Task and Stimulus Demands Influence Letter-case-Specific Priming in the Right Cerebral Hemisphere

A greater tendency to complete single-completion word stems (e.g. “BEY”) to form previously read whole words (e.g. “BEYOND”) was found when test stems were presented in the same letter case as their previously encoded words, compared with the different letter case, but only when stems were presented directly to the right hemisphere (i.e. in the left visual field) and not when they were presented directly to the left hemisphere (i.e. in the right visual field). This finding with single-completion stems was robust (i.e. observed for both lowercase and uppercase stems) when the initial encoding task was perceptually demanding, but it was test-case dependent (i.e. observed for uppercase but not lowercase stems) when the initial encoding task was not perceptually demanding. Results and theory help to explain why letter-case-specific priming in right-hemisphere test presentations is typically test-case dependent when priming is measured using perceptual identification at test, but is consistently robust when priming is measured using multiple-completion word stems (e.g.“BEA”) at test. Demands from both the stimuli and tasks affect the relative contributions of abstract and specific subsystems to the processing of visual forms.     

The neural substrates of visual implicit memory: do the two hemispheres play different roles?

Identification of visually presented words is facilitated by implicit memory, or visual priming, for past visual experiences with those words. There is disagreement over the neuro-anatomical substrates of this form of implicit memory. Several studies have suggested that this form of priming relies on a visual word-form system localized in the right occipital lobe, whereas other studies have indicated that both hemispheres are equally involved. The discrepancies may be related to the types of priming tasks that have been used because the former studies have relied primarily on word-stem completion tasks and the latter on tasks like word-fragment completion. The present experiments compared word-fragment and word-stem measurements of visual implicit memory in patients with right occipital lobe lesions and patients with complete callosotomies. The patients showed normal visual implicit memory on fragment completion tests, but essentially no visual priming on standard stem completion tests. However, when we used a set of word stems that had only one correct solution for each test item, as was true of the items in the fragment completion tests, the patients showed normal priming effects. The results indicate that visual implicit memory for words is not solely dependent upon the right hemisphere, rather it reflects changes in processing efficiency in bilateral visual regions involved in the initial processing of the items. However, under conditions of high lexical competition (i.e., multiple completion word stems), the lexical processes, which are dominant in the left hemisphere, overshadow the visual priming supported by the left hemisphere.     

Presentation of words to separate hemispheres prevents interword illusory conjunctions

We tested the hypothesis that division of inputs between the hemispheres could prevent interword letter migrations in the form of illusory conjunctions. The task was to decide whether a centrally-presented consonant-vowel-consonant (CVC) target word matched one of four CVC words presented to a single hemisphere or divided between the hemispheres in a subsequent test display. During half of the target-absent trials, known as conjunction trials, letters from two separate words (e.g., "tag" and "cop") in the test display could be mistaken for a target word (e.g., "top"). For the other half of the target-absent trails, the test display did not match any target consonants (Experiment 1, N = 16) or it matched one target consonant (Experiment 2, N = 29), the latter constituting true "feature" trials. Bi- as compared to unihemispheric presentation significantly reduced the number of conjunction, but not feature, errors. Illusory conjunctions did not occur when the words were presented to separate hemispheres.     

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

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

Interference and priming within and across visual fields in a lexical decision task

A lower-case target word or non-word was laterally presented by itself or simultaneously with an upper-case distractor word in the same or in the opposite visual field. The distractor words were either semantically associated or unrelated to the target. Subjects performed a lexical decision task on the target. Two kinds of selection within and across visual fields were investigated. Selection of an external target relevant for the response (filtering) and selection of an internal target by automatic priming. No lateral asymmetry was found for automatic priming. Filtering was equally efficient within hemispheres, but strongly asymmetrical across hemispheres. Unrelated distractor words presented to the right hemisphere interfered less with target processing in the left hemisphere than vice versa. Thus, all distractor words received equivalent semantic processing; those presented to the right hemisphere, however, were least costly to exclude from task-relevant processing in the left hemisphere.     

