不同的大脑半球专门化效应。上述研究表明: 大脑两半球均参与产生视觉心理表象, 但分工不同, 并表现出不同的单侧化效应: 大脑左半球通过运用类别空间关系产生表象更有效, 大脑右半球运用数量空间关系产生表象更有效。结果进一步拓展了Kosslyn关于视觉空间关系加工的大脑半球专门化观点。     

Judgments of relative position and distance on representations of spatial relations

This article examines Kosslyn's (1987) hypothesis of the unequal capacity of cerebral hemispheres to process categorical and coordinate spatial relations. Experiment 1 comprised 4 different tasks and failed to support this hypothesis in normal Ss. With the same stimulus patterns as in Kosslyn's study, the results failed to confirm cerebral asymmetry for representing the 2 types of spatial relations, in normal (Experiment 2) and commissurotomized (Experiment 3) Ss. In Experiment 4, a reduction in stimulus luminance produced a partial confirmation of the hypothesis as the right hemisphere proved more adept than the left hemisphere at operating on coordinate representations, whereas both were equally competent at processing categorical spatial-relation representations. The results suggest that the 2 hemispheres can operate on both types of spatial relations, but their respective efficiency depends on the quality of the representations to be processed.     

A Possible Connection between Categorical and Coordinate Spatial Relation Representations

Kosslyn (1987) theorized that the visual system uses two types of spatial relations. Categorical spatial relations represent a range of locations as an equivalence class, whereas coordinate spatial relations represent the precise distance between two objects. Data indicate a left hemisphere (LH) advantage for processing categorical spatial relations and a right hemisphere (RH) advantage for processing coordinate spatial relations. Although generally assumed to be independent processes, this article proposes a possible connection between categorical and coordinate spatial relations. Specifically, categorical spatial relations may be an initial stage in the formation of coordinate spatial relations. Three experiments tested the hypothesis that categorical information would benefit tasks that required coordinate judgments. Experiments 1 and 2 presented categorical information before participants made coordinate judgments and coordinate information before participants made categorical judgments. Categorical information sped the processing of a coordinate task under a range of experimental variables; however, coordinate information did not benefit categorical judgments. Experiment 3 used this priming paradigm to present stimuli in the left or right visual field. Although visual field differences were present in the third experiment, categorical information did not speed the processing of a coordinate task. The lack of priming effects in Experiment 3 may have been due to methodological changes. In general, support is provided that categorical spatial relations may act as an initial step in the formation of more precise distance representations, i.e., coordinate spatial relations.     

The development of categorical and coordinate spatial relations

Two classes of spatial relations can be distinguished in between and within object representations. Kosslyn [Kosslyn, S. M. (1987). Seeing and imagining in the cerebral hemispheres: A computational approach. Psychological Review, 94, 148–175] suggested that the right hemisphere (RH) is specialized for processing coordinate (metric) spatial information and the left hemisphere (LH) processes categorical (abstract) information more effectively. The present study examined the developmental pattern of spatial relation processing in 6–8-year old, 10–12-year old and adults. Using signal detection analyses we calculated sensitivity and bias scores for all age groups. The results indicated that older children and adults showed a greater response bias than younger children. Also, discrimination sensitivity for spatial relation changes clearly improved with age. For the oldest children (10–12-year old) and adults this improvement was accompanied by a RH specialization. In contrast with Kosslyn's claim, this RH advantage also applied to the processing of categorical spatial information. The results are discussed in terms of a right hemispheric specialization for spatial relation processing which matures with age.     

Hemispheric specialization for processing of visually presented verbal and spatial stimuli

Two “same-different” reaction time experiments, analogous in task demands made on the S, were designed to test laterality differences in. perception. Ten normal right-handed Ss performed a verbal task in which they decided whether or not two three-letter words belonged to the same conceptual class. Ten different Ss performed a spatial task in which they decided whether two 16-cell matrices with 3 blackened cells were identical. Reaction times were found to be sensitive to laterality differences in perception. Verbal stimuli were processed faster when presented in the right visual field, and thus projected directly to the left cerebral hemisphere; spatial stimuli were processed faster when presented in the left visual field, and thus projected directly to the right cerebral hemisphere. These results were analyzed in terms of implications regarding hemispheric asymmetries for processing of verbal and spatial material and the nature of interhemispheric transfer of information.     

