On hemispheric specialisation and visual field effects in the perception of print: A comment on Jordan, Patching, and Thomas

Introduction Due to structural characteristics of the visual pathways, stimuli that are presented in the right half of the visual field (RVF) are initially projected to the left cerebral hemisphere, while those presented in the left half of the visual field (LVF) are projected to the right cerebral hemisphere. This anatomical feature has frequently been taken to support the notion that the well-documented RVF advantage in recognising printed words is a reflection of functional differences between the two hemispheres; notably that of the dominance of the left hemisphere for processing language. Word stimuli that are sent straight to the left hemisphere are believed to profit from more efficient processing than those sent initially to the right hemisphere, because the latter stimuli must follow a longer and more noisy pathway before reaching the language-dominant hemisphere. In the work by Jordan, Patching, and Thomas (2003) the above notion is further developed to speculate that the point of entry of visual information into the cortex may determine the procedure that will underlie the ensuing word recognition process: "... the left hemisphere can process words by mapping orthographic information in parallel onto lexical entries whereas the right hemisphere has a more rudimentary process, that can only map orthographic information sequentially" (p. 50).     

Assessing the Role of Hemispheric Specialisation, Serial-Position Processing, and Retinal Eccentricity in Lateralised Word RecognitioN

The advantage for words in the right visual hemifield (RVF) has been assigned parallel orthographic processing by the left hemisphere and sequential by the right. However, an examination of previous studies of serial position performance suggests that orthographic processing in each hemifield is modulated by retinal eccentricity. To investigate this issue, we presented words at eccentricities of 1, 2, 3, and 4 degrees. Serial position performance was measured using the Reicher-Wheeler task to suppress influences of guesswork and an eye-tracker controlled fixation location. Greater eccentricities produced lower overall levels of performance in each hemifield although RVF advantages for words obtained at each eccentricity (Experiments 1 and 2). However, performance in both hemifields revealed similar U-shaped serial position performance at all eccentricities. Moreover, this performance was not influenced by lexical constraint (high, low; Experiment 2) or status (word, nonword; Experiment 3), although only words (not nonwords) produced an RVF advantage. These findings suggest that although each RVF advantage was produced by left-hemisphere function, the same pattern of orthographic analysis was used by each hemisphere at each eccentricity.     

Asymmetries and Eccentricities in Studies of Lateralised Word Recognition: A Response to Nazir

The anatomical arrangement of the human visual system offers considerable scope for investigating functional asymmetries in hemispheric processing. In particular, because each hemisphere receives information initially from the contralateral visual hemifield, visual stimuli presented to the left of a central fixation point can be projected directly to the right hemisphere and visual stimuli presented to the right of a central fixation point can be projected directly to the left hemisphere. Numerous studies using displays of this type suggest that, for the vast majority of individuals, written words produce different patterns of performance when presented to different hemifields and these findings have inspired considerable debate about the processes available for word recognition in each hemisphere.     

Asymmetries and eccentricities in studies of lateralised word recognition: a response to nazir

The anatomical arrangement of the human visual system offers considerable scope for investigating functional asymmetries in hemispheric processing. In particular, because each hemisphere receives information initially from the contralateral visual hemifield, visual stimuli presented to the left of a central fixation point can be projected directly to the right hemisphere and visual stimuli presented to the right of a central fixation point can be projected directly to the left hemisphere. Numerous studies using displays of this type suggest that, for the vast majority of individuals, written words produce different patterns of performance when presented to different hemifields and these findings have inspired considerable debate about the processes available for word recognition in each hemisphere.     

Hemispheric differences for identification of words and nonwords in Urdu-English bilinguals

Hemispheric asymmetry was examined for Urdu-English bilinguals identifying printed Urdu words and nonwords, separated Urdu letter strings, digits, and English nonwords. In all cases, fewer errors occurred when stimuli were presented to the right visual field/left hemisphere (RVF/LH) than to the left visual field/right hemisphere (LVF/RH). Qualitative error patterns suggested that separated Urdu letter strings were processed more serially than Urdu letter strings joined to form words or pronounceable nonwords and more serially on RVF/LH than on LVF/RH trials. This qualitative laterality effect is similar to that found for Hebrew and Arabic but opposite that found for English and suggests that the qualitative manner of processing printed verbal material is influenced by language-specific factors such as scanning direction, orthographic-to-phonological mapping rules, and morphology.     

