Right hemisphere reading mechanisms in a global alexic patient

We investigated the implicit, or covert, reading ability of a global alexic patient (EA) to help determine the contribution of the right hemisphere to reading. Previous studies of alexic patients with left hemisphere damage have suggested that the ability to derive meaning from printed words that cannot be read out loud may reflect right hemisphere reading mechanisms. Other investigators have argued that residual left hemisphere abilities are sufficient to account for implicit reading and moreover do not require the postulation of a right hemisphere system that has no role in normal reading processes. However, few studies have assessed covert reading in patients with lesions as extensive as the one in EA, which affected left medial, inferior temporal-occipital cortex, hippocampus, splenium, and dorsal white matter. EA was presented with lexical decision, semantic categorization, phonemic categorization, and letter matching tasks. Although EA was unable to access phonology and could not overtly name words or letters, she was nevertheless capable of making lexical and semantic decisions at above chance levels, with an advantage for concrete versus abstract words. Her oral and written spelling were relatively intact, suggesting that orthographic knowledge is retained, although inaccessible through the visual modality. Based on her ability to access lexical and semantic information without contacting phonological representations, we propose that EA's implicit reading emerges from, and is supported, by the right hemisphere. Finally, we conclude that her spelling and writing abilities are supported by left hemisphere mechanisms.     

Orthographic and phonological priming in the two cerebral hemispheres

The patterns of activation invoked in the two cerebral hemispheres by written words may be different. Two lexical decision experiments investigated several aspects of such activation patterns. Experiment 1 tested phonological and orthographic priming in the hemispheres, manipulating two levels of phonological and two levels of orthographic similarity. Orthographic priming in the left visual field (LVF) was significantly larger than in the right visual field (RVF). In Experiment 2, primes were phonologically identical to the targets (homophones) but differed in their orthographic similarity. For LVF targets, only orthographic priming was significant, whereas for RVF targets, the phonological primes were effective regardless of their orthographic similarity. The results imply that orthographic activation, though maintained by both hemispheres, is more characteristic of right hemisphere word recognition processes, whereas phonological priming is more characteristic of left hemisphere processes.     

Orthographic and phonological priming in the two cerebral hemispheres

The patterns of activation invoked in the two cerebral hemispheres by written words may be different. Two lexical decision experiments investigated several aspects of such activation patterns. Experiment 1 tested phonological and orthographic priming in the hemispheres, manipulating two levels of phonological and two levels of orthographic similarity. Orthographic priming in the left visual field (LVF) was significantly larger than in the right visual field (RVF). In Experiment 2, primes were phonologically identical to the targets (homophones) but differed in their orthographic similarity. For LVF targets, only orthographic priming was significant, whereas for RVF targets, the phonological primes were effective regardless of their orthographic similarity. The results imply that orthographic activation, though maintained by both hemispheres, is more characteristic of right hemisphere word recognition processes, whereas phonological priming is more characteristic of left hemisphere processes.     

Semantic category and rhyming processing in the left and right cerebral hemisphere

In this study, the question of hemispheric abilities in processing explicit semantic and phonological information was addressed by measuring response latencies and performances in two judgement tasks. In the semantic task, word pairs were sequentially presented to the left or right visual field and subjects were asked to judge whether these words were categorically related or not. In the phonological task, the same subjects were asked to decide whether pairs of orthographically dissimilar words rhymed or not. Statistical analysis showed that reaction times (RT) were significantly shorter in both tasks when words were presented to the right visual field. Furthermore, in the semantic task, faster responses were observed in both visual fields when the words were related than when they were unrelated. This response facilitation tended to be stronger when words were presented to the left visual field. This result is in accordance with other results obtained by lexical decision studies showing that priming in the right hemisphere is due to controlled processing. By contrast, RT differences between rhyming and non-rhyming word pairs in the phonological task did not reach significance. This supports previous findings showing that phonological facilitation cannot occur when orthographic and phonological indices are in conflict.     