Hemispheric asymmetries in memory processes as measured in a false recognition paradigm

Although memory differs in important ways between the left and right cerebral hemispheres, the nature of these differences remains controversial. We examined this issue in two experiments using a false memory paradigm that allowed novel tests of two theories that have not been assessed in a common paradigm previously. Lists of semantically related words (e.g., bed, rest, wake…), all highly associated to one “critical” word (e.g., sleep), were presented auditorily during a study phase. Memory for both the related words and the critical words was measured in a subsequent old/new recognition test using divided-visual- field word presentations. The most important results were that the ability to correctly reject previously unpresented words was greater when test items were presented to the right visual field/left hemisphere (RVF/LH) than to the left visual field/right hemisphere (LVF/RH) and that participants were more confident in correctly rejecting unpresented words when test items were presented to the RVF/LH than to the LVF/RH. Results were in line with the theory that associative activation of semantic information is restricted in the left hemisphere but diffuse in the right; however, these results contrasted with the theory that memory traces are interpretive in the left hemisphere but veridical in the right. A potential resolution to the seemingly contradictory theories of asymmetries in memory processing is briefly discussed.     

Multiple levels of stimulus representation in visual working memory

Object recognition presumably involves activation of multiple levels of representation. Here we use the encoding-related lateralization (ERL) method [Gratton, G. The contralateral organization of visual memory: A theoretical concept and a research tool. Psychophysiology, 35, 638-647, 1998] to describe the sequential activation of several of these levels. The ERL uses divided-field encoding to generate contralaterally biased representations in the brain. The presence and nature of these representations can be demonstrated by examining the event-related potentials (ERPs) elicited by centrally presented test probes for lateralized activity corresponding to the encoding side. We recorded ERPs during a memory-search task. Memory sets were composed of two or four uppercase letters displayed half to the left and half to the right of fixation. Probe stimuli were composed of one letter presented foveally in either upper- or lowercase. Letter case was manipulated to differentiate the time course of physical and symbolic levels of letter representation. Memory set size was manipulated to examine a relational level of letter representation. We found multiple ERLs in response to the probes: (1) An early (peak=170 msec) case-dependent (but set size independent) ERL, most evident at P7/P8, indexing the availability of a physical level of letter representation; (2) a later (200-400 msec) more diffusedly distributed ERL, independent of both letter case and set size, indexing a symbolic level of letter representation; (3) a long-latency (400-600 msec) ERL occurring at posterior sites, larger for the case match, Set Size 2 condition, indexing competition for neural representation across multiple letters. By assuming that these ERL activities track the progression of letter representation over time, we propose a model of letter processing in the context of visual working memory.     

Analysis of standard and non-standard visual word format in the two hemispheres

A dual-route model for the recognition of written words is hypothesised. The model postulates that the two cerebral hemispheres differ in their sensitivity to the visual format of verbal stimuli. Such stimuli may be 'standard', that is in Standard Visual Format (SVF), or "non-standard", i.e. in non-standard Visual Format (NSVF). In the left hemisphere (LH), SVF words are routed directly to the lexicon, while NSVF words must go through an additional encoding phase prior to lexicon access. In the right hemisphere (RH) all words (SVF and NSVF) must proceed to an additional encoding phase prior to accessing the lexicon. Two lateralized lexical decision experiments are reported. In Section 2, angle of orientation was used to define SVF and NSVF. In Section 3, presentation of verbal stimuli was preceded by a cue indicating whether the coming lexical stimulus was in SVF or NSVF, while SVF was defined as in Section 2. The results showed that the difference in RT, favouring RVF stimulation was found for orientations up to 30 degrees. Greater angles of presentation significantly reduced RVF superiority. The largest differences between responses to stimuli presented to the two visual fields was found when word stimuli were in SVF (up to 30 degrees ) and presented after a priming cue. The results are discussed in terms of the model, but alternative explanations are suggested.     

Evaluating a split fovea model of visual word recognition: effects of case alternation in the two visual fields and in the left and right halves of words presented at the fovea

Two experiments are reported exploring the effect of cAsE aLtErNaTiOn on lexical decisions to words and nonwords presented laterally or centrally. In line with previous research, Experiment 1 found that case alternation slowed lexical decision responses to words more in the right visual field (RVF) than in the left visual field (LVF). In Experiment 2, the words and nonwords were all presented centrally. There were three conditions, a condition in which the word and nonwords were presented in lower case letters, a condition in which the letters to the left of the central fixation were case alternated (e.g., aMbItion, mOdLants) and a condition in which the letters to the right of fixation were case alternated (e.g., collApSe, pireNtOl). Alternating the case of letters to the right of fixation slowed lexical decision responses more than alternating letter case to the left of fixation. The results provide further support for a split fovea account of visual word recognition according to which those letters of a centrally-fixated word that fall to the left of fixation are processed initially by the right cerebral hemisphere while those letters that fall to the right of fixation are processed initially by the left cerebral hemisphere, with the characteristics of the left and right hemispheres being revealed in the processing of initial and final letters in centrally presented words.     

