How does interhemispheric communication in visual word recognition work? Deciding between early and late integration accounts of the split fovea theory

It has recently been shown that interhemispheric communication is needed for the processing of foveally presented words. In this study, we examine whether the integration of information happens at an early stage, before word recognition proper starts, or whether the integration is part of the recognition process itself. Two lexical decision experiments are reported in which words were presented at different fixation positions. In Experiment 1, a masked form priming task was used with primes that had two adjacent letters transposed. The results showed that although the fixation position had a substantial influence on the transposed letter priming effect, the priming was not smaller when the transposed letters were sent to different hemispheres than when they were projected to the same hemisphere. In Experiment 2, stimuli were presented that either had high frequency hemifield competitors or could be identified unambiguously on the basis of the information in one hemifield. Again, the lexical decision times did not vary as a function of hemifield competitors. These results are consistent with the early integration account, as presented in the SERIOL model of visual word recognition.     

Hemispheric refractoriness and control of reaction time

Experiments are described which attempt to assess the relative efficiency of the hemispheres and their relationship in performance on complex RT tasks. A divided visual field method was used to direct signals to the temporal or nasal retinae of each eye thus passing information to separate hemispheres. A comparison of the separate response times was used to assess the relative efficiency of each hemisphere but significant differences were not observed. This suggests that each may be the equal of the other in organizing simple responses. A method was used to examine more complex RT by presenting the subject with two simultaneous signals for response. When pairs of signals are directed to separate hemispheres, response times are at their lowest value. When signals are directed to separate hemispheres through the same eye, a significant increase in RT occurs. A source of mutual interference appears to exist at the level of the eye. Response times are extended to their greatest value, however, when both signals are directed to the same hemisphere. This block to function has been described as “hemispheric refractoriness”, and is different for the two hemispheres. While each show a distinct block to function the extent of this is greater in the right or minor hemisphere than it is in the left or major hemisphere.     

Hemispheric differences for the integration of stimulus levels and their contents: evidence from bilateral presentations

Several studies have demonstrated that hemispheric differences for the processing of hierarchical letter stimuli are more likely to occur when the letters at the levels are associated with conflicting responses. Typically, a single stimulus is presented, so that the conflict occurs between the global and the local levels of the same stimulus. Our hypothesis is that in this situation, conflict resolution requires integration of the letters and their respective levels and that the hemispheres differ in this integration process. According to this integration theory, the favorable effect of response conflict on hemispheric differences should vanish if other features, such as location, can also serve for conflict resolution. This prediction was tested in the present study by simultaneously presenting an individual hierarchical stimulus to each visual field. Conflicting letters either were arranged within one stimulus or were placed in different stimuli. In the latter case, a response conflict could also be resolved by integrating letters and locations. As was expected, there were no visual field effects in these conditions. On the other hand, visual field effects showed up when the conflicting letters were located within the same stimulus. These results support the idea that the hemispheres differ in their capacity for integrating level and form.     

The cerebral balance of power: confrontation or cooperation?

Two visual search experiments were carried out using as stimuli large letters made of small identical letters presented in right, or left, or central visual fields. Considering the spatial frequency contents of the stimuli as the critical variable, Experiment 1 showed that a left-field superiority could be obtained whenever a decision had to be made on a large (low frequency) letter alone, and a right-field advantage emerged when a small (high frequency) letter had to be processed. Experiment 2 showed that the two levels of structure of the stimulus were not encoded at the same rate and that at very brief exposure, only the large letter could be accurately identified. This was accompanied by a left-field superiority, whether or not the stimulus contained the target. These results are interpreted as revealing a differential sensitivity of the hemispheres to the spatial frequency contents of a visual image, the right hemisphere being more adept at processing early-available low frequencies and the left hemisphere operating more efficiently on later-available low frequencies. From these and other experiments reviewed, it is suggested that (a) cerebral lateralization of cognitive functions results from differences in sensorimotor resolution capacities of the hemispheres; (b) both hemispheres can process verbal and visuospatial information, analytically and holistically; (c) respective hemispheric competence is a function of the level of sensorimotor resolution required for processing the information available.     

