Detecting violations of sensory expectancies following cerebellar degeneration: a mismatch negativity study

Two hypotheses concerning cerebellar function and predictive behavior are the sensory prediction hypothesis and the timing hypothesis. The former postulates that the cerebellum is critical in generating expectancies regarding forthcoming sensory information. The latter postulates that this structure is critical in generating expectancies that are precisely timed; for example, the expected duration of an event or the time between events. As such, the timing hypothesis constitutes a more specific form of prediction. The present experiment contrasted these two hypotheses by examining the mismatch negativity (MMN) response in patients with cerebellar cortical atrophy and matched controls. While watching a silent movie, a stream of task-irrelevant sounds was presented. A standard sound was presented 60% of the time, whereas the remaining sounds deviated from the standard on one of four dimensions: duration, intensity, pitch, or location. The timing between stimuli was either periodic or aperiodic. Based on the sensory prediction hypothesis, the MMN for the patients should be abnormal across all four dimensions. In contrast, the timing hypothesis would predict a selective impairment of the duration MMN. Moreover, the timing hypothesis would also predict that the enhancement of the MMN observed in controls when the stimuli are presented periodically should be attenuated in the patients. Compared to controls, the patients exhibited a delayed latency in the MMN to duration deviants and a similar trend for the intensity deviants, while pitch and location MMNs did not differ between groups. Periodicity had limited and somewhat inconsistent effects. The present results are at odds with a general role for the cerebellum in sensory prediction and provide partial support for the timing hypothesis.     

Automatic detection of motion direction changes in the human brain

The possibility that the visual system is able to register unattended changes is still debated in the literature. However, it is difficult to understand how a sensory system becomes aware of unexpected salient changes in the environment if attention is required for detecting them. The ability to automatically detect unusual changes in the sensory environment is an adaptive function which has been confirmed in other sensory modalities (i.e. audition). This deviance detector mechanism has proven to be based on a preattentive nonrefractory memory-comparison process. To investigate whether such automatic change detection mechanism exists in the human visual system, we recorded event-related potentials to sudden changes in a biologically important feature, motion direction. Unattended sinusoidal gratings varying in motion direction in the peripheral field were presented while subjects performed a central task with two levels of difficulty. We found a larger negative displacement in the electrophysiological response elicited by less frequent stimuli (deviant) at posterior scalp locations. Within the latency range of the visual evoked component N2, this differential response was elicited independently of the direction of motion and processing load. Moreover, the results showed that the negativity elicited by deviants was not related to a differential refractory state between the electrophysiological responses to frequent and infrequent directions of motion, and that it was restricted to scalp locations related to motion processing areas. The present results suggest that a change-detection mechanism sensitive to unattended changes in motion direction may exist in the human visual system.     

Retinotopic organization of early visual spatial attention effects as revealed by PET and ERPs

Cerebral blood flow PET scans and high-density event-related potentials (ERPs) were recorded (separate sessions) while subjects viewed rapidly-presented, lower-visual-field, bilateral stimuli. Active attention to a designated side of the stimuli (relative to passive-viewing conditions) resulted in an enhanced ERP positivity (P1 effect) from 80–150 msec over occipital scalp areas contralateral to the direction of attention. In PET scans, active attention vs. passive showed strong activation in the contralateral dorsal occipital cortex, thus following the retinotopic organization of the early extrastriate visual sensory areas, with some weaker activation in the contralateral fusiform. Dipole modeling seeded by the dorsal occipital PET foci yielded an excellent fit for the P1 attention effect. In contrast, dipoles constrained to the fusiform foci fit the P1 effect poorly, and, when the location constraints were released, moved upward to the dorsal occipital locations during iterative dipole fitting. These results argue that the early ERP P1 attention effects for lower-visual-field stimuli arise mainly from these dorsal occipital areas and thus also follow the retinotopic organization of the visual sensory input pathways. These combined PET/ERP data therefore provide strong evidence that sustained visual spatial attention results in a preset, top-down biasing of the early sensory input channels in a retinotopically organized way. Hum. Brain Mapping 5:280–286, 1997. © 1997 Wiley-Liss, Inc.     

