Bilateral Stimuli Research Articles

Bilateral redundancy gain and callosal integrity in a man with callosal lipoma: A diffusion-tensor imaging study

We investigated whether abnormalities in the structural organization of the corpus callosum in the presence of curvilinear lipoma are associated with increased facilitation of response time to bilateral stimuli, an effect known as the redundancy gain (RG). A patient (A.J.) with a curvilinear lipoma of the corpus callosum, his genetically-identical twin, and age-matched control participants made speeded responses to luminant stimuli. Structural organization of callosal regions was assessed with diffusion-tensor imaging. A.J. was found to have reduced structural integrity in the splenium of the corpus callosum and produced a large RG suggestive of neural summation.

Are objects the same as groups? ERP correlates of spatial attentional guidance by irrelevant feature similarity

It has been proposed that the most fundamental units of attentional selection are “objects” that are grouped according to Gestalt factors such as similarity or connectedness. Previous studies using event-related potentials (ERPs) have shown that object-based attention is associated with modulations of the visual-evoked N1 component, which reflects an early cortical mechanism that is shared with spatial attention. However, these studies only examined the case of perceptually continuous objects. The present study examined the case of separate objects that are grouped according to feature similarity (color, shape) by indexing lateralized potentials at posterior sites in a sustained-attention task that involved bilateral stimulus arrays. A behavioral object effect was found only for task-relevant shape similarity. Electrophysiological results indicated that attention was guided to the task-irrelevant side of the visual field due to achromatic-color similarity in N1 (155–205 ms post-stimulus) and early N2 (210–260 ms) and due to shape similarity in early N2 and late N2 (280–400 ms) latency ranges. These results are discussed in terms of selection mechanisms and object/group representations.

Auditory Rightward Spatial Bias Varies as a Function of Age

Age-related changes in auditory spatial perception of linguistic and non-linguistic stimuli in participants between 5 and 79 years of age were studied. The results show that the strength of the rightward perceptual bias in linguistic bilateral (dichotic) stimulus condition changes as a function of age. In childhood and old age also other rightward spatial biases were observed in linguistic as well as in non-linguistic stimulus conditions. We propose that the auditory rightward spatial biases are not specific to the language and are probably modulated by the early development and late decline of the executive functions.

Perceptual congruency of audio-visual speech affects ventriloquism with bilateral visual stimuli

Many studies on multisensory processes have focused on performance in simplified experimental situations, with a single stimulus in each sensory modality. However, these results cannot necessarily be applied to explain our perceptual behavior in natural scenes where various signals exist within one sensory modality. We investigated the role of audio-visual syllable congruency on participants' auditory localization bias or the ventriloquism effect using spoken utterances and two videos of a talking face. Salience of facial movements was also manipulated. Results indicated that more salient visual utterances attracted participants' auditory localization. Congruent pairing of audio-visual utterances elicited greater localization bias than incongruent pairing, while previous studies have reported little dependency on the reality of stimuli in ventriloquism. Moreover, audio-visual illusory congruency, owing to the McGurk effect, caused substantial visual interference on auditory localization. Multisensory performance appears more flexible and adaptive in this complex environment than in previous studies.

Spatial attention influences trial-by-trial relationships between response time and functional connectivity in the visual cortex

Variations of response time (RT) in selective attention tasks are often associated with variations of activity and functional connectivity in sensory cortices that process relevant stimuli. Here, we investigated whether such relationships are influenced by spatial attention. To investigate this hypothesis, we asked fourteen healthy adults to perform a covert spatial attention task, which made use of bilateral stimulus displays, while we recorded their brain activity using functional magnetic resonance imaging (fMRI). As expected, activity in the middle occipital gyrus increased when spatial attention was directed to the contralateral (versus the ipsilateral) visual field. Surprisingly, variations of RT were not associated with variations in the magnitude of this attentional enhancement. As predicted, however, they were linked to opposing variations of functional connectivity between middle occipital regions contralateral (but not ispilateral) to the attended visual field and the left fusiform gyrus, which is thought to figure prominently in the perceptual processing of visually presented letters. These findings suggest that trial-by-trial variations of RT reflect, at least partially, trial-by-trial variations in the extent to which spatial attention enhances functional connectivity between sensory regions that process relevant stimuli.