Some evidence for a selfterminating process in simple visual search tasks

Two experiments were conducted to investigate the unsettled question of whether the visual search process is selfterminating or exhaustive. In the experiments three letters were placed on an imaginary circle round the fixation point. Two different letters were used, one of which was defined as the ‘signal’. Ss had to respond ‘yes’ when one or more signals were in the display, ‘no’ otherwise. In both experiments the number of signals in the display was varied from 0 to 3.Decreasing latencies with increasing number of signals were observed in both experiments, indicating a selfterminating visual search. In experiment 1 a significant increase in latencies with increasing visual angle was found, in experiment 2 an increase in latency resulting from neighbouring contours. Both factors probably contribute to the slope of the function relating positive responses to display size and to the slope of the function relating negative responses to display size when stimuli are presented in linear arrays as in the experiment by Atkinson et al. (1969). As a result the slopes become more equal, falsely suggesting an exhaustive visual search process.     

Some evidence for a limited capacity parallel selfterminating process in simple visual search tasks.

An experiment was carried out in order to investigate the questions of exhaustive vs selfterminating and serial vs parallel processing in a simple visual search task. In the experiment, 1, 2 or 3 letters were placed on an imaginary circle round the fixation point. Two different letters were used, one of which was defined as the ‘signal’. Ss had to respond ‘yes’ when one or more signals were in the display, ‘no’ otherwise. The number of signals in the display was varied from ‘no signals’ to ‘all signals’.A decrease in latencies with increasing number of signals for a fixed number of elements presented was observed indicating a selfterminating search. A decrease in latencies with increasing number of elements when only signals were presented was taken as evidence for a parallel selfterminating process. A further analysis of the error data showed that a limited capacity system had to be assumed. It was shown that it was possible to construct the overall pattern of latencies over conditions from the error data obtained.     

Effects of visual-field and matching instruction on event-related potentials and reaction time

Vertical letter pairs were presented randomly in the left and right visual hemifields in a physical identity match and name identity match condition. The reaction times showed a right visual field superiority for name matches, and a left visual field superiority for physical matches. Event-related potentials to letter pairs showed a sequence of three waves: a negative wave (N2, around 270 msec), a positive wave (P3, around 500 msec), and a broad positive slow wave (SW, around 600-700 msec), respectively. P3 and SW amplitudes were consistently larger at the left hemisphere than at the right hemisphere, regardless of the field of stimulation. At both hemispheres, N2 waves were always larger to stimuli presented in the visual field contralateral to a hemisphere than stimuli presented in the visual field ipsilateral to a hemisphere. The positive waves (P3, SW) showed the opposite pattern: smaller amplitudes to stimuli that were presented contralaterally than stimuli that were presented ipsilaterally to a given hemisphere. These results were attributed to a shift in sustained negativity on the directly stimulated hemisphere, relative to the indirectly stimulated hemisphere, reflecting either sensory at attentional processes in the posterior cerebral hemispheres.     

Reaction times and evoked potentials as indicators of hemispheric differences for laterally presented name and physical matches

Letter pairs, which could be name matches, physical matches, or mismatches, were presented at fixation or 2.5 degrees left or right of fixation. During different experimental sessions, the locations and the types of matches were fixed (and therefore known in advance by the subject) or were randomized. Right visual field superiority in reaction time occurred for name matches only when location was randomized, and then the extent of the superiority depended on whether the types of match called for were predictable. Evoked potentials to the letter pairs during this task revealed hemispheric and neural pathway differences that were independent of expectancy condition. Right hemisphere responses were larger than left. For some components, amplitudes were smaller and latencies were shorter for direct than for indirect projection of stimuli to each cerebral hemisphere. Indirect-direct differences in P300 amplitude varied for each cerebral hemisphere according to whether a physical or name match occurred. The P130 and N170 components manifested hemispheric differences that depended on whether the two letters of a pair were in the same or different cases.     

