Lateralized interhemispheric transfer of color cues: evidence for dynamic coding principles of visual lateralization in pigeons

Visual feature discrimination tasks in pigeons reveal a right eye/left hemisphere dominance at the population level. Anatomical studies and lesion data show this visual lateralization to be related to asymmetries of the tectofugal system, which ascends from the tectum over the n. rotundus to the forebrain. Anatomically, this system is characterized by numerous morphological and connectional asymmetries which result in a bilateral visual representation in the dominant left hemisphere and a mostly contralateral representation in the subdominant right hemisphere. Ontogenetically, visual lateralization starts with an asymmetrical embryonic position within the egg, which leads to asymmetries of light stimulation. Differences in exposure to light stimulation between the eyes result in activity differences between the ascending tectofugal pathways of the left and the right hemisphere, which are transcribed during a critical time span into morphological asymmetries. The asymmetries established after this transient period finally start to determine the lateralized processes of the visual system for the entire life span of the individual. We now can show that these anatomical lateralizations are accompanied by asymmetries of interocular transfer, which enable a faster shift of learned color cues from the dominant right to the left eye than vice versa. In summary, our data provide evidence that cerebral asymmetries are based both on "static" anatomical and on "dynamic" process-dependent principles.     

Right-hemispheric dominance for processing extended non-linguistic frequency transitions

The left hemisphere is specialized for most linguistic tasks and the right hemisphere is specialized for many non-linguistic tasks, but the cause of these functional asymmetries is unknown. One of the stimulus factors that appears to influence these asymmetries is the rate at which stimuli change. In the present experiment, 41 participants completed the Fused Dichotic Words Test (FDWT) and a non-linguistic Frequency Transition Task (FTT) wherein the Frequency Transitions (FTs) were either rapid (40 ms) or relatively slow (200 ms). There was a right hemisphere advantage for slow FTs when the change was at the front of the stimulus, but no corresponding left hemisphere advantage for the rapid FTs. There was no relationship between either FTT and the left hemisphere advantage exhibited on the FDWT. This finding provides support for the position that the right hemisphere dominates tasks that require temporal processing over relatively long periods of time.     

Hemispheric asymmetries for gap detection depend on noise type

Studies of temporal processing asymmetries in the auditory modality have not produced consistent results. Some investigators have found a left hemisphere advantage, but others have failed to replicate this result. The present experiment investigated the possibility that differing properties of the noise employed between these experiments could be responsible for the conflicting results. Short bursts (300 ms) of white or brown noise were delivered monaurally to 42 participants. Half of the stimuli contained 3-5 ms gaps of silence. A right ear (left hemisphere) advantage in accuracy was observed in the white noise condition, but there was no such difference in the brown noise condition. This result demonstrates that the different acoustic properties of the stimuli between experiments can account for some of the discrepancies in their findings, and supports the position that the left hemisphere is superior at processing rapid temporal changes.     

Are the predictions of the dynamic dominance model of laterality applicable to the lower limbs?

Investigation of manual actions has supported the proposition that the right and left cerebral hemispheres have complementary specializations relevant for movement control. To test the extent to which hemisphere specialization affect lower limb control, we compared performance between the legs in two motor tasks. A pedal aiming task was employed to test the notion of left hemisphere specialization for dynamic control, and unipedal balance was employed to test the notion of right hemisphere specialization for impedance control. Evaluation was conducted on young adults, in the contexts of separate (Experiment 1) and integrated (Experiment 2) performance of the probing tasks. Results from the aiming task showed equivalent movement linearity toward the target between the right and left feet across experiments. Analysis of unipedal balance revealed that increased stance stability when supported on the left leg was observed when performing simultaneously the aiming task with the contralateral foot, but not in the context of isolated task performance. These results are inconsistent with the proposition of left hemisphere specialization for dynamic control in the lower limbs, and suggest that specialization of the right hemisphere for impedance control can be observed in balance control when stance is associated with voluntary movements of the contralateral lower limb.     

