Crossed-uncrossed Difference Research Articles

Does CUD measure interhemispheric transfer time? The allocation of attention influences the Poffenberger effect

A simple reaction time (SRT) difference between responses to visual stimuli presented ipsilaterally and contralaterally to the responding hand, known as the Poffenberger effect or the crossed–uncrossed difference (CUD), has been commonly interpreted as a measure of interhemispheric transfer time (IHTT). However, the validity of this interpretation and the measure's reliability have been debated. The present study aimed at obtaining reliable evidence of the influence of spatial attention on the CUD, which would provide an argument against the classical interpretation of CUD. To meet the high statistical power requirements, over 100 thousand SRTs in total were collected from 12 participants. The task had three stimulus presentation conditions differing in the degree of stimulus location uncertainty: blocked (no uncertainty), randomized (full uncertainty), and mixed (25% uncertainty). The results showed robust effects of location uncertainty, proving spatial attention's contribution to the CUD. Further, we observed a strong visual-field asymmetry reflecting the right hemisphere specialization in target detection and spatial reorienting. Lastly, despite exceptional reliability of the component SRT measures, the CUD reliability was still too low to justify using this measure as an index of individual differences.

Motives and Laterality: Exploring the Links

ObjectivesWe explored associations between the needs for power, achievement, and affiliation and functional cerebral asymmetries (FCAs), guided by three established hypotheses about the nature of these associations.MethodsOne-hundred-and-seven participants completed picture-story measures of dispositional motives and activity inhibition (AI), a frequent moderator of motive-behavior associations, tasks measuring FCAs (line bisection, chimeric emotional face judgments, turning bias, perceptual and response asymmetries on the Poffenberger task), self-reported laterality preferences (handedness, footedness, ear and eye preference), and interhemispheric interaction (crossed-uncrossed difference). They also completed an experiment manipulating hand contractions (left, right, both, neither) while they worked on a second picture-story motive measure.ResultsDispositional power motivation was associated with stronger rightward asymmetry and less interhemispheric transfer in high-AI and stronger leftward asymmetry and more interhemispheric transfer in low-AI individuals. For the affiliation motive, findings were fewer and in the opposite direction of those for the power motive. These findings emerged for men, but not for women. Left- or right-hand contractions led to increases in power and achievement motivation, but not affiliation motivation. Only left-hand contractions led to decreased AI.ConclusionsWe discuss these findings in the context of sex-dimorphic organizing and activating effects of steroids on motives and laterality.

Underlying neural alpha frequency patterns associated with intra-hemispheric inhibition during an interhemispheric transfer task

Interhemispheric transfer measured via differences in right- or left-handed motoric responses to lateralized visual stimuli, known as the crossed-uncrossed difference (CUD), is one way of identifying patterns of processing that are vital for understanding the transfer of neural signals. Examination of interhemispheric transfer by means of the CUD is not entirely explained by simple measures of response time. Multiple processes contribute to wide variability observed in CUD reaction times. Prior research has suggested that intra-hemispheric inhibitory processes may be involved in regulation of speed of transfer. Our study examined electroencephalography recordings and time-locked alpha frequency activity while 18 participants responded to lateralized targets during performance of the Poffenberger Paradigm. Our results suggest that there are alpha frequency differences at fronto-central lateral electrodes based on target, hand-of-response, and receiving hemisphere. These findings suggest that early motoric inhibitory mechanisms may help explain the wide range of variability typically seen with the CUD.

Effects of stimulus pair orientation and hand switching on reaction time estimates of interhemispheric transfer.