Transposed-letter and laterality effects in lexical decision

Two divided visual field lexical decision experiments were conducted to examine the role of the cerebral hemispheres in transposed-letter similarity effects. In Experiment 1, we created two types of nonwords: nonadjacent transposed-letter nonwords (TRADEGIA; the base word was TRAGEDIA, the Spanish for TRAGEDY) and two-letter different nonwords (orthographic controls: TRATEPIA). In Experiment 2, the controls were one-letter different nonwords (TRAGEPIA) instead of two-letter different nonwords (TRATEPIA). The effect of transposed-letter similarity was substantially greater in the right visual field (left hemisphere) than in the left visual field. Furthermore, nonwords created by transposing two letters were more competitive than the nonwords created by substituting one or two letters of a target word. We examine the implications of these findings for the models of visual word recognition.     

Lateralized repetition priming for familiar faces: Evidence for asymmetric interhemispheric cooperation

Repetition priming refers to facilitated recognition of stimuli that have been seen previously. Although a great deal of work has examined the properties of repetition priming for familiar faces, little has examined the neuroanatomical basis of the effect. Two experiments are presented in this paper that combine the repetition priming paradigm with a divided visual field methodology to examine lateralized recognition of familiar faces. In the first experiment participants were presented with prime faces unilaterally to each visual field and target faces foveally. A significant priming effect was found for prime faces presented to the right hemisphere, but not for prime faces presented to the left hemisphere. In Experiment 2, prime and target faces were presented unilaterally, either to the same visual field or to the opposite visual field (i.e., either within hemisphere or across hemispheres). A significant priming effect was found for the within right hemisphere condition, but not for the within left hemisphere condition, replicating the findings of the first experiment. Priming was also found in both of the across hemispheres conditions, suggesting that interhemispheric cooperation occurs to aid recognition. Taken in combination these experiments provide two main findings. First, an asymmetric repetition priming effect was found, possibly as a result of asymmetric levels of activation following recognition of a prime face, with greater priming occurring within the right hemisphere. Second, there is evidence for asymmetric interhemispheric cooperation with transfer of information from the right hemisphere to the left hemisphere to facilitate recognition.     

Lateralized repetition priming for familiar faces: Evidence for asymmetric interhemispheric cooperation

Repetition priming refers to facilitated recognition of stimuli that have been seen previously. Although a great deal of work has examined the properties of repetition priming for familiar faces, little has examined the neuroanatomical basis of the effect. Two experiments are presented in this paper that combine the repetition priming paradigm with a divided visual field methodology to examine lateralized recognition of familiar faces. In the first experiment participants were presented with prime faces unilaterally to each visual field and target faces foveally. A significant priming effect was found for prime faces presented to the right hemisphere, but not for prime faces presented to the left hemisphere. In Experiment 2, prime and target faces were presented unilaterally, either to the same visual field or to the opposite visual field (i.e., either within hemisphere or across hemispheres). A significant priming effect was found for the within right hemisphere condition, but not for the within left hemisphere condition, replicating the findings of the first experiment. Priming was also found in both of the across hemispheres conditions, suggesting that interhemispheric cooperation occurs to aid recognition. Taken in combination these experiments provide two main findings. First, an asymmetric repetition priming effect was found, possibly as a result of asymmetric levels of activation following recognition of a prime face, with greater priming occurring within the right hemisphere. Second, there is evidence for asymmetric interhemispheric cooperation with transfer of information from the right hemisphere to the left hemisphere to facilitate recognition.     

Effects of concurrent spatial and verbal memory loads on serial position functions of laterally presented letter strings

In this study the influence of visuospatial and verbal memory loads on the identification of laterally presented letter strings was investigated. In the asymmetry task without concurrent loads, a clear right visual field advantage for letter identification was obtained. Hemisphere-specific effects due to concurrent loads were particularly observed when the difficulty of the load tasks was increased. The effects of visuospatial loads were found to be sex related, suggesting that under heavy load conditions mental rotation selectively overloads the processing capacity of the right hemisphere in males, while in females capacity limitations were observed in both hemispheres. Concurrent visuospatial loads produced more facilitation (or less interference) for letters in the outermost positions of each visual field than for letters in the innermost positions of each visual field. The results of the verbal memory load tasks revealed that an easy verbal load task facilitated performance which was particularly manifest for the right-most letter of both the left visual field and the right visual field. A difficult verbal memory load task interfered with recognition accuracy of letters which was most marked for the center letter in the right visual field. Letter position effects obtained in this study were interpreted in terms of various processing mechanisms influencing the serial position curves.     