Cerebral asymmetry for American Sign Language: The effects of moving stimuli

Previous studies of cerebral asymmetry for the perception of American Sign Language (ASL) have used only static representations of signs; in this study we present moving signs. Congenitally deaf, native ASL signers identified moving signs, static representations of signs, and English words. The stimuli were presented rapidly by motion picture to each visual hemifield. Normally hearing English speakers also identified the English words. Consistent with previous findings, both the deaf and the hearing subjects showed a left-hemisphere advantage to the English words; likewise, the deaf subjects showed a right hemisphere advantage to the statically presented signs. With the moving signs, the deaf showed no lateral asymmetry. The shift from right dominance to a more balanced hemispheric involvement with the change from static to moving signs is consistent with Kimura's position that the left hemisphere predominates in the analysis of skilled motor sequencing (Kimura 1976). The results also indicate that ASL may be more bilaterally represented than is English and that the spatial component of language stimuli can greatly influence lateral asymmetries.     

Individual differences in spatial relation processing: effects of strategy, ability, and gender

Numerous studies have focused on the distinction between categorical and coordinate spatial relations. Categorical relations are propositional and abstract, and often related to a left hemisphere advantage. Coordinate relations specify the metric information of the relative locations of objects, and can be linked to right hemisphere processing. Yet, not all studies have reported such a clear double dissociation; in particular the categorical left hemisphere advantage is not always reported. In the current study we investigated whether verbal and spatial strategies, verbal and spatial cognitive abilities, and gender could account for the discrepancies observed in hemispheric lateralization of spatial relations. Seventy-five participants performed two visual half field, match-to-sample tasks (Van der Ham, van Wezel, Oleksiak, & Postma, 2007; Van der Ham, Raemaekers, van Wezel, Oleksiak, and Postma, 2009) to study the lateralization of categorical and coordinate relation processing. For each participant we determined the strategy they used in each of the two tasks. Consistent with previous findings, we found an overall categorical left hemisphere advantage and coordinate right hemisphere advantage. The lateralization pattern was affected selectively by the degree to which participants used a spatial strategy and by none of the other variables (i.e., verbal strategy, cognitive abilities, and gender). Critically, the categorical left hemisphere advantage was observed only for participants that relied strongly on a spatial strategy. This result is another piece of evidence that categorical spatial relation processing relies on spatial and not verbal processes.     

Heart-Rate responses (HRR) to lateralized visual stimuli

Direction of changes in heart-rate responses (HRR) were investigated in three separate experiments as a measure of differential cognitive and emotional specialization of the cerebral hemispheres. Visual stimuli were presented via the visual half-field technique in all three experiments. Slides with different contents were flashed for 200 msec on each trial either to the left or right of a center LED fixation point. The LED went on 5 seconds prior to slide onset. HR changes were scored as second-by-second deviations during 10 seconds after LED onset from pre-LED base line. In the first experiment it was hypothesized that emotionally relevant stimuli initially projected to only the right hemisphere would result in more anticipatory acceleration than when the same stimulus was initially projected to the left hemisphere. A picture of a snake and of a geometric figure were repeatedly briefly flashed to the right of the LED for half of the subjects, and to the left for the other half. There were 25 trials with an intertriai interval of 25–40 seconds. Results showed significant effects of deceleration as a function of the slide stimulus in all groups on seconds 5, 6, or 7 after onset of the center LED. Furthermore, an anticipatory acceleration was observed during the first trial-block on seconds 3 and 4 in the right hemisphere groups only with no differences between the neutral and emotional stimuli. In Experiments 2 and 3, a letter-string of six letters and a complex symmetric pattern were used as stimuli. These stimuli were chosen because previous research has clearly implicated the hemispheres to be differentially specialized in their ability to process verbal and visuo-spatial stimuli. The set-up was identical to Experiment 1, with the exception that differences in response to the two types of stimuli were evaluated on a within-subjects basis. The results from Experiments 2 and 3 showed stimulus-related deceleration, peaking on seconds 5–7 in all groups and an anticipatory acceleration peaking on seconds 3 and 4 in the right hemisphere groups, with decelerations during the corresponding seconds in the left hemisphere groups. The results are discussed in relation to recent findings by Walker and Sandman (1982) about the possibility of hemispheric specialization in psychologic influences on heart rate changes in response to environmental demands.     