Lateralization of Interimplant Timing and Level Differences in Children Who Use Bilateral Cochlear Implants

Interaural level differences (ILD) and interaural timing differences (ITD) are important cues for locating sounds in space. Adult bilateral cochlear implant (CI) users use ILDs more effectively than ITDs. Few studies investigated the ability of children who use bilateral CIs to make use of these binaural cues. Our working hypothesis was that children using bilateral CIs are able to perceive changes in ITDs and ILDs similar to their normal-hearing (NH) peers. Participants were two groups of children; 19 bilateral implant users (CI) and nine NH children. The children in the CI group had received a second CI after 4.9 +/- 2.8 yrs of unilateral use. Children performed a four alternative forced-choice lateralization task in which they were asked to describe stimuli as coming from the left side, right side, middle of the head, or from both sides simultaneously. Stimuli were 500 msec trains of electrical pulses delivered to apical electrode no. 18 (CI group) or clicks (NH group) presented 11 times per second with either ITDs (0, 400, 1000, or 2000 microsec delay between sides) or level differences (0, 10, or 20 Current Units (CI group) or 0, 10, or 20 dB (NH group) difference between sides). ITDs were presented using current levels that were balanced using left and right electrically evoked brain stem responses. Stimulus levels evoking response amplitudes that were most similar were used. Responses from children in the CI group changed significantly with changes in ILD of bilateral stimuli, but not with changes in ITD. Responses from children in the CI group were significantly different from those in the NH group in three ways. Children in the CI group perceived bilaterally presented electrical pulses: (1) to come from the second implanted side more often than the first, (2) to rarely come from the middle, and (3) to come from both sides of the head simultaneously. Perceived changes in lateralization with ILD changes were correlated with differences in amplitudes of electrically evoked brain stem responses by the left versus right CI. The results of this study illustrate that children who use bilateral CIs can lateralize stimuli on the basis of level cues, but have difficulty interpreting interimplant timing differences. Perceived lateralization of bilaterally presented stimuli to the second implanted side in many of the stimulus conditions may relate to the use of different device generations between sides. Further differences from normal lateralization responses could be due to abnormal binaural processing, possibly resulting from a period of unilateral hearing before the provision of a second implant or due to insufficiently matched interimplant stimuli. It may be possible to use objective measures such as electrically evoked auditory brain stem responses wave eV amplitudes to provide balanced levels of bilateral stimulation in children who have had no binaural hearing experience.

Automatic attention lateral asymmetry in visual discrimination tasks

There is evidence that automatic visual attention favors the right side. This study investigated whether this lateral asymmetry interacts with the right hemisphere dominance for visual location processing and left hemisphere dominance for visual shape processing. Volunteers were tested in a location discrimination task and a shape discrimination task. The target stimuli (S2) could occur in the left or right hemifield. They were preceded by an ipsilateral, contralateral or bilateral prime stimulus (S1). The attentional effect produced by the right S1 was larger than that produced by the left S1. This lateral asymmetry was similar between the two tasks suggesting that the hemispheric asymmetries of visual mechanisms do not contribute to it. The finding that it was basically due to a longer reaction time to the left S2 than to the right S2 for the contralateral S1 condition suggests that the inhibitory component of attention is laterally asymmetric.

Between-object action coupling influences visual selection: Neuropsychological evidence

Visual extinction, associated with unilateral parietal damage, occurs when a patient can report a single unilateral stimulus but fails to report the same stimulus under bilateral presentation conditions. Prior studies have shwn that extinction can be reduced if bilateral stimuli group to form parts of a single object. In this study we demonstrate that extinction can also be reduced in parietal patients when separate objects fall in appropriate spatial relations for action. We failed to find similar reductions in extinction when stimuli were associatively but not action-related, and we failed to find effects of action relations when words rather than objects were presented to patients. The evidence suggests that action coupling can be used to ‘glue’ objects together for visual selection.