A shift of visual spatial attention is selectively associated with human EEG alpha activity

Event-related potentials and ongoing oscillatory electroencephalogram (EEG) activity were measured while subjects performed a cued visual spatial attention task. They were instructed to shift their attention to either the left or right visual hemifield according to a cue, which could be valid or invalid. Thereafter, a peripheral target had to be evaluated. At posterior parietal brain areas early components of the event-related potential (P1 and N1) were higher when the cue had been valid compared with invalid. An anticipatory attention effect was found in EEG alpha magnitude at parieto-occipital electrode sites. Starting 200 ms before target onset alpha amplitudes were significantly stronger suppressed at sites contralateral to the attended visual hemifield than ipsilateral to it. In addition, phase coupling between prefrontal and posterior parietal electrode sites was calculated. It was found that prefrontal cortex shows stronger phase coupling with posterior sites that are contralateral to the attended hemifield than ipsilateral sites. The results suggest that a shift of attention selectively modulates excitability of the contralateral posterior parietal cortex and that this posterior modulation of alpha activity is controlled by prefrontal regions.     

Hemifield effects of spatial attention in early human visual cortex

Early visual areas (V1, V2, V3/VP, V4v) contain representations of the contralateral hemifield within each hemisphere. Little is known about the role of the visual hemifields along the visuo-spatial attention processing hierarchy. It is hypothesized that attentional information processing is more efficient across the hemifields (known as bilateral field advantage) and that the integration of information is greater within one hemifield as compared with across the hemifields. Using functional magnetic resonance imaging we examined the effect of distance and hemifield on parallel attentional processing in the early visual areas (V1-V4v) at individually mapped retinotopic locations aligned adjacently or separately within or across the hemifields. We found that the bilateral field advantage in parallel attentional processing over separated attended locations can be assigned, at least partly, to differences in distractor position integration in early visual areas. These results provide evidence for a greater integration of locations between two attended locations within one hemifield than across both hemifields. This nicely correlates with behavioral findings of a bilateral field advantage in parallel attentional processing (when distractors in between cannot be excluded) and a unilateral field advantage if attention has to be shifted across separated locations (when locations in between were integrated).     

Executive control of spatial attention shifts in the auditory compared to the visual modality

Voluntary orienting of visual spatial attention has been shown to be associated with activation in a distributed network of frontal and parietal brain areas. Neuropsychological data suggest that at least some of these areas should be sensitive to the direction in which attention is shifted. The aim of this study was to use rapid event-related functional magnetic resonance imaging to investigate whether spatial attention in the auditory modality is subserved by the same or different brain areas as in the visual modality, and whether the auditory and visual attention networks show any degree of hemispheric lateralisation or sensitivity to the direction of attention shifts. The results suggest that auditory and visual spatial attention shifts are controlled by a supramodal network of frontal, parietal and temporal areas. Areas activated included the precuneus and superior parietal cortex, the inferior parietal cortex and temporo-parietal junction, as well as the premotor and supplementary motor areas and dorsolateral prefrontal cortex (DLPFC). In the auditory task, some of these areas, in particular the precuneus as well as the inferior parietal cortex and temporo-parietal junction, showed 'relative' asymmetry, in that they responded more strongly to attention shifts towards the contralateral than the ipsilateral hemispace. Some areas, such as the right superior parietal cortex and left DLPFC, showed 'absolute' asymmetry, in that they responded more strongly in one than in the other hemisphere.     