Two behavioral estimates of interhemispheric transfer time, the crossed-uncrossed difference (CUD) and the unilateral field advantage (UFA), are thought to, respectively, index transfer of premotor and visual information across the corpus callosum in neurotypical participants. However, no attempt to manipulate visual and motor contingencies in a set of tasks while measuring the CUD and the UFA has yet been reported. In two go/no-go comparison experiments, stimulus pair orientations were manipulated. The hand of response changed after each correct response in the second, but not the first experiment. No correlation was found between the CUD and the UFA, supporting the hypothesis that these two measures index different types of information transfer across hemispheres. An effect of manipulation of stimulus pair orientation on UFAs was attributed to the homotopy of callosal fibers transferring visual information, while an effect of hand switching on CUDs was attributed mostly to spatial compatibility.

Visual and visuomotor interhemispheric transfer time in older adults

Older adults typically experience reductions in the structural integrity of the anterior channels of the corpus callosum. Despite preserved structural integrity in central and posterior channels, many studies have reported that interhemispheric transfer, a function attributed to these regions, is detrimentally affected by aging. In this study, we use a constrained event-related potential analysis in the theta and alpha frequency bands to determine whether interhemispheric transfer is affected in older adults. The crossed-uncrossed difference and lateralized visual evoked potentials were used to assess interhemispheric transfer in young (18–27) and older adults (63–80). We observed no differences in the crossed-uncrossed difference measure between young and older groups. Older adults appeared to have elongated transfer in the theta band potentials, but this effect was driven by shortened contralateral peak latencies, rather than delayed ipsilateral latencies. In the alpha band, there was a trend toward quicker transfer in older adults. We conclude that older adults do not experience elongated interhemispheric transfer in the visuomotor or visual domains and that these functions are likely attributed to posterior sections of the corpus callosum, which are unaffected by aging.

Vision of the body improves inter-hemispheric integration of tactile-motor responses

Sensory input from and motor output to the two sides of the body needs to be continuously integrated between the two cerebral hemispheres. This integration can be measured through its cost in terms of processing speed. In simple detection tasks, reaction times (RTs) are faster when stimuli are presented to the side of the body ipsilateral to the body part used to respond. This advantage – the contralateral-ipsilateral difference (also known as the crossed-uncrossed difference: CUD) – is thought to reflect inter-hemispheric interactions needed for sensorimotor information to be integrated between the two hemispheres. Several studies have shown that non-informative vision of the body enhances performance in tactile tasks. However, it is unknown whether the CUD can be similarly affected by vision. Here, we investigated whether the CUD is modulated by vision of the body (i.e., the stimulated hand) by presenting tactile stimuli unpredictably on the middle fingers when one hand was visible (i.e., either the right or left hand). Participants detected the stimulus and responded as fast as possible using either their left or right foot. Consistent with previous results, a clear CUD (5.8ms) was apparent on the unseen hand. Critically, however, no such effect was found on the hand that was visible (−2.2ms). Thus, when touch is delivered to a seen hand, the usual cost in processing speed of responding with a contralateral effector is eliminated. This result suggests that vision of the body improves the interhemispheric integration of tactile-motor responses.

Measurement of Peripheral Vision Reaction Time Identifies White Matter Disruption in Patients with Mild Traumatic Brain Injury.

This study examined whether peripheral vision reaction time (PVRT) in patients with mild traumatic brain injury (mTBI) correlated with white matter abnormalities in centroaxial structures and impairments in neuropsychological testing. Within 24 h after mTBI, crossed reaction times (CRT), uncrossed reaction times (URT), and crossed-uncrossed difference (CUD) were measured in 23 patients using a laptop computer that displayed visual stimuli predominantly to either the left or the right visual field of the retina. The CUD is a surrogate marker of the interhemispheric transfer time (ITT). Within 7 days after the injury, patients received a diffusion tensor-MRI (DTI) scan and a battery of neuropsychological tests. Nine uninjured control subjects received similar testing. Patients 18-50 years of age were included if they had a post-resuscitation Glasgow Coma Scale >13 and an injury mechanism compatible with mTBI. Healthy controls were either age- and gender-matched family members of the TBI patients or healthy volunteers. CUD deficits >2 standard deviations (SD) were seen in 40.9% of patients. The CUD of injured patients correlated with mean diffusivity (MD) (p < 0.001, ρ = -0.811) in the posterior corpus callosum. Patients could be stratified on the basis of CUD on the Stroop 1, Controlled Oral Word Association Test (COWAT), and the obsessive-compulsive component of the Basic Symptom Inventory tests. These studies suggest that the PVRT indirectly measures white matter integrity in the posterior corpus callosum, a brain region frequently damaged by mTBI.