Attention-spreading Based on Hierarchical Spatial Representations for Connected Objects

Attention selects objects or groups as the most fundamental unit, and this may be achieved through a process in which attention automatically spreads throughout their entire region. Previously, we found that a lateralized potential relative to an attended hemifield at occipito-temporal electrode sites reflects attention-spreading in response to connected bilateral stimuli [Kasai, T., & Kondo, M. Electrophysiological correlates of attention-spreading in visual grouping. NeuroReport, 18, 93-98, 2007]. The present study examined the nature of object representations by manipulating the extent of grouping through connectedness, while controlling the symmetrical structure of bilateral stimuli. The electrophysiological results of two experiments consistently indicated that attention was guided twice in association with perceptual grouping in the early phase (N1, 150-200 msec poststimulus) and with the unity of an object in the later phase (N2pc, 310/330-390 msec). This suggests that there are two processes in object-based spatial selection, and these are discussed with regard to their cognitive mechanisms and object representations.

The fluid dynamics of canine olfaction: unique nasal airflow patterns as an explanation of macrosmia

The canine nasal cavity contains hundreds of millions of sensory neurons, located in the olfactory epithelium that lines convoluted nasal turbinates recessed in the rear of the nose. Traditional explanations for canine olfactory acuity, which include large sensory organ size and receptor gene repertoire, overlook the fluid dynamics of odorant transport during sniffing. But odorant transport to the sensory part of the nose is the first critical step in olfaction. Here we report new experimental data on canine sniffing and demonstrate allometric scaling of sniff frequency, inspiratory airflow rate and tidal volume with body mass. Next, a computational fluid dynamics simulation of airflow in an anatomically accurate three-dimensional model of the canine nasal cavity, reconstructed from high-resolution magnetic resonance imaging scans, reveals that, during sniffing, spatially separate odour samples are acquired by each nostril that may be used for bilateral stimulus intensity comparison and odour source localization. Inside the nose, the computation shows that a unique nasal airflow pattern develops during sniffing, which is optimized for odorant transport to the olfactory part of the nose. These results contrast sharply with nasal airflow in the human. We propose that mammalian olfactory function and acuity may largely depend on odorant transport by nasal airflow patterns resulting from either the presence of a highly developed olfactory recess (in macrosmats such as the canine) or the lack of one (in microsmats including humans).

Loss of Somatosensory-evoked Potentials and the Timing of Perception

To determine if patients with brain lesions who have a unilateral loss of their primary somatosensory-evoked potential (SSEP) have altered temporal perception. Benjamin Libet postulated that the neural processing of stimuli to reach the conscious awareness takes 300 to 500 milliseconds and that accurate temporal perception of stimulus onset requires a retroactive computation. Although Libet proposed that the primary SSEP acts as a timing marker for this backward referral of perceived stimulus onset time, there has not been a systematic study of the necessity of the primary SSEP for perceptual timing. Participants were 10 healthy older adults and 10 stroke patients with hemisensory deficits. SSEPs were recorded from each hemisphere using median nerve stimulation. The participants' temporal perception of sensory stimuli was determined by asking them the temporal order of bilateral hand stimuli over varying interstimulus intervals. Patients with unilateral loss of SSEPs had a significantly greater mean delay in perception of stimuli from their contralesional arm than participants with intact bilateral SSEPs [mean delay (+/-standard deviation): 134 (+/-142) msec vs. 2.5 (+/-13) msec; P=0.03]. These results demonstrate that loss of SSEP is associated with a delay in perceptual awareness. This observation is consistent with the hypotheses that the SSEP acts as a marker for cortical events important for perceptual timing.