Spatial constraints on attention to speech in the blind

When sighted persons try to identify one of two speech utterances coming from different directions, they display both a frontal position advantage, i.e., better recognition of inputs from the front than of those from the rear, and a right-side advantage, better recognition of inputs from the right than of those from the left. The present study demonstrates a dissociation of the two effects in blind subjects (N = 10) who showed no frontal position advantage together with a right-side advantage superior to that of control sighted subjects (N = 16). There was no systematic difference between congenitally blind subjects and noncongenitals. The absence of frontal position advantage in the blind is consistent with the notion that this effect originates in the habit of sighted listeners to orient toward the source of heard speech. The occurrence of at least normal right-side advantage in the blind does not support recent suggestions of reduced lateralization of language functions in such subjects.     

Manual asymmetries in bimanual reaching: the influence of spatial compatibility and visuospatial attention

The goal of the present investigation was to explore the possible expression of hemispheric-specific processing during the planning and execution of a bimanual reaching task. Participants (N = 9) completed 80 bimanual reaching movements (requiring simultaneous, bilateral production of arm movements) to peripherally presented targets while selectively attending to either their left or right hand. Further, targets were presented in spatially compatible (ipsilateral to the aiming limb) and incompatible (contralateral to the aiming limb) response contexts. It was found that the left hand exhibited temporal superiority over the right hand in the response planning phase of bimanual reaching, indicating a left hand/right hemisphere advantage in the preparation of a bimanual response. During response execution, and consistent with the view that interhemispheric processing time (Barthelemy & Boulinguez, 2002) or biomechanical constraints (Carey, Hargreaves, & Goodale, 1996) generate temporal delays, longer movement times were observed in response to spatially incompatible target positions. However, no hemisphere-specific benefit was demonstrated for response execution. Based on these findings, we propose lateralized processing is present at the time of response planning (i.e., left hand/right hemisphere processing advantage); however, lateralized specialization appears to be annulled during dynamic execution of a bimanual reaching task.     

Effect of temporal constraints on hemispheric asymmetries during spatial frequency processing

Studies on functional hemispheric asymmetries have suggested that the right vs. left hemisphere should be predominantly involved in low vs. high spatial frequency (SF) analysis, respectively. By manipulating exposure duration of filtered natural scene images, we examined whether the temporal characteristics of SF analysis (i.e., the temporal precedence of low on high spatial frequencies) may interfere with hemispheric specialization. Results showed the classical hemispheric specialization pattern for brief exposure duration and a trend to a right hemisphere advantage irrespective of the SF content for longer exposure duration. The present study suggests that the hemispheric specialization pattern for visual information processing should be considered as a dynamic system, wherein the superiority of one hemisphere over the other could change according to the level of temporal constraints: the higher the temporal constraints of the task, the more the hemispheres are specialized in SF processing.     

Cerebral asymmetry for American Sign Language: The effects of moving stimuli

Previous studies of cerebral asymmetry for the perception of American Sign Language (ASL) have used only static representations of signs; in this study we present moving signs. Congenitally deaf, native ASL signers identified moving signs, static representations of signs, and English words. The stimuli were presented rapidly by motion picture to each visual hemifield. Normally hearing English speakers also identified the English words. Consistent with previous findings, both the deaf and the hearing subjects showed a left-hemisphere advantage to the English words; likewise, the deaf subjects showed a right hemisphere advantage to the statically presented signs. With the moving signs, the deaf showed no lateral asymmetry. The shift from right dominance to a more balanced hemispheric involvement with the change from static to moving signs is consistent with Kimura's position that the left hemisphere predominates in the analysis of skilled motor sequencing (Kimura 1976). The results also indicate that ASL may be more bilaterally represented than is English and that the spatial component of language stimuli can greatly influence lateral asymmetries.     

Manual asymmetries in the preparation and control of goal-directed movements

The primary purpose of this experiment was to determine if left hand reaction time advantages in manual aiming result from a right hemisphere attentional advantage or an early right hemisphere role in movement preparation. Right-handed participants were required to either make rapid goal-directed movements to small targets or simply lift their hand upon target illumination. The amount of advance information about the target for a particular trial was manipulated by precuing a subset of potential targets prior to the reaction time interval. When participants were required to make aiming movements to targets in left space, the left hand enjoyed a reaction advantage that was not present for aiming in right space or simple finger lifts. This advantage was independent of the amount or type of advance information provided by the precue. This finding supports the movement planning hypothesis. With respect to movement execution, participants completed their aiming movements more quickly when aiming with their right hand, particularly in right space. This right hand advantage in right space was due to the time required to decelerate the movement and to make feedback-based adjustments late in the movement trajectory.     

