Hemispheric differences in attentional orienting by social cues

Research points to a right hemisphere bias for processing social stimuli. Hemispheric specialization for attention shifts cued by social stimuli, however, has been rarely studied. We examined the capacity of each hemisphere to orient attention in response to social and nonsocial cues using a lateralized spatial cueing paradigm. We compared the up/down orienting effects of eye gaze cues, arrow cues, and peripheral cues (change in luminance). Results revealed similar cueing effects in each visual field for nonsocial cues, but asymmetric effects for social cues. At both short (150 ms) and long (950 ms) cue-target intervals, gaze cueing was significant in the LVF, but not in the RVF. Thus, there is a right hemisphere bias for attentional orienting cued by social stimuli, but not for attentional orienting cued by nonsocial stimuli. This supports a theory of a separate neural system for socially cued orienting of attention, as well as a theory of separate parallel and simultaneous neural systems for attention in the two cerebral hemispheres.     

Hemispheric specialization for local and global processing.

In an experiment on normal adults it was demonstrated that local processing of a linguistic stimulus is superior in the left hemisphere, whereas global processing does not appear to be strongly lateralized.     

The time course of hemispheric differences in categorical and coordinate spatial processing

Spatial relations between objects can be represented either categorically or coordinately. The metric, coordinate representation is associated with predominant right hemisphere activity, while the abstract, qualitative categorical representation is thought to be processed more in the left hemisphere [Kosslyn, S. M. (1987). Seeing and imagining in the cerebral hemispheres: A computational analysis. Psychological Review, 94, 148-175]. This hypothesized lateralization effect has been found in a number of studies, along with indications that specific task demands can be crucial for these outcomes. In the current experiment a new visual half field task was used which explores these hemispheric differences and their time course by means of a match-to-sample design. Within retention intervals that were brief (500 ms), intermediate (2000 ms), or long (5000 ms), the processing of categorical and coordinate representations was studied. In the 500 ms interval, the hemispheric effect suggested by Kosslyn (1987) was found, but in the longer intervals it was absent. This pattern of the lateralization effect is proposed to be caused by the differential effect the retention interval has on coordinate and categorical representations. Coordinate spatial relations appear susceptible to changes in retention interval and decay very quickly over time, congruent with previous findings about accurate location memory. The processing of categorical spatial relations showed less decay and only between 2000 ms and 5000 ms. Qualitative self reports suggest that the decay found for categorical relations might be caused by a switch from a visual to a more verbal memorization strategy.     

Lateralised processing of positive facial emotion: sex differences in strength of hemispheric dominance

Sex differences in lateralisation have been examined frequently, but have found varying and contradictory results. The experiment presented in this paper examines the lateralisation of processing positive facial emotion in 276 right handed undergraduates (138 males, 138 females). All participants completed two behavioural tests of lateralisation: a handedness preference questionnaire and a chimeric faces emotion judgement task, which measured strength of lateralisation for the perception of positive facial emotion. A highly significant difference was found for the chimeric faces task only: males were more strongly lateralised than females, although both males and females tended to be right hemisphere dominant. The results suggest that females are more bilaterally distributed and hence have greater access to mechanisms located in each hemisphere.     

Hemispheric integration and differences in perception of a line-motion illusion in the divided brain

Five people lacking the corpus callosum (two callosotomized, three with agenesis of the corpus callosum) and neurologically normal subjects were shown vertical lines that appeared instantaneously between pairs of rectangles in one or other visual field. When one of the rectangles flashed prior to the presentation of the line, and the line was in the same visual field, all subjects perceived the line as spreading from the flashed rectangle to the other. Normal subjects and one of the callosotomized subjects showed a slight but significant right visual-field advantage, perhaps reflecting a left-hemispheric superiority in processing rapid temporal events. The illusion was also induced when the line and the flash were in opposite visual fields in one of the callosotomized, one of the acallosal subjects, and about half of the normal subjects, implying interhemispheric integration even in the absence of the corpus callosum.     

On the role of response conflicts and stimulus position for hemispheric differences in global/local processing: an ERP study

It is widely assumed that the local and global levels of hierarchical stimuli are processed more efficiently in the left and right cerebral hemispheres, respectively. However, corresponding effects were not observed under all circumstances. In ERP studies, they occurred more often with centrally presented stimuli than with laterally presented ones, whereas reaction-time studies revealed that a response conflict between the levels is relevant. The present study examines which of these two factors is more important by presenting conflicting and non-conflicting stimuli to the left or right visual field and recording ERPs as well as collecting behavioral data. If a central stimulus position is crucial, then no effects should show up. Contrary to this prediction, the expected hemispheric differences were observed in the behavioral data as well as in the later occurring (N2 and P3) ERP amplitudes. However, in all variables, the effects were more pronounced for conflicting stimuli. The results suggest that response conflicts are more important for obtaining hemispheric differences in global/local processing than a central stimulus presentation. This is interpreted in the way that hemispheric differences vary with respect to the stimulus representation that is needed to select a proper response.     