Hemispheric processing of anaphoric inferences: the activation of multiple antecedents

This research investigates the hemispheric processing of anaphors when readers activate multiple antecedents. Participants read texts promoting an anaphoric inference and performed a lexical decision task to inference-related target words that were consistent (Experiment 1) or inconsistent (Experiment 2) with the text. These targets were preceded by constrained or less constraining text and were presented to participants' right visual field-left hemisphere or to their left visual field-right hemisphere. In Experiment 1, both hemispheres showed facilitation for consistent antecedents and the right hemisphere showed an advantage over the left hemisphere in processing antecedents when preceded by less constrained text. In Experiment 2, the left hemisphere only showed negative facilitation for inconsistent antecedents. When readers comprehend text with multiple antecedents: both hemispheres process consistent information, the left hemisphere inhibits inconsistent information, and the right hemisphere processes less constrained information.     

Length, formats, neighbours, hemispheres, and the processing of words presented laterally or at fixation

It has long been known that the number of letters in a word has more of an effect on recognition speed and accuracy in the left visual field (LVF) than in the right visual field (RVF) provided that the word is presented in a standard, horizontal format. After considering the basis of the length by visual field interaction two further differences between the visual fields/hemispheres are discussed: (a) the greater impact of format distortion (including case alternation) in the RVF than in the LVF and (b) the greater facilitation of lexical decision by orthographic neighbourhood size (N) in the LVF than in the RVF. In the context of split fovea accounts of word recognition, evidence is summarised which indicates that the processing of words presented at fixation is affected by the number of letters to the left of fixation but not by the number of letters to the right and by the number of orthographic neighbours activated by letters to the left of fixation but not by the number of orthographic neighbours activated by letters to the right of fixation. A model of word recognition is presented which incorporates the notion that the left hemisphere has sole access to a mode of word recognition that involves parallel access from letter forms to the visual input lexicon, is disrupted by format distortion, and does not employ top-down support of the letter level by the word level.     

Hemispheric processing differences revealed by differential conditioning and reaction time performance.

Two different experimental procedures were used to examine (a) information-processing differences between two groups of subjects (Cs versus Vs) identified by the form of their conditioned eyeblinks; (b) information-processing differences between the right and left cerebral hemispheres; and (c) parallels between hypothesized C-V differences and right-left hemisphere differences. In the first experiment, the evocative command words BLINK and DON'T BLINK served as positive and negative conditioned stimuli. It was found that Vs gave more conditioned eyeblinks than Cs and that differential eyelid conditioning of Vs more than Cs was influenced by the semantic content of the stimuli. More importantly, the conditioning performance of Cs was more influenced by the semantic attributes of the stimuli when they were presented directly to the right visual field (left hemisphere) than when they were presented directly to the left visual field (right hemisphere). In contrast, the conditioning performance of Vs was equally influenced by the semantic attributes regardless of which hemisphere received direct stimulation. A second experiment was designed to determine whether such hemisphere-of-presentation differences for Cs versus Vs could also be obtained in a very different task. Subjects classified as Cs or Vs during a differential eyelid conditioning task then performed two same-different reaction time (RT) tasks that required discrimination of complex polygons in one case and the names of letters in another. On each RT trial both stimuli of a pair appeared briefly either in the center, left, or right visual field. For both Cs and Vs RTs to complex polygon pairs averaged 20 msec faster on left visual field trials than on right visual field trials, consistent with current hypotheses about right-hemisphere specialization for visuospatial processing. In contrast, the results for letter pairs generally confirmed the C-V differences found in Experiment 1. That is, the right visual field (left-hemisphere) advantage for these verbal stimuli was once again larger for Cs than for Vs. The present results suggest that the two groups of subjects (Cs versus Vs) differ qualitatively in the modes of information processing that they typically employ. The results also suggest that these different modes of processing are related to aspects of cerebral hemisphere organization and that even right-handed individuals may differ from each other in the extent to which each cerebral hemisphere is mobilized for a given experimental task. Such individual differences must be incorporated into both models of classical eyelid conditioning and models of cerebral hemisphere specialization.     