Name and face matching in one or two visual fields: a test of models of hemispheric specialization

In two experiments a name and a face (each male or female) were simultaneously flashed to either the same or opposite visual fields (left or right), for matching congruent (same sex) or incongruent (opposite sex), to test the predictions of various models of hemispheric specialization. While overall best performance occurred with a face in the left visual field (LVF) and a name in the right visual field (RVF), and worst with the opposite configuration, the general pattern of results was incompatible with either a direct access model or an activational/attentional account. The results were, however, most compatible with the predictions of a semispecialized hemispheres account, whereby cerebral asymmetries are seen as relative rather than absolute, either hemisphere being capable of processing either kind of material (verbal or visuospatial), but to different levels of efficiency. However, despite the fact that the stimulus materials had previously been shown to produce stable and consistent lateral asymmetries in the predicted directions when presented in isolation, in the composite, integrative matching task the position of the name seemed to be the major determinant of the resultant asymmetries. It would seem therefore that when such stimuli are to be cross matched, either left hemisphere (language) processes somehow dominate right hemisphere (visuospatial) processing (though not in the way that would be predicted by a simple activational/attentional account) or the left hemisphere's greater capacity predominates.     

Evidence for cerebral asymmetries in a finger sequencing task

Visual input was lateralized using a specially designed contact lens system. Subjects performed a sequence of two keypresses in response to a light stimulus with either the left or the right hand in a choice reaction time paradigm. Two choice reaction time conditions were used: (A) hand certainty, sequence uncertainty and (B) hand uncertainty, sequence certainty. Reaction time (RT) results indicate that there are no significant differences between the left and right hemisphere in selecting a sequential response in either of the two conditions. Interfinger time (IFT) results show a relative left eye (right hemisphere)-left hand advantage when there was hand certainty, sequence uncertainty and a relative left eye (right hemisphere) disadvantage for both hands when there was hand uncertainty, sequence certainty. The RT results do not support the concept of a center in the left hemisphere for selection of the components of a two-element sequential keypress, prior to movement initiation. However, the IFT results indicate that there are differences in the processing ability of the left and right hemispheres in a sequencing task, after movement initiation.     

Hemispheric differences in object identification

Although the right hemisphere is thought to be preferentially involved in visuospatial processing, the specialization of the two hemispheres with respect to object identification is unclear. The present study investigated the effects of hemifield presentation on object and word identification by presenting objects (Experiment 1) and words (Experiment 2) in a rapid visual stream of distracters. In Experiment 1, object images presented in the left visual field (i.e., to the right hemisphere) were identified with shorter display times. In addition, the left visual field advantage was greater for inverted objects. In Experiment 2, words presented in the right visual field (i.e., to the left hemisphere) under similar conditions were identified with shorter display times. These results support the idea that the right hemisphere is specialized with regard to object identification.     

Kinematic Analyses of Manual Asymmetries in Visual Aiming Movements

The right hand advantage has been thought to arise from the greater efficiency of the right hand/left hemisphere system in processing visual feedback information. This hypothesis was examined using kinematic analyses of aiming performance, focusing particularly on time after peak velocity which has been shown to be sensitive to visual feedback processing demands. Eight right-handed subjects pointed at two targets with their left and right hands with or without vision available and either as accurately or as fast as possible. Pointing errors and movement time were found to be smaller with the right hand. Analyses of the temporal componenets of movement time revealed that the hands differed only in time after peak velocity (in deceleration), with the right hand spending significantly less time. This advantage for the right hand, however, was apparent whether or not vision was available and only when accuracy was emphasized in performance. These findings suggest that the right hand system may be more efficient at processing feedback information whether this be visual or nonvisual (e.g., proprioceptive).     