Hemispheric asymmetry in spatial attention across the menstrual cycle

Functional cerebral asymmetries (FCAs) are known to fluctuate across the menstrual cycle. The mechanisms of these sex hormonal modulations are poorly understood. It has been suggested that gonadal steroid hormones might suppress or specifically activate one hemisphere. However, recent studies suggest that high levels of gonadal steroid hormones reduce FCAs by its modulating effects on cortico-cortical transmission. To investigate the activating effects of gonadal steroid hormones on the interhemispheric interaction, a visual line-bisection task was administered to normally cycling women during menses and the midluteal cycle phase as well as to similar-aged healthy men. The results replicate previous findings of a sex difference in line-bisection as a function of hand-use and show that the hand-use effect fluctuates across the menstrual cycle. High levels of estradiol during the midluteal phase were related to a decrease of the hand-use effect. It is concluded that cycle-related fluctuations in levels of gonadal steroid hormones affect hemispheric asymmetry of spatial attention, presumably by interhemispheric spreading of neuronal activation.     

Hemispheric asymmetries in categorical perception of orientation in infants and adults

Orientation CP is the faster or more accurate discrimination of two orientations from different categories (e.g., oblique1 and vertical1) compared to two orientations from the same category (e.g., oblique1 and oblique2), even when the degree of difference is equated across conditions. Here, we assess whether there are hemispheric asymmetries in this effect for adults and 5-month-old infants. Experiment 1 identified the location of the vertical-oblique category boundary. Experiment 2, using a visual search task with oriented lines found that adult search was more accurate when the target and distractors were from different orientation categories, compared to targets and distractors of an equivalent physical difference taken from the same category. This effect was stronger for targets lateralized to the left visual field (LVF) than the right visual field (RVF), indicating a right hemisphere (RH) bias in adult orientation CP. Experiment 3, replicated the RH bias using different stimuli and also investigated the impact of visual and verbal interference on the category effect. Experiment 4, using the same visual search task, found that infant search was also faster when the target and distractors were from different orientation categories than the same, yet this category effect was stronger for RVF than LVF lateralized targets, indicating a LH bias in orientation CP at 5 months. These findings are contrasted to equivalent studies on the lateralization of color CP (e.g., Gilbert, Regier, Kay, & Ivry, 2005). The implications for theories on the contribution of the left and right hemispheres of the infant and adult brain to categorical computations are discussed.     

Differential processing of hierarchical visual stimuli in young and older healthy adults: implications for pathology

Hierarchical figures in which large (global) forms are constructed from smaller (local) forms (Navon, 1977) have proved valuable in studies of perceptual organisation and hemispheric specialisation in both healthy volunteers and a wide range of neurological and psychiatric patients. In studies using Navon figures, normal young adults typically identify global forms faster than local forms. When the global and local forms are incongruent (e.g., a large E made of smaller Rs), global forms often interfere with local form identification more than vice versa. In two conditions on the same subjects, we contrasted the performance of young (mean age 22 years) and older (mean age 58 years) healthy volunteers on global and local processing. In the directed attention task, subjects were instructed to detect a target letter that occurred at the prespecified local or global level. The young subjects showed, as expected, faster reaction times (RTs) to detect global targets. In contrast, the older subjects showed significantly faster RTs to the local targets. Likewise, in a divided attention task, in which subjects were instructed to detect a target letter that could occur at either the local or the global level, the young adults were slightly quicker to detect the global targets and the older subjects were significantly quicker to detect the local targets. Error rates were generally low and there was no significant speed/accuracy trade-off in either condition. The observed local precedence effects in healthy older subjects were unexpected and are discussed in reference to previous work on differential hemispheric aging. That work has suggested that the left hemisphere is preferentially biased toward local processing and ages relatively slowly while the right hemisphere is biased toward global processing and ages relatively quickly. The implications of such putative differential aging for the interpretation of pathological local/global processing in neurological and psychiatric diseases are also emphasised.     