The role of sound and spelling in auditory word recognition: Further evidence from brain-damaged patients

This follow - up investigation explored the effects of phonological and orthographic relatedness on auditory lexical access in left - and right - hemisphere - damaged individuals. Participants listened to prime–target pairs that shared word - initial phonology (e.g., definite–deaf), initial orthography (e.g., logic–log), both initial phonology and orthography (e.g., message–mess), or were unrelated (e.g., castle– green), presented at two different inter - stimulus intervals. All groups of subjects demonstrated facilitation of lexical decision latencies due to the combined influence of both orthography and phonology, confirming earlier findings concerning rime relations. The findings are briefly discussed in relation to the neural representation of formal lexical codes.     

Semantic and phonemic priming in the cerebral hemispheres

Representation of semantic and phonemic codes in the cerebral hemispheres was investigated in two priming experiments where prime and target words were independently projected to the left or right visual fields. The first experiment, using phonemic primes, confirmed the view that phonological information is not accessible to the right hemisphere. Priming effects were obtained only when the prime and target were both projected to the right visual field. The second experiment, employing category exemplars as primes, again found the left hemisphere to be the principal locus of the priming effects. The right hemisphere was unable, by itself, to activate words related to the exemplar prime. However, projection of the prime to the right visual field significantly facilitated responses to left visual field targets. The present findings support the view advanced by Drews (Neuropsychologia 25, 419-427, 1987) and Levy and Trevarthen (J. exp. Psychol., Hum. Percep. Perform. 2,299-312, 1976) that the left lexicon is structured in accordance with an hierarchy of logical semantic relationships, while the right lexicon is organized on the basis of simple associations between concepts. It is suggested, furthermore, that the patterns of semantic, but not phonological, activation invoked by a prime may be relayed between the two lexicons.     

Orthography influences the perception of speech in alexic patients

Interactive models of reading propose that phonological representations directly activate and/or constrain orthographic representations through feedback. These models also predict that spoken words should activate their orthographic forms. The effect of word orthography on auditory lexical access was investigated in two patients with alexia without agraphia. Several theories of alexia suggest that letter-by-letter reading results from impaired access to orthographic representations. Although alexics can often correctly identify orally spelled words and spell to dictation, it is unknown whether they can access the whole orthographic "word-form" as a unit via auditory presentation. The nonobligatory activation of orthography was examined in an auditory lexical decision task, in which the orthographic and phonological similarity between prime and target was manipulated. In controls, the combined effect of phonological and orthographic relatedness (OP) produced greater facilitation than phonological relatedness alone, indicating that orthography can influence auditory lexical decisions. The alexics displayed patterns of facilitation comparable to controls, suggesting they can quickly access whole-word orthographic information via the auditory modality. An alternate account posits that the OP advantage does not require on-line access of orthography, but instead is a developmental by-product of learning to read an orthographically inconsistent language. The results have implications for cognitive theories of alexia and provide support for interactive models of word recognition.     

Developmental differences of neurocognitive networks for phonological and semantic processing in Chinese word reading

Developmental differences in the neurocognitive networks for phonological and semantic processing in Chinese word reading were examined in 13 adults and 13 children using functional magnetic resonance imaging (fMRI). Rhyming and semantic association judgments were made to two-character words that were presented sequentially in the visual modality. These lexical tasks were compared with a nonlinguistic control task involving judgment of line patterns. The first main finding was that adults showed greater activation than children in right middle occipital gyrus on both the meaning and rhyming task, suggesting adults more effectively engage right hemisphere brain regions involved in the visual-spatial analysis of Chinese characters. The second main finding was that adults showed greater activation than children in left inferior parietal lobule for the rhyming as compared with the meaning task, suggesting greater specialization of phonological processing in adults. The third main finding was that children who had better performance in the rhyming task on characters with conflicting orthographic and phonological information relative to characters with nonconflicting information showed greater activation in left middle frontal gyrus, suggesting greater engagement of brain regions involved in the integration of orthography and phonology.     