The Contribution of Attention to the Right Visual Field Advantage for Word Recognition

Perceptual asymmetries have been explained by structural, attentional bias and attentional advantage models. Structural models focus on asymmetries in the physical access information has to the hemispheres, whereas attentional models focus on asymmetries in the operation of attentional processes. A series of experiments was conducted to assess the contribution of attentional mechanisms to the right visual field (RVF) advantage found for word recognition. Valid, invalid and neutral peripheral cues were presented at a variety of stimulus onset asynchronies to manipulate spatial attention. Results indicated a significant RVF advantage and cueing effect. The effect of the cue was stronger for the left visual field than the RVF. This interaction supports the attentional advantage model which suggests that the left hemisphere requires less attention to process words. The attentional asymmetry is interpreted in terms of the different word processing styles used by the left and right hemispheres. These results have ramifications for the methodology used in divided visual field research and the interpretation of this research.     

Hemispace Asymmetries and Laterality Effects in Arm Positioning

Hemispace asymmetries and laterality effects were examined on an arm positioning reproduction task. Sixteen male subjects were asked to reproduce both abductive and adductive positioning movements with the left or right arm within either the left or the right hemispace. Hemispace was manipulated using a 90 degrees head-rotation paradigm. A left hemispace advantage in positioning accuracy was predicted for both left and right arm movements on the grounds that the perceptual-motor control of positioning movements made in left hemispace is primarily mediated by the right hemisphere which is known to be advantageous for tasks which are spatial in nature (Heilman, Bowers, & Watson, 1984). No arm laterality effects were predicted to occur because the proximal musculature involved in the control of arm movements is innervated from both contralateral and ipsilateral cerebral hemispheres (Brinkman & Kuypers, 1973). Results showed that the predicted left hemispace advantage was evident for the right arm on the positioning variability measure alone, whereas it was absent for all other possible conditions on all error measures. Laterality (arm) effects were absent as predicted. The experiment also demonstrated a greater degradation of reproduction performance under the ′crossed" arm-hemispace conditions than under the ′uncrossed" conditions. A plausible explanation for the uncrossed advantage for the task is that under normal conditions, a single hemisphere is primarily responsible for both controlling the contralateral arm and directing attention to the contralateral hemispace, and consequently potential interhemispheric interference is minimized. A clear response bias effect in movement reproduction was also evident as a function of the direction of concurrent arm movement and head rotation. Arm movements made in the same direction as head rotation were systematically undershot in reproduction to a much greater degree than arm movements made in the opposite direction to head rotation.     

Cerebral Organization of Motor Imagery: Contralateral Control of Grip Selection in Mentally Represented Prehension

The principle of contralateral organization of the visual and motor systems was exploited to investigate contributions of the cerebral hemispheres to the mental representation of prehension in healthy, right-handed human subjects. Graphically rendered dowels were presented to either the left or right visual field in a variety of different orientations, and times to determine whether an underhand or overhand grip would be preferred for engaging these stimuli were measured. Although no actual reaching movements were performed, a significant advantage in grip-selection time was found when information was presented to the cerebral hemisphere contralateral to the designated response hand. Results are consistent with the position that motor imagery recruits neurocognitive mechanisms involved in movement planning. More precisely, these findings indicate that processes within each cerebral hemisphere participate in mentally representing object-oriented actions of the contralateral hand.     

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.     

Asymmetries for the visual expression and perception of speech

This study explored asymmetries for movement, expression and perception of visual speech. Sixteen dextral models were videoed as they articulated: 'bat,' 'cat,' 'fat,' and 'sat.' Measurements revealed that the right side of the mouth was opened wider and for a longer period than the left. The asymmetry was accentuated at the beginning and ends of the vocalization and was attenuated for words where the lips did not articulate the first consonant. To measure asymmetries in expressivity, 20 dextral observers watched silent videos and reported what was said. The model's mouth was covered so that the left, right or both sides were visible. Fewer errors were made when the right mouth was visible compared to the left--suggesting that the right side is more visually expressive of speech. Investigation of asymmetries in perception using mirror-reversed clips revealed that participants did not preferentially attend to one side of the speaker's face. A correlational analysis revealed an association between movement and expressivity whereby a more motile right mouth led to stronger visual expressivity of the right mouth. The asymmetries are most likely driven by left hemisphere specialization for language, which causes a rightward motoric bias.     

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.