The effects of focal and global attentional systems on spatial biases

The background page on which a stimulus is presented can influence the allocation of attention to that stimulus. The purpose of this study was to learn if there are hemispheric asymmetries in how background distraction affects attentional processing. Asymmetries were investigated by having right eye dominant subjects perform line bisections and manipulating the side of background distraction (right versus left), the eye of regard (right versus left), and the type of attention allocated (focal versus global). Overall subjects bisected lines to the left of center (pseudoneglect) and when viewing with the right eye (versus left) deviated more to the left. Subjects had more background distraction when viewing symbol than solid lines. Although overall, bias did not differ with the side of background distraction or the line being on one side or the other, when subjects viewed symbol, but not solid lines, this leftward bias was increased when the line was displaced to the right, thereby increasing the size of the left sided background. These findings suggest that when engaging the left hemisphere by using focused attention and placing the line on the right side, there is more distraction than when the right hemisphere is engaged.     

Neuroendocrine hemisphere asymmetries: Salivary cortisol secretion during lateralized viewing of emotion-related and neutral films

The study set out to examine whether the cerebral hemispheres differ in their ability to regulate cortisol secretion during emotion-related situations. One hundred twenty-three adult subjects were shown either an emotionally aversive or a neutral film in the left or right hemisphere by means of a technique for lateralizing visual input that allows prolonged viewing while permitting free ocular scanning. The film-related changes of cortisol secretion were determined by salivary cortisol radioimmunoassay. Right hemispheric viewing of the emotionally aversive film resulted in a significantly higher increase of cortisol secretion than left hemispheric viewing of the same film. No such differences were observed with respect to the neutral film. Comparing the effects of the two films separately for each hemisphere revealed that only the right hemisphere was able to respond neuroendocrinologically in a different manner to the emotional and the neutral film. Therefore, it is concluded that cortical regulation of cortisol secretion in emotion-related situations is under primary control of the right hemisphere. The potential implications of asymmetric control of cortisol secretion with respect to the pathogenesis of psychosomatic and immunological disorders are discussed.     

Dissociable Neural Subsystems Underlie Abstract and Specific Object Recognition

Participants named objects presented in the left or right visual field during a test phase, after viewing centrally presented same-exemplar objects, different-exemplar objects, and words that name objects during an initial encoding phase. In two experiments, repetition priming was exemplar-abstract yet visual when test objects were presented directly to the left cerebral hemisphere, but exemplar-specific when test objects were presented directly to the right cerebral hemisphere, contrary to predictions from single-system theories of object recognition. In two other experiments, stimulus degradation during encoding and task demands during test modulated these results in predicted ways. The results support the theory that dissociable neural subsystems operate in parallel (not in sequence) to underlie visual object recognition: An abstract-category subsystem operates more effectively than a specific-exemplar subsystem in the left hemisphere, and a specific-exemplar subsystem operates more effectively than an abstract-category subsystem in the right hemisphere.     

The valence-specific laterality effect in free viewing conditions: The influence of sex, handedness, and response bias

The right hemisphere has often been viewed as having a dominant role in the processing of emotional information. Other evidence indicates that both hemispheres process emotional information but their involvement is valence specific, with the right hemisphere dealing with negative emotions and the left hemisphere preferentially processing positive emotions. This has been found under both restricted (Reuter-Lorenz & Davidson, 1981) and free viewing conditions (Jansari, Tranel, & Adophs, 2000). It remains unclear whether the valence-specific laterality effect is also sex specific or is influenced by the handedness of participants. To explore this issue we repeated Jansari et al.'s free-viewing laterality task with 78 participants. We found a valence-specific laterality effect in women but not men, with women discriminating negative emotional expressions more accurately when the face was presented on the left-hand side and discriminating positive emotions more accurately when those faces were presented on the right-hand side. These results indicate that under free viewing conditions women are more lateralised for the processing of facial emotion than are men. Handedness did not affect the lateralised processing of facial emotion. Finally, participants demonstrated a response bias on control trials, where facial emotion did not differ between the faces. Participants selected the left-hand side more frequently when they believed the expression was negative and the right-hand side more frequently when they believed the expression was positive. This response bias can cause a spurious valence-specific laterality effect which might have contributed to the conflicting findings within the literature.     