Hemispheric differences in global versus local processing of hierarchical visual stimuli by normal subjects: new data and a meta-analysis of previous studies

Twenty-four normal right-handed male subjects classified hierarchical visual stimuli (small letters arranged to form larger letters), which were presented separately to each cerebral hemisphere. Previous studies using similar paradigms have yielded mixed evidence for the specialization of local, analytic processing in the left hemisphere and global, holistic processing in the right hemisphere. No statistically significant evidence for such a pattern of specialization was evident in the results of the present study. However, when meta-analytic techniques were used to combine the present results with those from previous studies, there was evidence that the hypothesized left-hemisphere/local, right-hemisphere/global specialization does in fact exist.     

Hemispheric differences in processing dichotic meaningful and non-meaningful words

Classic dichotic-listening paradigms reveal a right-ear advantage (REA) for speech sounds as compared to non-speech sounds. This REA is assumed to be associated with a left-hemisphere dominance for meaningful speech processing. This study objectively probed the relationship between ear advantage and hemispheric dominance in a dichotic-listening situation, using event-related potentials (ERPs). The mismatch negativity (MMN) and a late negativity (LN) were measured for bisyllabic meaningful words and non-meaningful pseudowords, which differed in their second syllable. Eighteen normal-hearing listeners were presented with a repeating diotic standard ([beI-gi:] or [leI-gi:]) and an occasional dichotic deviant (a standard presented to one ear and a deviant [beI-bi:], [beI-di:], [leI-bi:] or [leI-di:] presented to the opposite ear). As predicted there was a REA for meaningful words compared to non-meaningful words. Also, dipole source analysis suggested that dipole strength was stronger in the left than the right cortical region for meaningful words. However, there were differences in response within meaningful words as well as between meaningful and non-meaningful words which may be explained by the characteristics of embedded words and the position-specific probability of phoneme occurrence in words.     

The time course of spatial memory processing in the two hemispheres

Previous studies have shown that memories for positions are often distorted in systematic ways, indicating the influence of categorical positions codes which can bias responses in object-relocation tasks towards stored spatial prototypes. In the present study, we examined the time course of these categorical influences. Subjects had to relocate the position of a tachistoscopically presented dot within a circle, which could appear in either the left visual field (i.e. initially to the right hemisphere) or the right visual field (i.e. initially to the left hemisphere). Three retention intervals between presentation and relocation were used: 500, 2000 and 5000 ms. Performance was most accurate with left visual field/right hemisphere presentation. Systematic distortions were found for angular errors (dot relocations regressed towards the 45 degrees with a quadrant) as well as for radial errors (dots were replaced in the direction of the circle's circumference, and this more so when the dot was further away from the circumference). Importantly, these categorical biases became stronger with retention interval and initial left hemispheric processing. These results suggest that categorical spatial coding might be the default manner in which spatial information is remembered over time. Finally, the left hemisphere may play an important role for such a categorical spatial coding.     

Hemispheric specialization for categorical and coordinate spatial relations during an image generation task: evidence from children and adults

Hemispheric specialization of categorical and coordinate image generation was assessed in adults, 8-year-old and 10-year-old children. In a standardized image generation task, participants decided whether probes, presented in a blank grid (categorical task) or bracketed square (coordinate task), would have appeared on a previously studied letter. To ensure that participants mentally generated the target letter, probe location was varied. "Early" probes appeared on letter segments that are first produced when the letter is drawn; while "late" probes appeared on later produced segments. Like previous adult studies, the grid task elicited a left hemisphere "categorical" strategy; while the bracket task elicits a right hemisphere "coordinate" strategy. However, contrary to previous research, the results reveal the significant and complex effects of probe location on categorical and coordinate image generation abilities. Specifically, early probes elicited a strong right hemisphere advantage for both tasks across all ages, whereas late probes produced a left hemisphere dissociation between categorical and coordinate processing. The left hemisphere dissociation was evident only for 10-year-olds and adults, suggesting that younger children are not yet proficient in generating spatial representations.     

Hemispheric asymmetries in hierarchical stimulus processing are modulated by stimulus categories and their predictability

Hemispheric dominance has been behaviourally documented for the local (left hemisphere, LH) or global (right hemisphere, RH) processing of hierarchical letters. However, Fink et al. (1997) indicated that stimulus category modulates this hemispheric asymmetry. The purpose of this study was to investigate the influence of the category (letters versus objects) on hemispheric specialisation for global and local processing using a visual half-field presentation in a task where participants ignored whether the target appeared at the global or local level. In Experiment 1 we replicated the classic hemispheric asymmetry for global/local processing of hierarchical letters. In Experiment 2, which consisted of hierarchical object processing, a RH dominance for the local level was observed. In Experiment 3 a within-participant design was used where anticipation about the stimulus category was precluded, resulting in the classic RH and LH specialisations for global and local processing for both letter-based and object-based stimuli. Taken together, these results suggest that the highly demanding local processing stage engages one hemisphere more than the other, according to the lateralisation of cerebral networks specialised for stimulus category. In addition, the direction of lateralisation for the local level was also modulated by the predictability of the stimulus category.     