The effect of phonological repetition on cortical magnetic responses evoked by visually presented words

Neuroimaging studies have reported that the left superior temporal cortical area is activated by visually presented words. In the present study, we recorded cortical magnetic responses evoked by visual words and examined the effect of phonological repetition (e.g., hair-hare) on left superior temporal cortical activity, using pairs of homophonic Japanese words as stimuli. Unlike English, Japanese has a large number of homophone pairs with a totally different orthography. By taking advantage of this feature of the Japanese writing system, the effect of phonological repetition can be solely examined without being confounded by the effect of orthographic similarity. Magnetic responses were recorded over the bilateral temporal sites of the brain while subjects silently read words. The words were presented one by one; a quarter of them was immediately followed by a homophonic word. Clear magnetic responses in the latency range of 300-600 msec were observed in the left hemisphere, and the responses to the homophones were smaller than those to the first presented words. In the right hemisphere, clear responses were not consistently recorded in the same latency range, and no effect of phonological repetition was observed. The sources of the responses recorded over the left hemisphere were estimated to be in the left superior temporal cortical area adjacent to the auditory cortex and the source strength as well as the magnetic responses showed a reduction by phonological repetition. This result suggests that the activity in the left superior temporal cortical area is associated with access to the phonological representation of words.     

Hemispheric asymmetry in lexical access and phonological encoding

In a lexical decision task, a right visual field (RVF) advantage was found for all types of stimuli, irrespective of right- or left-handed responding, and nonwords displayed a clear effect of phonological encoding. It is argued that the left hemisphere is specialised for lexical access and is responsible for phonological encoding of visually-presented words.     

Effects of spelling-to-sound regularity on word identification following brief presentation in the right or left visual field

In this experiment subjects were briefly presented with a vertically written word to either the left or right visual field. On half the critical trials the target word was irregular in its spelling-to-sound correspondence (e.g. PINT) and in the other half it was regular (e.g. LIME). With right visual field presentation substantially more regular than irregular words were reported correctly but with left visual field presentation no such effect was found. The data also showed the presence of far more "regularization" errors following right visual field presentation. The data are seen as consistent with the view that the right hemisphere is not capable of generating phonology.     

Interhemispheric cooperation and activation in integration of verbal information

Subjects had to judge the semantic relation between two tachistoscopically presented German adjectives. In Exp. I, the two words were projected simultaneously, either both to the left visual field (i.e. to the right hemisphere, LVF-RH), or both to the right visual field (i.e. to the left hemisphere, RVF-LH), or one word to each visual field (i.e. to both hemispheres, EVF-BH). The results yielded a significantly higher performance in the condition EVF-BH than in the two unilateral conditions, without a significant difference between the latter two conditions. In Exp. II, the two words were shown successively. With bilateral presentation, a distinction was made as to whether the left visual field (EVF-BH-L) or the right visual field (EVF-BH-R) was stimulated first. The significantly highest performance was shown in conditions RVF-LH and EVF-BH-R compared with the condition EVF-BH-L, and the significantly poorest performance in the condition LVF-RH compared with the condition EVF-BH-L. The results provide evidence for (1) an intrahemispheric interference with simultaneous unilateral presentation, (2) a cooperative interhemispheric interaction with simultaneous bilateral presentation, and (3) an interhemispheric priming effect with successive bilateral presentation when the left hemisphere was stimulated first.     

Superior Written Over Spoken Picture Naming in a Case of Frontotemporal Dementia

Two main hypotheses have been proposed regarding the role of phonology in written word production. According to the phonological mediation hypothesis, the retrieval of the lexical phonological representation of a word is an obligatory prerequisite to the retrieval of its spelling. Therefore, deficits to the phonological lexicon should affect both spoken and written picture naming. In contrast, the orthographic autonomy hypothesis posits that the lexical orthographic representations of words can be accessed without any necessary phonological mediation. In support of this view, cases of preserved written naming despite impaired lexical phonology have been reported following brain damage. In this report, we replicate this basic pattern of performance in case YP, a 60-year-old woman with a pattern of frontotemporal dementia. As her disease progressed, YP’s ability to write down the names of pictures remained very good despite a severe decline in oral naming. Further testing indicated that this deficit was not primarily due to an articulatory or post-lexical phonological deficit. YP’s case provides strong additional support for the orthographic autonomy hypothesis. The significance of this case with respect to the characterization of dementia syndromes is discussed.     

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.     