Lateralisation of emotions: evidence from pupil size measurement

The way our brain processes emotional stimuli has been studied intensively. One of the main issues still under debate is the laterality of valence processing. Herein, we employed the fact that pupil size increases under conditions of higher mental effort and during emotional processing, in order to contrast three proposed hypotheses in the field. We used different manual response mapping for emotional stimuli: Participants responded with their right hand for positive and with their left hand for negative facial expressions, or vice versa. The hands position was either regular (Experiment 1) or crossed (Experiment 2) in order to rule out a “spatial-valence association” alternate explanation. A third experiment was conducted by employing a passive viewing procedure of peripheral emotional stimuli. In the first two experiments, pupil size was larger when participants responded to positive stimuli with their left hand and to negative with their right hand, compared with the opposite mapping. Results of Experiment 3 strengthen the findings of Experiments 1 and 2. These findings provide significant psychophysiological evidence for the valence hypothesis: Processing positive stimuli involves the left hemisphere, while processing negative stimuli involves the right hemisphere. These results are discussed in relation to contemporary theories of emotion processing.     

SEMANTIC NETWORKS IN THE DIVIDED CEREBRAL HEMISPHERES

Abstract— Hemispheric differences in the recognition and manipulation of meaning may be based on distinctions in size, composition, or organization of the right and left semantic networks. The present study describes these features of pictorially based semantic networks in 3 subjects with complete forebrain commissurotomy. Stimuli were presented for prolonged viewing to the left and right visual hemifields. For each trial, the subjects chose from a 20-choice array all pictures that were associated with a target, then indicated the member of each pair of chosen associates that was more closely related to the target. The hemispheres' networks were found to be of similar size and composition, but were organized differently. The right hemisphere more often produced linear rankings of semantic associates to a target than did the left, and rankings by the two hemispheres were not strongly correlated. Hemispheric differences in semantic organization mirror differences in perceptual organization, with the right hemisphere specialized for conventional meaning and the left hemisphere specialized for detecting and processing deviations from standard meaning.     

No strong evidence for lateralisation of word reading and face recognition deficits following posterior brain injury

Face recognition and word reading are thought to be mediated by relatively independent cognitive systems lateralised to the right and left hemispheres, respectively. In this case, we should expect a higher incidence of face recognition problems in patients with right hemisphere injury and a higher incidence of reading problems in patients with left hemisphere injury. We tested this hypothesis in a group of 31 patients with unilateral right or left hemisphere infarcts in the territory of the posterior cerebral arteries. In most domains tested (e.g., visual attention, object recognition, visuo-construction, motion perception), we found that both patient groups performed significantly worse than a matched control group. In particular, we found a significant number of face recognition deficits in patients with left hemisphere injury and a significant number of patients with word reading deficits following right hemisphere injury. This suggests that face recognition and word reading may be mediated by more bilaterally distributed neural systems than is commonly assumed.     

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.     

Bilateral capacity for speech sound processing in auditory comprehension: evidence from Wada procedures

Data from lesion studies suggest that the ability to perceive speech sounds, as measured by auditory comprehension tasks, is supported by temporal lobe systems in both the left and right hemisphere. For example, patients with left temporal lobe damage and auditory comprehension deficits (i.e., Wernicke's aphasics), nonetheless comprehend isolated words better than one would expect if their speech perception system had been largely destroyed (70-80% accuracy). Further, when comprehension fails in such patients their errors are more often semantically-based, than-phonemically based. The question addressed by the present study is whether this ability of the right hemisphere to process speech sounds is a result of plastic reorganization following chronic left hemisphere damage, or whether the ability exists in undamaged language systems. We sought to test these possibilities by studying auditory comprehension in acute left versus right hemisphere deactivation during Wada procedures. A series of 20 patients undergoing clinically indicated Wada procedures were asked to listen to an auditorily presented stimulus word, and then point to its matching picture on a card that contained the target picture, a semantic foil, a phonemic foil, and an unrelated foil. This task was performed under three conditions, baseline, during left carotid injection of sodium amytal, and during right carotid injection of sodium amytal. Overall, left hemisphere injection led to a significantly higher error rate than right hemisphere injection. However, consistent with lesion work, the majority (75%) of these errors were semantic in nature. These findings suggest that auditory comprehension deficits are predominantly semantic in nature, even following acute left hemisphere disruption. This, in turn, supports the hypothesis that the right hemisphere is capable of speech sound processing in the intact brain.     

Hemispheric lateralization in 47,XXY Klinefelter's syndrome boys

Thirty-two boys with a 47,XXY karyotype were compared with chromosomally normal male controls in their performance on six tasks of hemispheric specialization. The results revealed that the 47,XXY subjects had smaller asymmetries on left hemisphere tasks and larger asymmetries on right hemisphere tasks than controls. Analyses of individual right and left side scores revealed that the atypical lateral asymmetries of the 47,XXYs were due to a shift toward greater right hemisphere involvement on four of the six measures. It was postulated that the slower fetal growth rates of the extra X chromosome group might contribute to their atypical hemispheric specialization and the failure of their left hemisphere to gain dominance over their right in language processing.     