Word position affects stimulus recognition: Evidence for early ERP short-term plastic modulation

The present study was aimed at investigating the short-term plastic changes that follow word learning at a neurophysiological level. The main hypothesis was that word position (left or right visual field, LVF/RH or RVF/LH) in the initial learning phase would leave a trace that affected, in the subsequent recognition phase, the Recognition Potential (i.e., the first negative component distinguishing words from other stimuli) elicited 220–240 ms after centrally presented stimuli. Forty-eight students were administered, in the learning phase, 125 words for 4 s, randomly presented half in the left and half in the right visual field. In the recognition phase, participants were split into two equal groups, one was assigned to the Word task, the other to the Picture task (in which half of the 125 pictures were new, and half matched prior studied words). During the Word task, old RVF/LH words elicited significantly greater negativity in left posterior sites with respect to old LVF/RH words, which in turn showed the same pattern of activation evoked by new words. Therefore, correspondence between stimulus spatial position and hemisphere specialized in automatic word recognition created a robust prime for subsequent recognition. During the Picture task, pictures matching old RVF/LH words showed no differences compared with new pictures, but evoked significantly greater negativity than pictures matching old LVF/RH words. Thus, the priming effect vanished when the task required a switch from visual analysis to stored linguistic information, whereas the lack of correspondence between stimulus position and network specialized in automatic word recognition (i.e., when words were presented to the LVF/RH) revealed the implicit costs for recognition. Results support the view that short-term plastic changes occurring in a linguistic learning task interact with both stimulus position and modality (written word vs. picture representation).     

Efficiency of the forebrain commissures: memory for stimuli seen by the other hemisphere

How well will one cerebral hemisphere recognize items viewed initially via the other? Nonverbalizable images or words were presented to one visual field and memory for them tested in the same or the other visual field. The initially viewing hemisphere subsequently had no secure advantage in accuracy, and only for images was there a 30-ms (ca 3%) penalty in reaction time for viewing with the "other" hemisphere. Interhemispheric mnemonic communication is thus highly reliable. At longer retention intervals (1-2 min vs. 4-30 s, with accumulating added stimuli), however, recognition of words was asymmetric as to hemisphere, in that initial viewing via the right hemisphere was subsequently (and paradoxically) much better recognized via the left (other) hemisphere than was the converse situation. This suggests that the initial engram with right hemispheric viewing of words ultimately becomes established in the left, and that the right has less accurate access to a previous "left hemispheric view".     

PERCEPTUA.ND LEXICA.ACTORS IN A CAS.F LETTER-BY-LETTER READING

We report the case of a letter-by-letter reader (MJ) who showed normal processing of single letters and who could normally access the orthographic input lexicon when presented with letter names for aural recognition, or when allow ed enough tim e to process a visually presented letter string. However, MJ showed severe difficulties in simultaneously processing multiple letters and other simple visual stimuli. Furthermore, she does not have normal access to lexical orthographic representations and their meanings when stimuli are presented for too brief a time to allow for serial processing of the letter string. We found no evidence of (partial) lexical or semantic access without corresponding recognition of the letters in a word: No signs of implicit reading were observed w hen the input stimuli were controlled for the relevant visual features; “implicit reading” was only obtained under conditions that allowed sophisticated guessing. This pattern of results is interpreted as indicating that LBL reading (at least in MJ) results from damage to prelexical processing mechanisms. In MJ's case, the deficit reflects the degraded transfer of information from a normal visual processing system in the right hemisphere to a normal language processing system in the left hemisphere.     

Chlorpromazine and the lateralisation of the perception of emotion.

In a double-blind design, sixteen volunteer students were administered 50 mg of chlorpromazine or a placebo in tablet form on separate occasions, two hours prior to testing. The test was a lexical decision paradigm involving unilateral presentation of pairs of neutral, positive and negative emotional words and nonwords to the left or right visual fields. Reaction time to identify words and nonwords was recorded. Neutral stimuli were processed faster in the left hemisphere, while negative stimuli were processed faster in the right hemisphere. Chlorpromazine improved speed of response for neutral stimuli presented to the right hemisphere and for affective stimuli presented to the left hemisphere. Thus chlorpromazine seems to benefit right hemisphere processing of nonaffective stimuli and benefit left hemisphere processing of affective stimuli. Chlorpromazine appeared to have a different impact on neutral and affective words, and on the right and the left hemispheres. The general effect of chlorpromazine was to reduce lateralisation. It was suggested that this occurred because chlorpromazine blocked dopamine and possibly other neurotransmitters.     