Attentional and sensory effects of lowered levels of intrinsic alertness

Low levels of intrinsic alertness are associated with lateralized performance in visual tasks, similar to neglect of the left (ipsilesional) visual hemi-field. However, it is unclear whether reduced alertness produces a specific lateralization of spatial-attentional processes in terms of the prioritization of right- over left-side stimuli, or whether it affects more basic functions of visuo-sensory coding, and/or higher function of the top-down control of selection, of stimuli on the left side. To decide between these alternatives, the present study examined the effects of lowered alertness, induced by a 50-min vigilance task, in a partial-report paradigm of briefly presented letter displays. With only one (unilateral) stimulus in display, no specific hemi-field effects were found under low-alertness conditions, indicating that reduced alertness impairs neither sensory effectiveness nor the top-town control of selection. However, with dual, bilateral stimuli, report accuracy was specifically affected for left-side targets (in subjects who showed comparable performance for both sides under normal-alertness conditions). This pattern can be interpreted in terms of a specific bias in spatial-attentional weighting, where prioritization of stimuli on the right leads to (mild) extinction of targets on the left. Moreover, participants who had a lower general level of alertness also showed a more pronounced re-distribution of weights, evidenced by a more severe imbalance in report accuracy, in a low compared to a normal state of alertness. This suggests that a low general level of intrinsic alertness engenders a specific vulnerability to neglect-like performance with a (mild) left-side extinction.

A choice reaction time index of callosal anatomical homotopy

Tachistoscopically presented bilateral stimulus pairs not parallel to the meridian produced significantly longer RTs on a task requiring discrimination of shapes (Go/no-Go) than pairs emplaced symmetrically on each side of the meridian in Desjardins and Braun [Desjardins, S., & Braun, C. M. J. (2006). Homotopy and heterotopy and the bilateral field advantage in the Dimond paradigm. Acta Psychologica, 121, 125–136]. This was explained by the fact that there are more homotopic than heterotopic fibers in the corpus callosum. However: (1) different parts of the visual field were not equiprobably stimulated, possibly causing subtle biases, (2) the predicted cost of vertical asymmetry was tested only with bilateral stimuli, and (3) interstimulus distance was at the outer limit of callosal midline fusion (10.6°). Here, a tachistoscopic experiment with 24 normal participants replicated the between-field vertical symmetry advantage [Desjardins, S., & Braun, C. M. J. (2006). Homotopy and heterotopy and the bilateral field advantage in the Dimond paradigm. Acta Psychologica, 121, 125–136.], but without irrelevant stimulation conditions and with more proximal stimuli. In addition, a significant specific cost of vertical asymmetry of 7 ms was found for between-field integration over within-field integration. As far as we know, this is the first demonstration of an effect of callosal anatomical homotopy with reaction time.

Parallel interhemispheric processing in hemineglect: Relation to visual field defects

Parallel interhemispheric processing is required to explore our visual environment and to integrate visual information from both hemifields simultaneously. Damage to the right temporo-parietal cortex can disrupt such parallel processes and result in neglect and visual extinction of stimuli in the left contralesional visual space. Neglected or extinguished stimuli can still be processed, yet without reaching the patient's awareness. Such unconscious processing has been attributed to structurally intact primary visual areas in neglect. To study whether unconscious parallel processing depends on visual functional integrity, we compared the performance of neglect patients with visual field defects (VFDs) (n=11) and hemianopic patients with partial or complete blindness of one visual hemifield (n=11) on redundant targets effects (RTE). The RTE manifests as faster reaction times to redundant paired (two stimuli, one in each hemifield) than single stimulation (in one hemifield). We found RTEs, i.e., unconscious processing, in neglect patients but not in hemianopic patients. Furthermore, neglect patients showed large crossed-uncrossed differences (CUDs), i.e., faster response times to ipsi- than contralesional hemifield stimulation, reflecting a difference in processing speed for single stimuli in the two hemispheres that were correlated with VFDs and visual extinction. The finding that extinction, but not RTE, was correlated with the CUD suggests that under competitive bilateral stimulus conditions the delayed contralesional visual field input may not be detected by the intact left hemisphere, which presumably mediates the task given the impairment of the right hemisphere. By contrast, unconscious parallel processing of contralesional stimuli (RTE) occurred even when contralesional visual field input is lacking (VFD) or delayed (CUD) and is possibly mediated via subcortical visual pathways.