Timing the fearful brain: unspecific hypervigilance and spatial attention in early visual perception

Numerous studies suggest that anxious individuals are more hypervigilant to threat in their environment than nonanxious individuals. In the present event-related potential (ERP) study, we sought to investigate the extent to which afferent cortical processes, as indexed by the earliest visual component C1, are biased in observers high in fear of specific objects. In a visual search paradigm, ERPs were measured while spider-fearful participants and controls searched for discrepant objects (e.g. spiders, butterflies, flowers) in visual arrays. Results showed enhanced C1 amplitudes in response to spatially directed target stimuli in spider-fearful participants only. Furthermore, enhanced C1 amplitudes were observed in response to all discrepant targets and distractors in spider-fearful compared with non-anxious participants, irrespective of fearful and non-fearful target contents. This pattern of results is in line with theoretical notions of heightened sensory sensitivity (hypervigilance) to external stimuli in high-fearful individuals. Specifically, the findings suggest that fear facilitates afferent cortical processing in the human visual cortex in a non-specific and temporally sustained fashion, when observers search for potential threat cues.     

Multisensory integration of speech sounds with letters vs. visual speech: only visual speech induces the mismatch negativity

Numerous studies have demonstrated that the vision of lip movements can alter the perception of auditory speech syllables (McGurk effect). While there is ample evidence for integration of text and auditory speech, there are only a few studies on the orthographic equivalent of the McGurk effect. Here, we examined whether written text, like visual speech, can induce an illusory change in the perception of speech sounds on both the behavioural and neural levels. In a sound categorization task, we found that both text and visual speech changed the identity of speech sounds from an /aba/‐/ada/ continuum, but the size of this audiovisual effect was considerably smaller for text than visual speech. To examine at which level in the information processing hierarchy these multisensory interactions occur, we recorded electroencephalography in an audiovisual mismatch negativity (MMN, a component of the event‐related potential reflecting preattentive auditory change detection) paradigm in which deviant text or visual speech was used to induce an illusory change in a sequence of ambiguous sounds halfway between /aba/ and /ada/. We found that only deviant visual speech induced an MMN, but not deviant text, which induced a late P3‐like positive potential. These results demonstrate that text has much weaker effects on sound processing than visual speech does, possibly because text has different biological roots than visual speech.     

Age-related changes in the dynamics of human albino visual pathways

A deficiency of melanin in the retinal pigment epithelium, which regulates the development of neural retina, leads to chiasmal misrouting such that the uncrossed pathway (to the ipsilateral hemisphere) is reduced relative to the crossed pathway (to the contralateral hemisphere). This study examines age-related changes in the flash and pattern appearance visual evoked potentials (VEP) of human albinos. Scalp recorded cortical VEPs to flash (FVEP) and pattern appearance stimulation were recorded in 58 albino (8 months to 60 years) and 34 normal subjects (4-55 years). VEPs were analysed by amplitude and latency. The contralateral hemisphere FVEP amplitude decreased with age in albino subjects, as in both hemispheres in normals. However, the ipsilateral hemisphere FVEP amplitude was significantly lower in young albino subjects, initially giving a marked interhemispheric asymmetry, but this normalized with age. Significant interhemispheric FVEP latency asymmetries were not observed. The contralateral pattern appearance VEP latency in albino subjects decreased with age, as in both hemispheres in normals; the ipsilateral latency increased significantly with age. Significant interhemispheric pattern appearance VEP amplitude asymmetries were not observed. These novel and unexpected observations indicate significant age-related changes in the retinocortical pathways of the human albino. These changes have implications for our understanding of development and plasticity of the central visual pathways.     

Comparative study of visual evoked potentials in spinocerebellar ataxias and multiple sclerosis

Visual evoked potentials (VEPs) were delayed in 11 out of 18 patients with Friedreich's ataxia, in 1 out of 8 patients with Strumpell's hereditary spastic ataxia, in 2 out of 5 cases with cerebellar atrophy and in 42 out of 50 patients with multiple sclerosis (MS). Responses were normal in 5 cases with Pierre Marie's disease. Amplitude and temporal dispersion were statistically analyzed in the above-mentioned groups of patients with respect to controls. An abnormal temporal dispersion, also considered as interpeak N1P2, was frequently found in MS but only occasionally in spinocerebellar ataxias. Amplitude was statistically reduced in Friedreich's ataxia group, where an inverse relationship between latency and amplitude was found. No relation was found between VEP delay and duration of the disease, in any group considered.     