Inter-hemispheric integration of tactile-motor responses across body parts.

In simple detection tasks, reaction times (RTs) are faster when stimuli are presented to the visual field or side of the body ipsilateral to the body part used to respond. This advantage, the crossed-uncrossed difference (CUD), is thought to reflect inter-hemispheric interactions needed for sensorimotor information to be integrated between the two cerebral hemispheres. However, it is unknown whether the tactile CUD is invariant when different body parts are stimulated. The most likely structure mediating such processing is thought to be the corpus callosum (CC). Neurophysiological studies have shown that there are denser callosal connections between regions that represent proximal parts of the body near the body midline and more sparse connections for regions representing distal extremities. Therefore, if the information transfer between the two hemispheres is affected by the density of callosal connections, stimuli presented on more distal regions of the body should produce a greater CUD compared to stimuli presented on more proximal regions. This is because interhemispheric transfer of information from regions with sparse callosal connections will be less efficient, and hence slower. Here, we investigated whether the CUD is modulated as a function of the different body parts stimulated by presenting tactile stimuli unpredictably on body parts at different distances from the body midline (i.e., Middle Finger, Forearm, or Forehead of each side of the body). Participants detected the stimulus and responded as fast as possible using either their left or right foot. Results showed that the magnitude of the CUD was larger on the finger (~2.6 ms) and forearm (~1.8 ms) than on the forehead (≃0.9 ms). This result suggests that the interhemispheric transfer of tactile stimuli varies as a function of the strength of callosal connections of the body parts.

Auditory and Visual Interhemispheric Communication in Musicians and Non-Musicians

The corpus callosum (CC) is a brain structure composed of axon fibres linking the right and left hemispheres. Musical training is associated with larger midsagittal cross-sectional area of the CC, suggesting that interhemispheric communication may be faster in musicians. Here we compared interhemispheric transmission times (ITTs) for musicians and non-musicians. ITT was measured by comparing simple reaction times to stimuli presented to the same hemisphere that controlled a button-press response (uncrossed reaction time), or to the contralateral hemisphere (crossed reaction time). Both visual and auditory stimuli were tested. We predicted that the crossed-uncrossed difference (CUD) for musicians would be smaller than for non-musicians as a result of faster interhemispheric transfer times. We did not expect a difference in CUDs between the visual and auditory modalities for either musicians or non-musicians, as previous work indicates that interhemispheric transfer may happen through the genu of the CC, which contains motor fibres rather than sensory fibres. There were no significant differences in CUDs between musicians and non-musicians. However, auditory CUDs were significantly smaller than visual CUDs. Although this auditory-visual difference was larger in musicians than non-musicians, the interaction between modality and musical training was not significant. Therefore, although musical training does not significantly affect ITT, the crossing of auditory information between hemispheres appears to be faster than visual information, perhaps because subcortical pathways play a greater role for auditory interhemispheric transfer.

Interhemispheric vs. stimulus-response spatial compatibility effects in bimanual reaction times to lateralized visual stimuli

In the present study, we tested right- and left-handed participants in a Poffenberger paradigm with bimanual responses and hands either in an anatomical or in a left-right inverted posture. We observed a significant positive crossed-uncrossed difference (CUD) in RTs for both manual dominance groups and both response postures. These results rule out an explanation of the CUD in terms of stimulus-response spatial compatibility (SRSC) and provide convincing evidence on the important role of interhemispheric callosal transfer in bimanual responding in right- as well as left-handed individuals.