Hemispheric differences in processing tone frequency and amplitude modulations.

Transient frequency and amplitude modulations (FMs, AMs) of sound are requisite to speech recognition. We recorded whole-head magnetoencephalographic signals from seven subjects to binaural 620 ms 667 Hz tones, with 3, 30, or 300 ms FMs or AMs in the beginning or middle of the tone. Responses were significantly larger and earlier for FMs than AMs, for rapid than slow modulations, and for modulations at the beginning (BEG) than in the middle (MID) of the sound. BEG 3 ms FMs elicited strongest signals in the left and MID 3 ms FMs in the right hemisphere. Fast MID modulations produced significantly stronger responses in the right than left hemisphere. These differences may reflect different functions of the left and right hemisphere in speech perception.     

Hemispheric differences in the recruitment of semantic processing mechanisms

This study examined how the two cerebral hemispheres recruit semantic processing mechanisms by combining event-related potential measures and visual half-field methods in a word priming paradigm in which semantic strength and predictability were manipulated using lexically associated word pairs. Activation patterns on the late positive complex (LPC), linked to controlled aspects of processing, showed that previously documented left hemisphere (LH) processing benefits for word pairs with a weak forward but strong backward association stem from the ability to appreciate meaning relations in an order-independent fashion and/or strategically reorder them. Whereas there is a LH benefit for such strategic processing during comprehension in passive tasks, the present study further showed that the right hemisphere (RH) is also able to make use of these mechanisms when explicit semantic judgments are required. In both hemispheres, N400 responses, linked to initial semantic activation, were largely graded by association strength, with more amplitude reduction for forward associates and strong, symmetrically associated pairs compared to backward associates and matched weak, symmetrically associated pairs. However, responses to moderately associated pairs were more facilitated after initial presentation to the LH than to the RH. This pattern converges with sentence-processing findings that point to LH advantages for using context information to predict features of likely upcoming words. Together, the results suggest that an important basis for hemispheric asymmetries in language comprehension arises from when and how each uses top-down semantic mechanisms to shape initial semantic activation over time.     

Hemispheric Specialization in Children with Unilateral Epileptic Focus, With and Without Computed Tomography-Demonstrated Lesion

Summary: We evaluated 24 testable children with unilateral epileptic foci, with and without radiologically demonstrable lesions, to determine if the normal pattern of cerebral dominance is changed by the presence of an epileptic focus. Verbal and figural stimuli were presented tachistoscopically to the right and left visual hemifields to investigate the specific abilities of the two hemispheres. Three blocks of stimuli consisting of two-letter patterns, three-letter patterns, and meaningful two-syllable words were used as verbal stimuli to assess the abilities of the left hemisphere. A test consisting of localizing a dot on a 3x3 matrix was used to assess the abilities of the right hemisphere. Six normal children were chosen as controls. In all groups, specialization of the epileptic hemisphere was lost. The presence or absence of a radiologically demonstrable lesion did not influence this pattern.     

Hemispheric dissociation of reward processing in humans: Insights from deep brain stimulation

Rewards have various effects on human behavior and multiple representations in the human brain. Behaviorally, rewards notably enhance response vigor in incentive motivation paradigms and bias subsequent choices in instrumental learning paradigms. Neurally, rewards affect activity in different fronto-striatal regions attached to different motor effectors, for instance in left and right hemispheres for the two hands. Here we address the question of whether manipulating reward-related brain activity has local or general effects, with respect to behavioral paradigms and motor effectors. Neuronal activity was manipulated in a single hemisphere using unilateral deep brain stimulation (DBS) in patients with Parkinson's disease. Results suggest that DBS amplifies the representation of reward magnitude within the targeted hemisphere, so as to affect the behavior of the contralateral hand specifically. These unilateral DBS effects on behavior include both boosting incentive motivation and biasing instrumental choices. Furthermore, using computational modeling we show that DBS effects on incentive motivation can predict DBS effects on instrumental learning (or vice versa). Thus, we demonstrate the feasibility of causally manipulating reward-related neuronal activity in humans, in a manner that is specific to a class of motor effectors but that generalizes to different computational processes. As these findings proved independent from therapeutic effects on parkinsonian motor symptoms, they might provide insight into DBS impact on non-motor disorders, such as apathy or hypomania.