Egocentric metric representations in peripersonal space: A bridge between motor resources and spatial memory

Research on visuospatial memory has shown that egocentric (subject-to-object) and allocentric (object-to-object) reference frames are connected to categorical (non-metric) and coordinate (metric) spatial relations, and that motor resources are recruited especially when processing spatial information in peripersonal (within arm reaching) than extrapersonal (outside arm reaching) space. In order to perform our daily-life activities, these spatial components cooperate along a continuum from recognition-related (e.g., recognizing stimuli) to action-related (e.g., reaching stimuli) purposes. Therefore, it is possible that some types of spatial representations rely more on action/motor processes than others. Here, we explored the role of motor resources in the combinations of these visuospatial memory components. A motor interference paradigm was adopted in which participants had their arms bent behind their back or free during a spatial memory task. This task consisted in memorizing triads of objects and then verbally judging what was the object: (1) closest to/farthest from the participant (egocentric coordinate); (2) to the right/left of the participant (egocentric categorical); (3) closest to/farthest from a target object (allocentric coordinate); and (4) on the right/left of a target object (allocentric categorical). The triads appeared in participants' peripersonal (Experiment 1) or extrapersonal (Experiment 2) space. The results of Experiment 1 showed that motor interference selectively damaged egocentric-coordinate judgements but not the other spatial combinations. The results of Experiment 2 showed that the interference effect disappeared when the objects were in the extrapersonal space. A third follow-up study using a within-subject design confirmed the overall pattern of results. Our findings provide evidence that motor resources play an important role in the combination of coordinate spatial relations and egocentric representations in peripersonal space.     

When phonology fails: orthographic neighbourhood effects in dyslexia

Both cerebral hemispheres contain phonological, orthographic and semantic representations of words, however there are between-hemisphere differences in the relative engagement and specialization of the different representations. Taking orthographic processing for example, previous studies suggest that orthographic neighbourhood size (N) has facilitatory effects in the right but not the left hemispheres. To pursue the nature of this asymmetric N effect, in particular whether there are individual differences in such specialisation, we examined N in a case of developmental dyslexia, FM. We first describe the nature of his difficulties, which are mainly severe phonological deficits. Employing the divided visual field paradigm with FM revealed a greater sensitivity in the right than in the left hemisphere to orthographic variables, with a significant inhibitory N effect in the left, but not right hemisphere. Such inhibition, to a lesser degree, was found among a group of adults with dyslexia but not among age-matched normal readers. We argue that enhanced sensitivity to orthographic cues is developed in some cases of dyslexia when a normal, phonology-based left hemisphere word recognition processing is not achieved. The interpretation presented here is cast in terms of differences between people with dyslexia and typical readers that originate in the atypical way in which orthographic representations are initially set up.     

Lateralization of lexical codes in auditory word recognition

Three experiments examined the lateralization of lexical codes in auditory word recognition. In Experiment 1 a word rhyming with a binaurally presented cue word was detected faster when the cue and target were spelled similarly than when they were spelled differently. This orthography effect was larger when the target was presented to the right ear than when it was presented to the left ear. Experiment 2 replicated the interaction between ear of presentation and orthography effect when the cue and target were spoken in different voices. In Experiment 3, subjects made lexical decisions to pairs of stimuli presented to the left or the right ear. Lexical decision times and the amount of facilitation which obtained when the target stimuli were semantically related words did not differ as a function of ear of presentation. The results suggest that the semantic, phonological, and orthographic codes for a word are represented in each hemisphere; however, orthographic and phonological representations are integrated only in the left hemisphere.     

Hemispheric asymmetry and electrodermal activity in orienting and pavlovian conditioning paradigms

Three experiments on lateralization of electrodermal orienting and conditioned behavior are reported. The basic findings show effects of hemispheric asymmetry on initial magnitude and rate of habituation of phasic stimulus-elicited electrodermal responses. In summary, responses to verbal stimuli repeatedly flashed to the right visual half-field are larger than corresponding responses in the left half-field. Similarily, spatially relevant stimuli initially presented to the left half-field result in larger responses than when the same stimuli are repeatedly presented in the right half-field. In the auditory modality, data show asymmetrical control of responding to verbal material in a dichotic conditioning paradigm with greater resistance to extinction when the conditioned stimulus is initially fed only to the left hemisphere as compared to when it is fed only to the right hemisphere. It is argued that the present approach taps on basic mechanisms for the lateralization of attentional and associative functions.     