From orthography to phonetics: ERP measures of grapheme-to-phoneme conversion mechanisms in reading

Neuroimaging has provided evidence that the first stages of visual word recognition activate a visual word-form center localized in the left extrastriate cortex (fusiform gyrus). Accordingly, neurological cases of patients suffering from pure alexia reported the left posterior occipital lobe as the possible locus of orthographic analysis. There is less agreement in the literature about which brain structures are involved in the subsequent stages of word processing and, in particular, their time course of activation. Functional magnetic resonance imaging and magnetic source imaging studies recently reported data that could indicate a dual route model of reading. These findings are particularly relevant to studies on the functional deficits associated with phonological and surface dyslexia. There is evidence for the existence of two different brain mechanisms supporting phonological processing in visual word recognition: one mechanism subserving "assembled phonology" for reading letter strings and another one subserving "addressed phonology" for reading meaningful words. However, available knowledge on the time course and neural locus of grapheme-to-phoneme conversion mechanisms in reading is still inadequate. In this study, we compared processing of meaningful and meaningless Italian words in a task requiring a phonemic/phonetic decision task. Stimuli were 1152 different orthographic stimuli presented in the central visual field. Half the stimuli were Italian words (with a high or low frequency of occurrence), the other half were meaningless strings of letters (legal pseudowords and letter strings). Event-related potentials were recorded from 28 scalp sites in 10 Italian university students. The task consisted of deciding about the presence/absence of a given "phone" in the hypothetical enunciation of word read: for example, "Is there a /k/ in cheese?". Results showed that lexical frequency and orthographical regularity affected linguistic processing within 150 msec poststimulus. Indeed, the amplitude of a centroparietal P150 varied as a function of stimulus type, being larger in response to high-frequency words than to low-frequency ones and to words and pseudowords than to letter strings. This component might index visual categorization processes and recognition of familiar objects, being highly sensitive to orthographic regularity and "ill-formedness" of words. The amplitude of the P150 was the same in response to well-formed meaningless and to meaningful words, when these latter had a low lexical frequency. This might indicate that highly familiar words are recognized as meaningful unitary visual objects at very early stages of processing, through a visual route to an orthographic input lexicon. Moreover, the amplitude of the negativity recorded between 250 and 350 msec showed an anteroposterior topographic dissociation for access to the phonemic representation of well- or ill-formed strings of characters. Brain responses were larger over the left occipito-temporal regions during reading of words and pseudowords and over the left frontal regions during reading of letter strings.     

Superior written over spoken picture naming in a case of frontotemporal dementia

Two main hypotheses have been proposed regarding the role of phonology in written word production. According to the phonological mediation hypothesis, the retrieval of the lexical phonological representation of a word is an obligatory prerequisite to the retrieval of its spelling. Therefore, deficits to the phonological lexicon should affect both spoken and written picture naming. In contrast, the orthographic autonomy hypothesis posits that the lexical orthographic representations of words can be accessed without any necessary phonological mediation. In support of this view, cases of preserved written naming despite impaired lexical phonology have been reported following brain damage. In this report, we replicate this basic pattern of performance in case YP, a 60-year-old woman with a pattern of frontotemporal dementia. As her disease progressed, YP's ability to write down the names of pictures remained very good despite a severe decline in oral naming. Further testing indicated that this deficit was not primarily due to an articulatory or post-lexical phonological deficit. YP's case provides strong additional support for the orthographic autonomy hypothesis. The significance of this case with respect to the characterization of dementia syndromes is discussed.     

Differences in semantic category priming in the left and right cerebral hemispheres under automatic and controlled processing conditions.

The contribution of each cerebral hemisphere to the generation of semantic category meanings at automatic and strategic levels of processing was investigated in a priming experiment where prime and target words were independently projected to the left or right visual fields (LVF or RVF). Non-associated category exemplars were employed as related pairs in a lexical decision task and presented in two experimental conditions. The first condition was designed to elicit automatic processing, so related pairs comprised 20% of the positive set, stimulus pairs were temporally separated by a stimulus onset asynchrony (SOA) of 250 ms, and there was no allusion to the presence of related pairs in the instructions to subjects. The second condition, designed to invoke controlled processing, incorporated a relatedness proportion of 50%, stimulus pairs separated by an SOA of 750 ms, and instructions which informed subjects of the presence and use of category exemplar pairs in the stimulus set. In the first condition, a prime directed to either visual field facilitated responses to categorically related targets subsequently projected to the RVF, while in the second condition a prime directed to either visual field facilitated responses to related targets projected to the LVF. The facilitation effects obtained in both conditions appeared to reflect automatic processes, while strategic processes were invoked in the left, but not the right hemisphere in the second condition. The results suggest that both hemispheres have automatic access to semantic category meanings, although the timecourse of activation of semantic category meanings is slower in the right hemisphere than in the left.     