Hemispheric differences for feature perception

Summary Hemisphere differences for featural processing were investigated in three experiments. Stimulus arrays composed of the same background letter were presented tachistoscopically and the subject instructed to detect an embedded target letter. Background and target letter features were manipulated across studies, while stimulus arrays were presented at different retinal loci within experiments. Signal detection analysis revealed that left hemisphere stimulus presentations demonstrated slightly better detection when target and background features were relatively dissimilar, while right hemisphere stimulus presentations demonstrated better detection when target and background features were highly similar. Retinal locus generally decreased detection performance when the stimulus letters were dissimilar and interacted with hemispheric advantages. Both of these factors also were affected by changes in response criterion across experiments which were linked to target/background perceptual confusions. The findings suggest that the left and right hemispheres differ in their feature extraction capabilities during the early stages of visual stimulus processing.     

Form-Specific Visual Priming in the Left and Right Hemispheres

Word fragment completion performance was examined for items that were presented in the same or different letter case at study and test. During the study phase words and nonwords were presented at central fixation, then during the test phase a divided visual field technique was used in which word fragments were presented briefly to the right hemisphere (left visual field) or the left hemisphere (right visual field). Previous research using the word stem completion task indicated that only the right hemisphere was sensitive to case changes in words from study to test. In contrast, the current results indicate that in the fragment completion task the priming effects for the test items presented to either hemisphere were greater when the fragments were in the same compared to different letter case at study and test. These results indicate that both hemispheres are capable of supporting form-specific visual implicit memory.     

Reaction time to letter name or letter case

Subjects were shown two letters from the set BDGbdg. In one condition, subjects were required to decide whether or not the letters had the same name. In the other condition, subjects decided whether or not the two letters were presented in the same case. Reaction times were always faster when the two letters on a given trial were physically identical. However, there was no difference in the speed of a name match or a case match when the two letters were not physically identical. Since subjects could have based a case match on the presence or absence of a single feature — a protruding vertical line — it was concluded that subjects are not able to selectively attend to a single visual feature in order to identify a letter. Rather, a subject analyzes several features in parallel in order to arrive at a simultaneous decision about a letter's name or case.     

Spatial frequencies and the cerebral hemispheres: contrast sensitivity, visible persistence, and letter classification

The hypothesis that the two cerebral hemispheres are specialized for processing different visual spatial frequencies was investigated in three experiments. No differences between the left and right visual fields were found for: (1) contrast-sensitivity functions measured binocularly with vertical gratings ranging from 0.5 to 12 cycles per degree (cpd); (2) visible persistence durations for 1- and 10-cpd gratings measured with a stimulus alternation method; and (3) accuracy (d') and reaction times to correctly identify digitally filtered letters as targets (L or H) or nontargets (T or F). One significant difference, however, was found: In Experiment 3, a higher decision criterion (beta) was used when filtered letters were identified in the right visual field than when they were identified in the left. The letters were filtered with annular, 1-octave band-pass filters with center spatial frequencies of 1, 2, 4, 8, and 16 cpd. Combining four center frequencies with three letter sizes (0.5 degrees, 1 degree, and 2 degrees high) made some stimuli equivalent in distal spatial frequency (cycles per object) and some equivalent in proximal spatial frequency (cycles per degree). The effective stimulus in the third experiment seemed to be proximal spatial frequency (cycles per degree) not distal (cycles per object). We conclude that each cerebral hemisphere processes visual spatial frequency information with equal accuracy but that different decision rules are used.