Prism adaptation and spatial attention: a study of visual search in normals and patients with unilateral neglect

Visuomotor adaptation to a prism-induced lateral displacement of the visual field induces mild perceptual biases in healthy individuals and improves symptoms of unilateral neglect. The present study employed a speeded visual search task to test the hypothesis that prism adaptation induces an adaptive redistribution of selective spatial attention. In Experiment 1, 32 neurologically healthy, right-handed participants were adapted to a 15° prism–induced lateral (left or right) displacement of the visual field. Spatial attention was measured by search time and error-rate in unique-feature (“preattentive”) and feature-absent (“serial”) visual search tasks, before and after prism adaptation. The single target appeared at different locations within arrays of 12, 24 or 48 items. Contrary to the attentional hypothesis, the pattern of search performance across the display remained unchanged following prism adaptation. In Experiment 2, we tested four patients with unilateral right hemisphere damage on the visual search tasks, before and after adaptation to 15° rightward-displacing prisms. All four patients showed a pathological gradient of spatial attention toward the ipsilesional side prior to adaptation. Consistent with the results from Experiment 1, the gradient in search performance shown by the patients did not change following prism adaptation. Taken together, these findings suggest that the perceptual aftereffects in normals and amelioration of unilateral neglect following prism adaptation are not mediated by an adaptive redistribution of spatial attention.     

Intracortical evoked potentials of cats elicited by punctate visual stimuli in receptive field peripheries

Intracortical visual evoked potentials, related to stimulus eccentricity from receptive field center, were investigated as a possible measure of interaction among adjacent populations of cortical neurons. Displacement of the stimuli from field center resulted in (1) a progressive decline in amplitude of the primary potential and (2) an enhancement or development of longer latency (60–100 ms) ‘secondary’ potentials often not apparent with field center stimulation.     

Preferential attention to animals and people is independent of the amygdala

The amygdala is thought to play a critical role in detecting salient stimuli. Several studies have taken ecological approaches to investigating such saliency, and argue for domain-specific effects for processing certain natural stimulus categories, in particular faces and animals. Linking this to the amygdala, neurons in the human amygdala have been found to respond strongly to faces and also to animals. However, the amygdala’s necessary role for such category-specific effects at the behavioral level remains untested. Here we tested four rare patients with bilateral amygdala lesions on an established change-detection protocol. Consistent with prior published studies, healthy controls showed reliably faster and more accurate detection of people and animals, as compared with artifacts and plants. So did all four amygdala patients: there were no differences in phenomenal change blindness, in behavioral reaction time to detect changes or in eye-tracking measures. The findings provide decisive evidence against a critical participation of the amygdala in rapid initial processing of attention to animate stimuli, suggesting that the necessary neural substrates for this phenomenon arise either in other subcortical structures (such as the pulvinar) or within the cortex itself.     

Event-related potential activity in the basal ganglia differentiates rewards from nonrewards: temporospatial principal components analysis and source localization of the feedback negativity

Event-related potential studies of reward processing have consistently identified the feedback negativity (FN), an early neural response that differentiates feedback indicating unfavorable versus favorable outcomes. Several important questions remain, however, about the nature of this response. In this study, the FN was recorded in response to monetary gains and losses during a laboratory gambling task, and temporospatial principal components analysis was used to separate the FN from overlapping responses. The FN was identified as a positive deflection at frontocentral recording sites that was enhanced for rewards compared with nonrewards. Furthermore, source localization techniques identified the striatum as a likely neural generator. These data indicate that this apparent FN reflects increased striatal activation in response to favorable outcomes that is reduced or absent for unfavorable outcomes, thereby providing unique information about the timing and nature of basal ganglia activity related to reward processing.     