Grammatical number agreement processing using the visual half-field paradigm: An event-related brain potential study

Despite indications in the split-brain and lesion literatures that the right hemisphere is capable of some syntactic analysis, few studies have investigated right hemisphere contributions to syntactic processing in people with intact brains. Here we used the visual half-field paradigm in healthy adults to examine each hemisphere's processing of correct and incorrect grammatical number agreement marked either lexically, e.g., antecedent/reflexive pronoun (“The grateful niece asked herself/*themselves…”) or morphologically, e.g., subject/verb (“Industrial scientists develop/*develops…”). For reflexives, response times and accuracy of grammaticality decisions suggested similar processing regardless of visual field of presentation. In the subject/verb condition, we observed similar response times and accuracies for central and right visual field (RVF) presentations. For left visual field (LVF) presentation, response times were longer and accuracy rates were reduced relative to RVF presentation. An event-related brain potential (ERP) study using the same materials revealed similar ERP responses to the reflexive pronouns in the two visual fields, but very different ERP effects to the subject/verb violations. For lexically marked violations on reflexives, P600 was elicited by stimuli in both the LVF and RVF; for morphologically marked violations on verbs, P600 was elicited only by RVF stimuli. These data suggest that both hemispheres can process lexically marked pronoun agreement violations, and do so in a similar fashion. Morphologically marked subject/verb agreement errors, however, showed a distinct LH advantage.     

Cross-language tests of hemispheric strategies in reading nonwords.

Four experiments explored the effects of specific language characteristics on hemispheric functioning in reading nonwords using a lateralized trigram identification task. Previous research using nonsense consonant-vowel-consonant (CVC) trigrams has shown that total error scores reveal a right visual field (RVF) advantage in Hebrew, Japanese, and English. Qualitative error patterns have shown that the right hemisphere uses a sequential strategy, whereas the left hemisphere uses a more parallel strategy in English but shows the opposite pattern in Hebrew. Experiment 1 tested whether this is due to the test language or to the native language of the participants. Results showed that native language had a stronger effect on hemispheric strategies than test language. Experiment 2 showed that latency to target letters in the CVCs revealed the same asymmetry as qualitative errors for Hebrew speakers but not for English speakers and that exposure duration of the stimuli affected misses differentially according to letter position. Experiment 3 used number trigrams to equate reading conventions in the 2 languages. Qualitative error scores still revealed opposing asymmetry patterns. Experiments 1-3 used vertical presentations. Experiment 4 used horizontal presentation, which eliminated sequential processing in both hemispheres in Hebrew speakers, whereas English speakers still showed sequential processing in both hemispheres. Comparison of the 2 presentations suggests that stimulus arrangement affected qualitative errors in the left visual field but not the RVF for English speakers and in both visual fields for Hebrew speakers. It is suggested that these differences result from orthographic and morphological differences between the languages: Reading Hebrew requires attention to be deployed to all the constituents of the stimulus in parallel, whereas reading English allows sequential processing of the letters in both hemispheres. Implications of cross-language studies for models of hemispheric function are discussed.     

Visual evoked response correlates of cerebral specialization after human commissurotomy

Visual evoked responses (VERs) were recorded from commissurotomy patients and normal subjects in order to investigate the electrophysiological correlates of cerebral lateralization and independent hemispheric processing. A verbal task consisted of detecting rhyming words and a spatial task consisted of comparing matching shapes. Stimuli were delivered, by means of a computer graphic display, into the right visual field, the left visual field, or both visual fields. Behavioral results and evoked response waveforms showed lateralization for verbal processing in the left hemisphere for the patient group. Analysis of variance of product moment correlation coefficients between VER waveforms indicated significant differences between unilateral and bilateral stimulation for both patients and normals. Commissurotomy and normal VERs showed similar patterns of correlation when verbal stimuli were delivered to the left hemisphere, but differed during right hemisphere stimulation. The results are discussed in terms of VER correlates of visual-verbal processing.     

Lateralization of repetition effects in event-related potentials to words in left- and right-handed women

In our previous event-related potential (ERP) study [A. Nowicka, I. Szatkowska, Memory-induced modulation of event-related potentials in frontal cortex of human subjects: a divided visual field study, Neurosci. Lett. 359 (2004) 171-174], word repetition effects in right-handed males were observed only in case of the direct stimulation of the left (competent) hemisphere. Since the left-handedness and the factor of female gender may determine the lateralization of verbal functions, the goal of the present study was to test the sensitivity of the ERP repetition effects to the visual field of word presentation in the group of left- and right-handed women. ERPs were recorded from symmetrical sites over the left and right hemisphere. Target words were presented in the left (LVF) or right (RVF) visual hemifield. Subjects' were instructed to recognize the target word on a response card. A substantial portion of words was repeated twice. Words elicited different ERPs at frontal sites when presented the first and second times: ERPs were more positive to the repeated stimulus than to its first occurrence. However, in right-handed participants, repetition effects were only observed when target words were presented to the RVF whereas in left-handed participants, repetition effects were only observed when target words were presented to the LVF. These findings are indicative of the asymmetrical involvement of the two hemispheres in memory-induced modulation of brain activity related to verbal processing in left- and right-handed women.     