A simple technique for lateralizing visual input that allows prolonged viewing

A simplified technique is described for obtaining lateralization of visual input with prolonged viewing. This technique is based on the presence of constant normal lateral limits for horizontal rotation of the eyes with reference to the head. With head movement prevented by use of a standard bite bar, and the eyes rotated to the left and held at their lateral limit, the temporal half of the visual field of the left eye may be used for lateralized input to the right hemisphere or vice versa for input to the left hemisphere. Any form of visual stimuli or visually monitored task can be used if confined within one of the extreme temporal hemifields. In comparison to previous methods, this technique is technically simple, inexpensive, without significant risk or discomfort to the subject, readily applicable to normal and various brain-lesioned subjects, and permits prolonged in-depth viewing. An alternative version of this technique uses a stabilized spectacle frame fitted with adjustable central occluders set to allow vision through only one or both of the extreme temporal hemifields.     

Aphasic patients exhibit a reversal of hemispheric asymmetries in categorical color discrimination

Patients with left hemisphere (LH) or right hemisphere (RH) brain injury due to stroke were tested on a speeded, color discrimination task in which two factors were manipulated: (1) the categorical relationship between the target and the distracters and (2) the visual field in which the target was presented. Similar to controls, the RH patients were faster in detecting targets in the right visual field when the target and distracters had different color names compared to when their names were the same. This effect was absent in the LH patients, consistent with the hypothesis that injury to the left hemisphere handicaps the automatic activation of lexical codes. Moreover, the LH patients showed a reversed effect, such that the advantage of different target-distracter names was now evident for targets in the left visual field. This reversal may suggest a reorganization of the color lexicon in the right hemisphere following left hemisphere brain injury and/or the unmasking of a heightened right hemisphere sensitivity to color categories.     

Evidence for right hemisphere phonology in a backward masking task

The extent to which orthographic and phonological processes are available during the initial moments of word recognition within each hemisphere is under specified, particularly for the right hemisphere. Few studies have investigated whether each hemisphere uses orthography and phonology under constraints that restrict the viewing time of words and reduce overt phonological demands. The current study used backward masking in the divided visual field paradigm to explore hemisphere differences in the availability of orthographic and phonological word recognition processes. A 20 ms and 60 ms SOA were used to track the time course of how these processes develop during pre-lexical moments of word recognition. Nonword masks varied in similarity to the target words such that there were four types: orthographically and phonologically similar, orthographically but not phonologically similar, phonologically but not orthographically similar and unrelated. The results showed the left hemisphere has access to both orthography and phonology early in the word recognition process. With more time to process the stimulus, the left hemisphere is able to use phonology which benefits word recognition to a larger extent than orthography. The right hemisphere also demonstrates access to both orthography and phonology in the initial moments of word recognition, however, orthographic similarity improves word recognition to a greater extent than phonological similarity.     

Remembering 1500 pictures: the right hemisphere remembers better than the left

We hypothesized that the right hemisphere would be superior to the left hemisphere in remembering having seen a specific picture before, given its superiority in perceptually encoding specific aspects of visual form. A large set of pictures (N=1500) of animals, human faces, artifacts, landscapes, and art paintings were shown for 2s in central vision, or tachistoscopically (for 100ms) in each half visual field, to normal participants who were then tested 1-6 days later for their recognition. Images that were presented initially to the right hemisphere were better recognized than those presented to the left hemisphere. These results, obtained with participants with intact brains, large number of stimuli, and long retention delays, are consistent with previously described hemispheric differences in the memory of split-brain patients.     

Social and affective impairments are important recovery after acquired stroke in childhood

Despite a congenital stroke, overall intelligence at school age is generally within the normal range. Language acquisition problems are more prominent when children are younger (<5 years of age) than when they are older. They are present after both right and left lesions, but appear to have different features. They are less apparent than in the child with a developmental language disorder. Acquired aphasia in childhood results in subtle and often persisting deficits. Children with congenital strokes are at risk for behavioral and psychiatric problems. Those with congenital right hemisphere strokes appear to be more difficult infants, but there is no clear side of lesion effect in older children. Children with congenital right hemisphere strokes have more prominent spatial difficulties than their left lesion counterparts. Evaluating both the process and the product highlights this. Increasing the difficulty of the task often brings out deficits in the right lesion group even when they seemingly recovered.     