Intact implicit memory in patients with frontal lobe lesions.

Patients with frontal lobe lesions and control subjects were administered tests of word-stem completion priming. In this implicit memory test, subjects are first presented words (e.g. MOTEL, PARADE) in an incidental learning paradigm. Following word presentation, subjects are shown word stems (e.g. MOT, PAR) and asked to produce the first word that comes to mind. Patients with frontal lobe lesions exhibited normal levels of word-stem completion. These findings indicate that implicit memory can operate normally despite damage to the prefrontal cortex. The present results substantiate previous neuropsychological and positron emission tomography findings which indicate that word priming depends critically on posterior cortical areas.     

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.     

Levels-of-processing effect on word recognition in schizophrenia.

BACKGROUND: Individuals with schizophrenia have difficulty organizing words semantically to facilitate encoding. This is commonly attributed to organizational rather than semantic processing limitations. By requiring participants to classify and encode words on either a shallow (e.g., uppercase/lowercase) or deep level (e.g., concrete/abstract), the levels-of-processing paradigm eliminates the need to generate organizational strategies. METHODS: This paradigm was administered to 30 patients with schizophrenia and 30 healthy comparison subjects to test whether providing a strategy would improve patient performance. RESULTS: Word classification during shallow and deep encoding was slower and less accurate in patients. Patients also responded slowly during recognition testing and maintained a more conservative response bias following deep encoding; however, both groups showed a robust levels-of-processing effect on recognition accuracy, with unimpaired patient performance following both shallow and deep encoding. CONCLUSIONS: This normal levels-of-processing effect in the patient sample suggests that semantic processing is sufficiently intact for patients to benefit from organizational cues. Memory remediation efforts may therefore be most successful if they focus on teaching patients to form organizational strategies during initial encoding.     

Rightward bisection errors for letter lines: the role of semantic information

When bisecting words in their middle, people reveal leftward bisection errors. This tendency might emerge from an attentional bias towards the beginning of the word. However, when longer meaningless letter strings are presented, people reveal a rightward bisection bias. To test the role of semantic information on leftward or rightward bisection biases, we tested letter line bisection performance in healthy right-handed students in four independent experiments. A third of the letter lines contained an embedded four-letter word to the left of true centre, another third contained an embedded four-letter word to the right of true centre, while the remaining lines contained no words. Half of these words were emotional words, the other half were neutral words. Results across experiments revealed a stronger rightward bisection bias: (i) for letter lines containing emotional as compared to neutral words, (ii) for letter lines containing words in the left as compared to right half of the lines, and (iii) for those experiments in which the spatial position of letter lines remained within a narrow body-centred space. Findings from this study suggest that letter line bisection performance might be only minimally determined by visuo-spatial attention. Rather, letter line perception might activate the left hemisphere more than the right hemisphere, shifting the subjective midpoint to the right of true centre. Leftward bisection biases for words only, as had been described in the literature, may thus have resulted from automated reading strategies rather than from attentional biases towards the left hemispace.     

Unconscious relational inference recruits the hippocampus

Relational inference denotes the capacity to encode, flexibly retrieve, and integrate multiple memories to combine past experiences to update knowledge and improve decision-making in new situations. Although relational inference is thought to depend on the hippocampus and consciousness, we now show in young, healthy men that it may occur outside consciousness but still recruits the hippocampus. In temporally distinct and unique subliminal episodes, we presented word pairs that either overlapped ("winter-red", "red-computer") or not. Effects of unconscious relational inference emerged in reaction times recorded during unconscious encoding and in the outcome of decisions made 1 min later at test, when participants judged the semantic relatedness of two supraliminal words. These words were either episodically related through a common word ("winter-computer" related through "red") or unrelated. Hippocampal activity increased during the unconscious encoding of overlapping versus nonoverlapping word pairs and during the unconscious retrieval of episodically related versus unrelated words. Furthermore, hippocampal activity during unconscious encoding predicted the outcome of decisions made at test. Hence, unconscious inference may influence decision-making in new situations.     