Morphological structure and hemispheric functioning: the contribution of the right hemisphere to reading in different languages

This study examined the relationship between morphological structure of languages and performance asymmetries of native speakers in lateralized tasks. In 2 experiments, native speakers of English (concatenative morphology stem plus affix) and of Hebrew and Arabic (nonconcatenative root plus word-form morphology) were presented with lateralized lexical decision tasks, in which the morphological structure of both words and nonwords was manipulated. In the 1st study, stimuli were presented unilaterally. In the 2nd study, 2 stimuli were presented bilaterally, and participants were cued to respond to 1 of them. Three different indexes of hemispheric integration were tested: processing dissociation, effects of distractor status, and the bilateral effect. Lateralization patterns in the 3 languages revealed both common and language-specific patterns. For English speakers, only the left hemisphere (LH) was sensitive to morphological structure, consistent with the hypothesis that the LH processes right visual field stimuli independently but that the right hemisphere uses LH abilities to process words in the left visual field. In Hebrew and Arabic, both hemispheres are sensitive to morphological structure, and interhemispheric transfer of information may be more symmetrical than in English. The relationship between universal and experience-specific effects on brain organization is discussed.     

Semantic priming within and between the visual fields: an event-related brain potential study

To investigate interhemispheric transfer of language information, event-related brain potentials (ERPs) were recorded during performance of a semantic matching task in which prime and target word pairs were sequentially presented to either the left (LVF) or the right (RVF) visual field. A posterior slow negative shift prior to the target presentation developed over both hemispheres for LVF primes, whereas for RVF primes the negative shift declined over the right hemisphere. An N400 attenuation was observed for targets semantically related to the primes and was predominant over the left parietal site. The N400 priming effect was significantly reduced for LVF targets preceded by RVF primes, compared to other patterns of presentation. These findings indicate that semantic priming is attenuated in the right hemisphere when information is transferred from the left hemisphere.     

Selectivity of N170 for visual words in the right hemisphere: Evidence from single‐trial analysis

Neuroimaging and neuropsychological studies have identified the involvement of the right posterior region in the processing of visual words. Interestingly, in contrast, ERP studies of the N170 typically demonstrate selectivity for words more strikingly over the left hemisphere. Why is right hemisphere selectivity for words during the N170 epoch typically not observed, despite the clear involvement of this region in word processing? One possibility is that amplitude differences measured on averaged ERPs in previous studies may have been obscured by variation in peak latency across trials. This study examined this possibility by using single‐trial analysis. Results show that words evoked greater single‐trial N170s than control stimuli in the right hemisphere. Additionally, we observed larger trial‐to‐trial variability on N170 peak latency for words as compared to control stimuli over the right hemisphere. Results demonstrate that, in contrast to much of the prior literature, the N170 can be selective to words over the right hemisphere. This discrepancy is explained in terms of variability in trial‐to‐trial peak latency for responses to words over the right hemisphere.     

Exploring cerebral asymmetries for the verb generation task

This study investigated potential right hemisphere involvement in the verb generation task. Six divided visual field experiments explored cerebral asymmetries for word retrieval in the verb generation task as well as in rhyme generation and immediate and delayed word pronunciation. The typical right visual field/left hemisphere (RVF/LH) advantage was observed for pronunciation and rhyme generation. For verb generation, the RVF/LH advantage was obtained only when stimulus items had a single prepotent response and not when there were multiple response alternatives. A semantic priming experiment suggested that activation for less common, related verbs was maintained for a longer time course within the right than within the left hemisphere. The authors suggest that the right hemisphere may play a role in continued activation of semantically related response alternatives in word generation and discuss methodological implications of their findings.     

Early and late VEPs for reading stimuli are altered by common binocular misalignments

Effects of mild binocular anomalies on early and late evoked response complexes to word stimuli were examined in visual evoked potentials (VEPs) recorded over occipital cortex, Wernicke's area, and its right side homolog in university students with (fixation disparity group) and without (normal group) fixation disparity. Stimuli were monocularly or binocularly viewed words of a paragraph presented individually for 100 ms, one per second, and a binocular control condition without linguistic content. An early complex (P125-N170) recorded at Oz and a late complex (N170-late P) recorded from the t4emporoparietal placements were analyzed. The early complex as not influenced by linguistic content. Binocular stimuli generally resulted in larger early and late VEP amplitudes than did monocular stimuli except that the fixation disparity group had no binocular amplitude enhancement at Oz. Left hemisphere amplitude was greater than right for all conditions, more so for words than for nonlinguistic stimuli. For words, the left hemisphere advantage was significant only for the normal group. Relationships between basic visual processing and language processing in relation to early and late complexes are considered from several perspectives.