Age-related differences in interhemispheric visuomotor integration measured by the redundant target effect.

Presentation of bilateral redundant visual stimuli produces faster reaction times (RT) than presentation of a single unilateral stimulus; an effect known as the redundant target effect (RTE; Miller, 1982), and is a means of testing interhemispheric visuomotor integration (Ouimet, 2009). RTEs that exceed expectations, based on Miller's race model of inequality (RMI), are referred to as "enhanced RTEs" and imply neural coactivation. Paradoxically, enhanced RTEs are observed in cases of corpus callosum disruption. The Hemispheric Coactivation Hypothesis accounts for this paradox by positing that bihemispheric processing occurs to both unilateral and bilateral stimuli in the normal brain, but occurs only with bilateral stimuli in the disconnected brain. Neuroimaging has revealed decreases in the microstructural integrity of the corpus callosum with age (Ota et al., 2006), but research investigating the bilateral RTE in healthy older individuals is lacking. The present study investigated the bilateral RTE in healthy younger and healthy older adults using simple RT and choice RT tasks. Our prediction that older individuals would show significantly larger RTEs than younger individuals was found to be true for both tasks. Tests of the RMI produced little evidence for coactivation. The crossed-uncrossed difference, generally used as a means of testing visuomotor interhemispheric transfer, was also investigated, but no age effects were found. The observation of greater RTE in age is congruent with the Hemispheric Coactivation hypothesis (Miller, 2004) in which callosal disconnection is associated with increased RTE.

Bimanual crossed–uncrossed difference and asynchrony of normal, anterior- and totally-split-brain individuals

This study investigated interhemispheric transfer in ten normal, four anterior-, and four totally-split-brain individuals, through measures of manual asynchrony and bimanually-recorded crossed–uncrossed difference (CUD). The CUD relied on the difference between crossed and uncrossed responses whereas the asynchrony measure relied on the reaction time difference between the two responding hands. Manipulations of sensory and attentional factors were assessed for both measures. We found a normal CUD (3.8ms) along with an exacerbated and more variable asynchrony for partial split-brain individuals (40.8ms) compared to normal individuals (CUD: 0.4ms, asynchrony: 13.8ms). In turn, the CUD of total split-brain individuals (20.4ms) was larger than that of partial split-brain and normal individuals. Also, the asynchrony of total split-brain individuals (57.6ms) was larger and more variable than that of normal individuals, and more variable than that of partial split-brain individuals. We interpret these results as behavioural evidence of independent mechanisms underlying the CUD and bimanual synchronization, as well as evidence of the joint involvement of both the anterior and the posterior portions of the corpus callosum in bimanual coordination.

Handedness effects on interhemispheric transfer time: A TMS study

The crossed–uncrossed difference (CUD) estimates the interhemispheric transfer time (ITT) through the corpus callosum. Previous research has shown that transcranial magnetic stimulation (TMS) to the occipital cortex determines an increased CUD during cognitive tasks. The aim of the present study was to investigate whether TMS stimulation applied at a motor stage can interfere with the ITT, comparing the performance of left- and right-handed people. Results showed a significant TMS effect, i.e. increasing reaction times were reported when stimulation was delivered on the left primary motor area. Effects were more evident when information was primarily perceived through the dominant hemisphere. Both left and right stimulations increased CUD times in right-handed subjects; however, left-handed subjects showed significant effects associated with left stimulation only. Furthermore, in both groups, TMS produced larger effects in the crossed than in the uncrossed condition. TMS stimulation increased reaction times, thus supporting the idea that the interhemispheric transfer of visuo-motor information occurs at a motor processing stage. The dominant hemisphere seems to play a major role within this process: our data indicates that left- and right-handed people have different ITT latencies associated with the transfer of information to the contralateral hemisphere.