Mapping the bilateral visual integration by EEG and fMRI

In the human visual system, the internal representation of the left and right visual hemifields is split at the midline of the two cerebral hemispheres. The present study aims to address the questions of when and where the lateralized cortical visual representations are merged to form an intact percept by using a multimodal neuroimaging approach. Visual evoked potential (VEP) and functional magnetic resonance imaging (fMRI) data were acquired from a group of healthy subjects presented with unilateral versus bilateral visual stimuli. Cortical activities involved in processing bilateral visual information are expected to be equally responsive to ipsilateral and contralateral stimuli, and demonstrate spatial nonlinearity in the response to bilateral stimuli. Utilizing these features, we performed integrative as well as separate analyses for both VEP and fMRI data. The present results suggest that i) the majority of cortical activity that integrates visual information across hemifields takes place at extrastriate areas during late visual processing, and that ii) the lateral occipito-temporal (LOT) regions (likely the MT+ complex) and the medial occipital cortex (i.e. V1) may contribute to bilateral visual integration during early visual processing. Our findings are generally in agreement with the bottom-up visual hierarchy, with the exception of the evidence suggesting an early activation of the higher-tier LOT areas and the influence from ipsilateral visual inputs upon the V1 response.

Optic Flow Stimuli in and Near the Visual Field Centre: A Group fMRI Study of Motion Sensitive Regions

Motion stimuli in one visual hemifield activate human primary visual areas of the contralateral side, but suppress activity of the corresponding ipsilateral regions. While hemifield motion is rare in everyday life, motion in both hemifields occurs regularly whenever we move. Consequently, during motion primary visual regions should simultaneously receive excitatory and inhibitory inputs. A comparison of primary and higher visual cortex activations induced by bilateral and unilateral motion stimuli is missing up to now. Many motion studies focused on the MT+ complex in the parieto-occipito-temporal cortex. In single human subjects MT+ has been subdivided in area MT, which was activated by motion stimuli in the contralateral visual field, and area MST, which responded to motion in both the contra- and ipsilateral field. In this study we investigated the cortical activation when excitatory and inhibitory inputs interfere with each other in primary visual regions and we present for the first time group results of the MT+ subregions, allowing for comparisons with the group results of other motion processing studies. Using functional magnetic resonance imaging (fMRI), we investigated whole brain activations in a large group of healthy humans by applying optic flow stimuli in and near the visual field centre and performed a second level analysis. Primary visual areas were activated exclusively by motion in the contralateral field but to our surprise not by central flow fields. Inhibitory inputs to primary visual regions appear to cancel simultaneously occurring excitatory inputs during central flow field stimulation. Within MT+ we identified two subregions. Putative area MST (pMST) was activated by ipsi- and contralateral stimulation and located in the anterior part of MT+. The second subregion was located in the more posterior part of MT+ (putative area MT, pMT).

Preattentive surface and contour grouping in Kanizsa figures: Evidence from parietal extinction

Visual extinction commonly occurs after unilateral, parietal brain damage and manifests in a failure to identify contralesional stimuli when presented simultaneously with other, ipsilesional stimuli – but full awareness for single stimulus presentations. However, extinction can be substantially reduced when preattentive grouping operations link fragmentary items across hemifields into a coherent object. For instance, one study demonstrated preserved access to bilateral stimulus segments when these could be grouped to form a Kanizsa square [Mattingley, J. B., Davis, G., & Driver, J. (1997). Preattentive filling-in of visual surfaces in parietal extinction. Science, 275, 671–674]. Here, we investigated the relative contributions of distinct object attributes to the spared access in Kanizsa figure completion in extinction, by systematically varying the degree to which bilateral surface filling-in and contour interpolations group disparate items. We demonstrate that surface information can substantially reduce extinction, whereas contour completions showed comparably smaller influences. In summary, such graded influences of object attributes support recurrent models of grouping, first, linking fragmentary parts into coherent surfaces and, second, interpolating the precise boundaries.