A valence-specific lateral bias for discriminating emotional facial expressions in free field

Findings from subjects with unilateral brain damage, as well as from normal subjects studied with tachistoscopic paradigms, argue that emotion is processed differently by each brain hemisphere. An open question concerns the extent to which such lateralised processing might occur under natural, freeviewing conditions. To explore this issue, we asked 28 normal subjects to discriminate emotions expressed by pairs of faces shown side-by-side, with no time or viewing constraints. Images of neutral expressions were shown paired with morphed images of very faint emotional expressions (happiness, surprise, disgust, fear, anger, or sadness). We found a surprising and robust laterality effect: When discriminating negative emotional expressions, subjects performed significantly better when the emotional face was to the left of the neutral face; conversely, when discriminating positive expressions, subjects performed better when the emotional face was to the right. We interpret this valence-specific laterality effect as consistent with the idea that the right hemisphere is specialised to process negative emotions, whereas the left is specialised to process positive emotions. The findings have important implications for how humans perceive facial emotion under natural conditions.     

Unilateral hemispheric activation does not affect free-viewing perceptual asymmetries

Strong leftward perceptual biases have been reported for the selection of the darker of two left/right mirror-reversed luminance gradients under free-viewing conditions. This study investigated the effect of unilateral hemispheric activation on this leftward bias in two groups of dextrals (N = 52 and N = 24). In Experiment 1, activation was manipulated by asking participants to tap with their left or right fingers along their midline. In Experiment 2, participants clenched their left or right hands in their respective hemispaces. Participants selected the stimulus that was darker on the left-hand side 73% of the time. Despite manipulations of activation strength and hemispace, activation had no effect on the asymmetry. If activation was important, the leftward bias should have been enhanced when the left hand/right hemisphere was active and reduced (or reversed) when the right hand/left hemisphere was active. The contribution of left-to-right scanning biases to free-viewing perceptual asymmetries is discussed as an alternative.     

Apparent shift in visual field preference after unilateral stroke

Patients with either a left- or a right-hemisphere stroke lesion scored higher in tasks of word-picture matching and of nonverbal shape matching when information was presented tachistoscopically (120 msec) to the visual field (VF) projecting to their undamaged hemisphere. Left-hemisphere stroke patients (n = 13) were dissociated from right-hemisphere stroke patients (n = 15) by low word recognition from memory and by low right VF but nearly normal left VF accuracy in word-picture matching or shape matching; the former appeared to rely upon processing of word meaning by the right hemisphere. In contrast, right-stroke patients had higher right than left VF scores in both tasks, and their discrimination of nonverbal shapes via the right VF was not different from that of controls (n = 15). Preferred processing by the VF projecting to the undamaged hemisphere appeared as a shift in perceptual asymmetry but may indicate, in support of a "direct access" model, that each hemisphere responds more or less efficiently to word and to nonverbal shape discriminations.     

The cost of action miscues: hemispheric asymmetries

Adaptive behaviors require preparation and when necessary inhibition or alteration of actions. The right hemisphere has been posited to be dominant for preparatory motor activation. This experiment was designed to learn if there are hemispheric asymmetries in the control of altered plans of actions. Cues, both valid and invalid, which indicate the hand most likely to be called onto respond, as well as the imperative stimuli that indicate the actual response hand, were presented to either the right or left visual fields of 14 normal right handed participants. The delay after a miscue is dependent on the time taken to inhibit the premotor and motor systems of the incorrectly activated hemisphere, as well as to activate the motor systems of the opposite hemisphere, which might have been interhemispherically inhibited by this miscue. Analyses of reaction times revealed that miscues presented in left hemispace (right hemisphere) cost more time than those miscues presented in right hemispace (left hemisphere), suggesting that activation of the preparatory systems controlled by the right hemisphere may take longer to reverse than those controlled by the left hemisphere. This asymmetry may be related to asymmetries in the strength of hemispheric activation with contralateral inhibition.     

The influence of emotional faces on the spatial allocation of attention

Background objectives: Studies suggest that the right hemisphere is dominant for emotional facial recognition. In addition, whereas some studies suggest the right hemisphere mediates the processing of all emotions (dominance hypothesis), other studies suggest that the left hemisphere mediates positive emotions the right mediates negative emotions (valence hypothesis). Since each hemisphere primarily attends to contralateral space, the goals of this study was to learn if emotional faces would induce a leftward deviation of attention and if the valence of facial emotional stimuli can influence the normal viewer’s spatial direction of attention. Methods: Seventeen normal right handed participants were asked to bisect horizontal lines that had all combinations of sad, happy or neutral faces at ends of these lines. During this task the subjects were never requested to look at these faces and there were no task demands that depended on viewing these faces. Results: Presentation of emotional faces induced a greater leftward deviation compared to neutral faces, independent of where (spatial position) these faces were presented. However, faces portraying negative emotions tended to induce a greater leftward bias than positive emotions. Conclusions: Independent of location, the presence of emotional faces influenced the spatial allocation of attention, such that normal subjects shift the direction of their attention toward left hemispace and this attentional shift appears to be greater for negative (sad) than positive faces (happy).