Recognition of letters traced in the right and left palms: Evidence for a process-oriented tactile asymmetry

Three experiments were conducted to examine the relative ability of the cerebral hemispheres to identify capital letters traced in the palms of the hands. In Experiment 1, letters were presented either right side up or upside down, and the subject's task was to name the letter aloud or point to an identical letter using the stimulated hand. Analysis of the accuracy data revealed that the left palm/right hemisphere (LP/RH) performed this task significantly better than did the right palm/left hemisphere (RP/LH), particularly when the stimuli were presented in the upside-down orientation. In Experiments 2 and 3, subjects performed the same letter identification task; however, on half the trials, they were required to maintain either a spatial or verbal concurrent memory load (i.e., a 24-point Vanderplas & Garvin form or six low-imagery nouns, respectively). In the no-load condition of Experiment 2 (spatial forms), the previously observed LP/RH advantage was replicated. However, in the load condition, this LP/RH superiority was no longer in evidence. In Experiment 3 (low-imagery nouns), the presence of a concurrent verbal task had minimal impact on the previously observed performance asymmetry as the LP/RH advantage was obtained in both the no-load and load conditions. The results of the three studies taken in composite suggest that (1) the operations utilized to identify letters traced in the palms of the hands are primarily spatial in nature and (2) that the observed performance asymmetry may be attributed to a right hemisphere superiority for the analysis and codification of information along a spatial dimension. These findings are discussed in terms of a "process-oriented" model of hemispheric asymmetry.     

HEMISPHERIC DIFFERENCES IN AUDITORY PERCEPTION ARE SIMILAR TO THOSE FOUND IN VISUAL PERCEPTION

Abstract— In a pitch discrimination task, subjects were faster and more accurate in judging low-frequency sounds when these stimuli were presented to the left ear, compared with the right ear. In contrast, a right-ear advantage was found with high-frequency sounds. The effect was in terms of relative frequency and not absolute frequency, suggesting that the effect arisen from pastsensory mechanisms. A simitar laterality effect has been reported in visual perception with stimuli varying in spatial frequency. These multimodal laterality effects may reflect a general computational difference between the two cerebral hemispheres, with the left hemisphere biased for processing high-frequency information and the right hemisphere biased for processing low-frequency information.     

Differential visual field-interhemispheric transfer: can it explain sex and handedness differences in lateralization?

104 men and women were tested for visual field-hemispheric transfer of spatial information on a dot-localization task. Right-handed subjects showed significant improvement when stimuli were presented to the left visual field of the right hemisphere (LVF-RH) after practice on the same task presented to the right visual field of the left hemisphere (RVF-LH) first. No improvement was found when the task was presented in the reverse order (LVF-RH first followed by RVF-LH). It was concluded that, for right-handers, transfer of spatial information to the right hemisphere is facilitated while transfer to the left hemisphere is inhibited. Left-handed subjects demonstrated no significant improvement in either condition, suggesting inhibition or lack of transfer of spatial information in either direction. No sex differences were found in either right-handed or left-handed subjects. The findings suggest that there may be different mechanisms underlying the similarities in functional lateralization of women and left-handers.     

Hemispheric Differences in the Recognition of Environmental Sounds

ABSTRACT— Recent work has found support for two dissociable and parallel neural subsystems underlying object and shape recognition in the visual domain: an abstract-category subsystem that operates more effectively in the left cerebral hemisphere than in the right, and a specific-exemplar subsystem that operates more effectively in the right hemisphere than in the left. Evidence of this asymmetry has been observed for linguistic stimuli (words, pseudoword forms) and nonlinguistic stimuli (objects). In the auditory domain, we previously found hemispheric asymmetries in priming effects using linguistic stimuli (spoken words). In the present study, we conducted four long-term repetition-priming experiments to investigate whether such hemispheric asymmetries would be observed for nonlinguistic auditory stimuli (environmental sounds) as well. The results support the dissociable-subsystems theory. Specificity effects were obtained when sounds were presented to the left ear (right hemisphere), but not when sounds were presented to the right ear (left hemisphere). Theoretical implications are discussed.     

Hemisphere-dependent holistic processing of familiar faces

In two behavioral experiments involving lateralized stimulus presentation, we tested whether one of the most commonly used measures of holistic face processing—the composite face effect—would be more pronounced for stimuli presented to the right as compared to the left hemisphere. In experiment 1, we investigated the composite face effect in a verbal identification task, similar to its original report (Young, Hellawell, & Hay, 1987). Aligning top and bottom halves of composite face stimuli led to performance decreases irrespective of hemifield, indicating holistic processing of comparable magnitude for inputs provided separately to either hemisphere. However, when matching of the same top parts was required in experiment 2, an alignment-dependent performance decrease was found for stimuli presented in the left, but not right visual field. These observations suggest that the right hemisphere dominates in early stages of holistic processing, as indexed by the composite face effect, but that later processes such as face identification and naming are based on unified representations that are independent of input lateralization. Moreover, the composite face effect may not rely on the exact same representation(s) when measured in matching and identification tasks.