Associative priming in the hemispheres as a function of SOA

The representation of associative codes in the cerebral hemisphere was investigated in two priming experiments where associated prime and target words were independently projected to the left or right visual fields. The first experiment, using a stimulus onset asynchrony of 250 msec, found priming in all visual field conditions, except that in which both prime and target were projected to the right hemisphere. A second experiment was conducted to determine whether this absence of right hemisphere priming was due to an inadequate interval of time between prime and target. In this experiment, a stimulus onset asynchrony of 450 msec revealed significant priming in all visual field conditions. The results suggest that lexical representations are activated more slowly in the right hemisphere than in the left.     

Dissociated roles of the middle frontal gyri in the processing of Chinese characters

The present study examined a hypothesis that the right middle frontal gyrus participates in processing orthography of Chinese characters, while the left middle frontal gyrus mediates access to phonology and semantics. Brain activation during three character tasks, which required processing orthography, phonology, or semantics of Chinese characters, respectively, was measured by functional magnetic resonance imaging. Comparable neural activity in the right middle frontal gyrus was observed in all three character tasks that always demand orthographical processing. In contrast, the left middle frontal gyrus showed greater activation in the phonological and semantic tasks than in the orthographic task. These results suggest that the right and left middle frontal gyrus have dissociable functions in achieving Chinese character recognition.     

ERP manifestations of processing printed words at different psycholinguistic levels: time course and scalp distribution.

The aim of the present study was to examine the time course and scalp distribution of electrophysiological manifestations of the visual word recognition mechanism. Event-related potentials (ERPs) elicited by visually presented lists of words were recorded while subjects were involved in a series of oddball tasks. The distinction between the designated target and nontarget stimuli was manipulated to induce a different level of processing in each session (visual, phonological/phonetic, phonological/lexical, and semantic). The ERPs of main interest in this study were those elicited by nontarget stimuli. In the visual task the targets were twice as big as the nontargets. Words, pseudowords, strings of consonants, strings of alphanumeric symbols, and strings of forms elicited a sharp negative peak at 170 msec (N170); their distribution was limited to the occipito-temporal sites. For the left hemisphere electrode sites, the N170 was larger for orthographic than for nonorthographic stimuli and vice versa for the right hemisphere. The ERPs elicited by all orthographic stimuli formed a clearly distinct cluster that was different from the ERPs elicited by nonorthographic stimuli. In the phonological/phonetic decision task the targets were words and pseudowords rhyming with the French word vitrail, whereas the nontargets were words, pseudowords, and strings of consonants that did not rhyme with vitrail. The most conspicuous potential was a negative peak at 320 msec, which was similarly elicited by pronounceable stimuli but not by nonpronounceable stimuli. The N320 was bilaterally distributed over the middle temporal lobe and was significantly larger over the left than over the right hemisphere. In the phonological/lexical processing task we compared the ERPs elicited by strings of consonants (among which words were selected), pseudowords (among which words were selected), and by words (among which pseudowords were selected). The most conspicuous potential in these tasks was a negative potential peaking at 350 msec (N350) elicited by phonologically legal but not by phonologically illegal stimuli. The distribution of the N350 was similar to that of the N320, but it was broader and including temporo-parietal areas that were not activated in the "rhyme" task. Finally, in the semantic task the targets were abstract words, and the nontargets were concrete words, pseudowords, and strings of consonants. The negative potential in this task peaked at 450 msec. Unlike the lexical decision, the negative peak in this task significantly distinguished not only between phonologically legal and illegal words but also between meaningful (words) and meaningless (pseudowords) phonologically legal structures. The distribution of the N450 included the areas activated in the lexical decision task but also areas in the fronto-central regions. The present data corroborated the functional neuroanatomy of word recognition systems suggested by other neuroimaging methods and described their timecourse, supporting a cascade-type process that involves different but interconnected neural modules, each responsible for a different level of processing word-related information.