Functional MRI of the primary somatosensory cortex in extinction to simultaneous bilateral tactile stimuli due to right temporal lobe stroke

Patients with right posterior temporoparietal cortical lesions often exhibit extinction to tactile double simultaneous stimuli (EDSS). It is not known whether sensory unawareness-extinction results from suppression of sensory input into the somatosensory cortex (SI), inhibition of SI, or interference which prevents SI output from activating and being fully processed by association areas. A patient with EDSS due to a right temporal stroke sparing SI and posterior parietal cortex and eight age-matched healthy controls were studied with fMRI during tactile stimulation. The volume of activation of SI during tactile stimulation of the right hand, the left hand and both hands was measured and the patient's volume of activation was compared to that of the control subjects in each of these stimulus conditions. Although the patient demonstrated behavioral EDSS, during fMRI the patient's activation of SI on both sides was within the range of the control participants' volumes of activation. These findings suggest that EDSS in patients with a right temporal lobe stroke results from processing abnormalities that occur after these afferent tactile stimuli are processed by SI.

Callosal function in MS patients with mild and severe callosal damage as reflected by diffusion tensor imaging

In this study, callosal function was behaviourally tested in MS patients with a redundant stimuli task. Reaction times to uni- and bilateral visual stimuli are recorded. Normal subjects respond faster to bilateral than to unilateral stimuli. This effect is called the redundancy gain effect. In patients with agenesis of the corpus callosum, the redundancy gain exceeds that predicted by probability summation, suggesting a mediating influence of the corpus callosum in healthy controls. The aim of this study is to investigate the effect of callosal damage on the redundancy gain in MS patients by investigating the probability summation model. Seventeen MS patients and as many matched healthy controls performed the redundancy gain task. In order to objectify callosal damage in our MS group, diffusion tensor imaging (DTI) derived measures such as fractional anisotropy (FA) and mean diffusivity (MD) in the corpus callosum were obtained. Callosal FA and MD significantly differed in our MS group compared to the healthy controls, indicating pathological callosal involvement. Since the amount of callosal damage was highly variable within the MS group, the MS cohort was split into a low and a high callosal-injured group as quantified by FA. The high FA group performed like the healthy controls, whereas violations of the probability ( race) model were found for the low FA group. We conclude that behavioural measures obtained by the redundancy gain paradigm reflect callosal pathology in MS as measured by DTI.

Enhanced redundancy gain in schizophrenics: A correlate of callosal dysfunction?

An abnormal pattern of hemispheric asymmetry, possibly as a result of disturbed interhemispheric communication, is widely, albeit by no means unanimously, held as a major cause of schizophrenia. To behaviourally test interhemispheric communication in schizophrenia we used a task that has been shown to be a reliable indicator of callosal functioning, namely, the redundant signals effect (RSE). It consists of the speeding of simple reaction time when responding to double as opposed to single visual stimuli. When the stimuli in a pair are presented to different hemispheres patients who underwent total commissurotomy or suffer from callosal agenesis show a paradoxically enhanced RSE with respect to healthy controls. Therefore, if schizophrenia patients have a callosal abnormality they ought to show a similar effect. In three experiments we tested a total of 55 patients with a diagnosis of schizophrenia and 51 healthy controls. In Experiment 1 we presented unilateral single stimuli and bilateral simultaneous double stimuli. The RSE was reliably larger in schizophrenics than in controls. In Experiment 2 the temporal interval between the two stimuli in a pair was varied. We found that while in controls the RSE disappeared with interstimulus intervals longer than 17 ms, in schizophrenia patients there was a RSE only for simultaneous double stimuli. Finally, in Experiment 3 we found that there was no enhanced redundancy gain in schizophrenics when the double stimuli were presented to one and the same hemisphere, and therefore, with no need for callosal transmission. All in all, the present results provide evidence of a callosal dysfunction in schizophrenia that impairs interhemispheric integration.