Crossed-uncrossed difference (CUD) in a new light: anatomy of the negative CUD in Poffenberger's paradigm

Crossed Uncrossed Differentials (CUDs) have long been used as surrogate for the interhemispheric transfer time (IHTT). Evidence is presented that macular vision is the province of the major hemisphere, wherein all commands are initiated regardless of the laterality of the effectors of such commands. Using clinical and time-resolved data it is shown also that the above arrangement (i.e. neural handedness) corresponds to the subject's behavioral avowed (avowed, self-declared) handedness only in a statistical sense; with a substantial minority of humanity displaying a disparity of neural and behavioral handedness. Evidence is provided that the negative CUD in previously reported studies was a reflection of such incongruity in those subjects studied. Thus, to lateralize the command center it is sufficient to determine the reaction time of two symmetrically located effectors on the body. The side with longer reaction time is ipsilateral to the major hemisphere, with the difference of the two sides commensurate to transcallosal IHTT.

Corpus Callosal Microstructural Integrity Influences Interhemispheric Processing: A Diffusion Tensor Imaging Study

Normal aging and chronic alcoholism result in disruption of brain white matter microstructure that does not typically cause complete lesions but may underlie degradation of functions requiring interhemispheric information transfer. We examined whether the microstructural integrity of the corpus callosum assessed with diffusion tensor imaging (DTI) would relate to interhemispheric processing speed. DTI yields estimates of fractional anisotropy (FA), a measure of orientation and intravoxel coherence of water diffusion usually in white matter fibers, and diffusivity (<D>), a measure of the amount of intracellular and extracellular fluid diffusion. We tested the hypothesis that FA and <D> would be correlated with (i) the crossed-uncrossed difference (CUD), testing visuomotor interhemispheric transfer; and (ii) the redundant targets effect (RTE), testing parallel processing of visual information presented to each cerebral hemisphere. FA was lower and <D> higher in alcoholics than in controls. In controls but not alcoholics, large CUDs correlated with low FA and high <D> in total corpus callosum and regionally in the genu and splenium. In alcoholics but not controls, small RTEs, elicited with equiluminant stimuli, correlated with low FA in genu and splenium and high <D> in the callosal body. The results provide in vivo evidence for disruption of corpus callosum microstructure in normal aging and alcoholism that has functional ramifications for efficiency in interhemispheric processing.

Magnetic stimulation and the crossed?uncrossed difference (CUD) paradigm: selective effects in the ipsilateral and contralateral hemispheres

When a visual target is presented to one hemifield, manual responses made to the target using the ipsilateral hand (uncrossed responses) are faster than responses using the contralateral hand (crossed response), because there is no need for visuomotor information to be transferred between the hemispheres. This difference in response times is termed the crossed-uncrossed difference (CUD) and is a valuable means of estimating interhemispheric transfer time. We aimed to investigate the CUD by applying repetitive transcranial magnetic stimulation (rTMS) over the left and right occipital cortex during a lateralized target-detection task. Eleven neurologically healthy subjects, all right-handed, participated in the study. Relative to sham TMS we increased the CUD, by inhibiting the crossed latencies, but only when rTMS was applied to the hemisphere receiving visual information. These results replicate and extend previous findings and suggest the inhibitory rTMS effect under the crossed condition might be because the weak visual output is unable to activate the crossed pathway.

Temporary and permanent signs of interhemispheric disconnection after traumatic brain injury