Lateralised covert attention in word identification

The right visual field superiority in word recognition has been attributed to an attentional advantage by the left brain hemisphere. We investigated whether such advantage involves lateralised covert attention, in the absence of overt fixations on prime words. In a lexical decision task target words were preceded by an identical or an unrelated prime word. Eye movements were monitored. In Experiment 1 lateralised (to the left or right of fixation) prime words were parafoveally visible but foveally masked, thus allowing for covert attention but preventing overt attention. In Experiment 2 prime words were presented at fixation, thus allowing for both overt and covert attention. Results revealed positive priming in the absence of fixations on the primes when these were presented in the right visual field. The effects of covertly attended primes were nevertheless significantly reduced in comparison with those of overtly attended primes. It is concluded that word identification can be accomplished to a significant extent by lateralised covert attention alone, with right visual field advantage.     

Lateralised covert attention in word identification

The right visual field superiority in word recognition has been attributed to an attentional advantage by the left brain hemisphere. We investigated whether such advantage involves lateralised covert attention, in the absence of overt fixations on prime words. In a lexical decision task target words were preceded by an identical or an unrelated prime word. Eye movements were monitored. In Experiment 1 lateralised (to the left or right of fixation) prime words were parafoveally visible but foveally masked, thus allowing for covert attention but preventing overt attention. In Experiment 2 prime words were presented at fixation, thus allowing for both overt and covert attention. Results revealed positive priming in the absence of fixations on the primes when these were presented in the right visual field. The effects of covertly attended primes were nevertheless significantly reduced in comparison with those of overtly attended primes. It is concluded that word identification can be accomplished to a significant extent by lateralised covert attention alone, with right visual field advantage.     

ERP evidence of morphological analysis from orthography: a masked priming study

There is broad consensus that the visual word recognition system is sensitive to morphological structure (e.g., "hunter" = "hunt" + "er"). Moreover, it has been assumed that the analysis of morphologically complex words (e.g., "hunter") occurs only if the meaning of the complex form can be derived from the meanings of its constituents (e.g., "hunt" and "er"). However, recent behavioral work using masked priming has suggested that morphological analysis can occur at an early, orthographic level, with little influence from semantics. The present investigation examined the neurophysiological correlates of masked priming in conditions of a genuine morphological relationship (e.g., "hunter"-"HUNT"), an apparent morphological relationship ("corner"-"CORN"), and no morphological relationship ("brothel"-"BROTH"). Neural priming was indexed by the reduction of the N400 ERP component associated with targets preceded by related primes, as compared to targets preceded by unrelated primes. The mere appearance of morphological structure ("corner"-"CORN") resulted in robust behavioral and neural priming, whose magnitude was similar to that observed in pairs with genuine morphological relationship and greater than that in the nonmorphological pairs. The results support a purely structural morphemic segmentation procedure operating in the early stages of visual word perception.     

Top‐down and bottom‐up influences on the left ventral occipito‐temporal cortex during visual word recognition: An analysis of effective connectivity

The functional role of the left ventral occipito‐temporal cortex (vOT) in visual word processing has been studied extensively. A prominent observation is higher activation for unfamiliar but pronounceable letter strings compared to regular words in this region. Some functional accounts have interpreted this finding as driven by top‐down influences (e.g., Dehaene and Cohen [ 2011 ]: Trends Cogn Sci 15:254–262; Price and Devlin [ 2011 ]: Trends Cogn Sci 15:246–253), while others have suggested a difference in bottom‐up processing (e.g., Glezer et al. [ 2009 ]: Neuron 62:199–204; Kronbichler et al. [ 2007 ]: J Cogn Neurosci 19:1584–1594). We used dynamic causal modeling for fMRI data to test bottom‐up and top‐down influences on the left vOT during visual processing of regular words and unfamiliar letter strings. Regular words (e.g., taxi) and unfamiliar letter strings of pseudohomophones (e.g., taksi) were presented in the context of a phonological lexical decision task (i.e., “Does the item sound like a word?”). We found no differences in top‐down signaling, but a strong increase in bottom‐up signaling from the occipital cortex to the left vOT for pseudohomophones compared to words. This finding can be linked to functional accounts which assume that the left vOT contains neurons tuned to complex orthographic features such as morphemes or words [e.g., Dehaene and Cohen [ 2011 ]: Trends Cogn Sci 15:254‐262; Kronbichler et al. [ 2007 ]: J Cogn Neurosci 19:1584–1594]: For words, bottom‐up signals converge onto a matching orthographic representation in the left vOT. For pseudohomophones, the propagated signals do not converge, but (partially) activate multiple orthographic word representations, reflected in increased effective connectivity. Hum Brain Mapp 35:1668–1680, 2014. © 2013 Wiley Periodicals, Inc.