The corpus callosum is frequently damaged by closed head traumas, and the resulting deficits of interhemispheric communication may vary according to the specific position of the lesion within the corpus callosum. This paper describes a single case who suffered a severe traumatic brain injury resulting in a lesion of the posterior body of the corpus callosum. Among the classical symptoms of interhemispheric disconnection, left hand anomia, left upper limb ideomotor dyspraxia, left visual field dyslexia and dysnomia, and left ear suppression in a dichotic listening task were observed shortly after the injury but recovered completely or almost completely with the passage of time. The only symptom of interhemispheric disconnection which was found to persist more than 4 years after the injury was an abnormal prolongation of the crossed-uncrossed difference in a simple visuomotor reaction time task. This prolongation was comparable with that observed in subjects with complete callosal lesions or agenesis. The results suggest that the posterior body of the corpus callosum may be an obligatory interhemispheric communication channel for mediating fast visuo-motor responses. The transient nature of other symptoms of interhemispheric disconnection suggests a relatively wide dispersion of fibers with different functions through the callosal body, such that parts of them can survive a restricted lesion and allow functional recovery of hemispheric interactions. An assessment of the evolution in time of symptoms of interhemispheric disconnection following restricted callosal lesions may reveal fine and coarse features of the anatomo-functional topography of the corpus callosum.

Interhemispheric transmission of visuomotor information in humans: fMRI evidence.

Normal human subjects underwent functional magnetic resonance imaging (fMRI) while performing a simple visual manual reaction-time (RT) task with lateralized brief stimuli, the so-called Poffenberger's paradigm. This paradigm was employed to measure interhemispheric transmission (IT) time by subtracting mean RT for the uncrossed hemifield-hand conditions, that is, those conditions not requiring an IT, from the crossed hemifield-hand conditions, that is, those conditions requiring an IT to relay visual information from the hemisphere of entry to the hemisphere subserving the response. The obtained difference is widely believed to reflect callosal conduction time, but so far there is no direct physiological evidence in humans. The aim of our experiment was twofold: first, to test the hypothesis that IT of visuomotor information requires the corpus callosum and to identify the cortical areas specifically activated during IT. Second, we sought to discover whether IT occurs mainly at premotor or perceptual stages of information processing. We found significant activations in a number of frontal, parietal, and temporal cortical areas and in the genu of the corpus callosum. These activations were present only in the crossed conditions and therefore were specifically related to IT. No selective activation was present in the uncrossed conditions. The location of the activated callosal and cortical areas suggests that IT occurs mainly, but not exclusively, at premotor level. These results provide clear cut evidence in favor of the hypothesis that the crossed-uncrossed difference in the Poffenberger paradigm depends on IT rather than on a differential hemispheric activation.

Changes in Interhemispheric Transfer Rate and the Development of Bimanual Coordination during Childhood

In this study, we investigated the effect of the development of interhemispheric communication on age-related change in bimanual coordination. Interhemispheric communication was assessed by comparing the latency of a manual response to a visual stimulus when the hemisphere perceiving the stimulus and the hemisphere controlling the manual response were the same (uncrossed condition) to the latency when they were different (crossed condition). In the first experiment (5- to 10-year-old children) we used a two-choice response-time task, and in the second experiment (3- to 7-year-old children) we used a simple response-time task. In both studies, bimanual coordination was tested on a line-drawing task, and we compared performance on mirror and parallel movements. The crossed-uncrossed difference decreased with age in both experiments. When estimated on the simple response-time task, the crossed-uncrossed difference was related to the difference in performance between mirror and parallel movements on the bimanual task. Thus, improved interhemispheric communication contributes to progress in bimanual coordination, especially that which requires resisting the attraction of mirror movements in order to rotate both hands with parallel movements.

The Stability of Compromised Interhemispheric Processing in Callosal Dysgenesis and Partial Commissurotomy

The persistence and stability of selective deficits in interhemispheric processing resulting from known callosal pathology have been monitored over periods ranging from ten to thirty five years. The present study included five patients: two with complete agenesis of the corpus callosum, one with partial dysgenesis, and two with a partial section of the corpus callosum. A crossed-uncrossed difference task and four bilateral visual matching tasks were administered to these patients and to groups of normal individuals matched on age and intelligence. As expected, all of the patients showed deficits in speed or accuracy relative to the performance of their control groups. The profile of performance for each patient across the five tasks demonstrated a systematic (but not perfectly consistent) relationship with the location and extent of callosal pathology.