大鼠诱发电位检测法及正常值探讨

目的 :探讨大鼠诱发电位检测方法及皮层诱发电位正常值。方法 :对 4 0只 Wistar大鼠进行 BAEP及CEP检测。结果 :BAEP的 I至 V波潜伏期 ( X± S)分别为 1 .5 9± 0 .2 ,2 .4 0± 0 .2 6 ,3.1 7± 0 .36 ,4 .2 5± 0 .37及 5 .2 5± 0 .36 ms波峰间期分别为 I- 1 .5 8± 0 .2 8ms、 -V 2 .0 9± 0 .3ms及 I-V3.6 7± 0 .31 ms、CEP主反应呈先正后负的 PN型 ,部分出现后发放 ,主反应潜伏期为 6 .6 3± 1 .33ms。结论 :用这种方法检则大鼠诱发电位 ,方法可靠 ,图形清晰 ,重复性好 ,数据相对较稳定 ,为今后大鼠脑诱发电位异常判断奠定了良好的基础。     

Covariations in ERP and PET measures of spatial selective attention in human extrastriate visual cortex

In a previous study using positron emission tomography (PET), we demonstrated that focused attention to a location in the visual field produced increased regional cerebral blood flow in the fusiform gyrus contralateral to the attended hemifield (Heinze et al. [1994]: Nature 372:543). We related these effects to modulations in the amplitude of the P1 component (80–130 msec latency) of the visual event-related brain potentials (ERPs) recorded from the same subjects, under the identical stimulus and task conditions. Here, we replicate and extend these findings by showing that attention effects in the fusiform gyrus and the P1 component were similarly modulated by the perceptual load of the task. When subjects performed a perceptually demanding symbol-matching task within the focus of spatial attention, the fusiform activity and P1 component of the ERP were of greater magnitude than when the subjects performed a less perceptually demanding task that required only luminance detection at the attended location. In the latter condition, both the PET and ERP attention effects were reduced. In addition, in the present data significant activations were also obtained in the middle occipital gyrus contralateral to the attended hemifield, thereby demonstrating that multiple regions of extrastriate visual cortex are modulated by spatial attention. The findings of covariations between the P1 attention effect and activity in the posterior fusiform gyrus reinforce our hypothesis that common neural sources exist for these complementary, but very different measures of human brain activity. Hum. Brain Mapping 5:273–279, 1997. © 1997 Wiley-Liss, Inc.     

Study of visual evoked potentials in diabetics without retinopathy: correlations with clinical findings and polyneuropathy

To evaluate central optic pathways' involvement in diabetics, visual evoked potentials (VEP), in particular the latency of positive peak (LP100), were studied in 35 patients without retinopathy (4 insulin-dependent, 31 non-insulin-dependent) and 35 normal controls using reversal pattern stimulation. LP100 was significantly delayed in diabetics at both binocular and monocular stimulation. Furthermore, the delay in LP100 was significantly longer in the diabetics with polyneuropathy than in those without, particularly after binocular stimulation. A positive correlation was found between latencies of VEP and HbA1, duration of disease, and neurologic score. VEP measurement seems a simple and sensitive method for detecting early alterations in central optic pathways in diabetics.     

Multifocal visual evoked potentials in optic neuritis and multiple sclerosis: A review

Multifocal visual evoked potential (mf-VEP) represents a new approach to the classical full field (ff-)VEP with separate responses from up to 60 sectors of the visual field. A thorough literature survey of the use of mf-VEP in optic neuritis (ON) and multiple sclerosis (MS) is presented (38 published studies were retrieved). Mf-VEP provides direct topographical information of specific lesions and facilitates investigations on structural-functional correlations thus providing new methods for exploring the interplay between demyelination, atrophy and remyelination in MS. Good correlation was shown between mf-VEP and OCT, ff-VEP, MRI (MTR, DTI), 30-2 standard automated perimetry and low-contrast-visual acuity. All but one study showed superior sensitivity and specificity compared to ff-VEP, especially with regards to small, peripheral lesions or lesions of the upper visual field. Mf-VEP has shown superior sensitivity and specificity than established methods in diagnosing optic nerve lesions and tracking functional recovery following lesions. Abnormal mf-VEP responses in the fellow, non-ON afflicted eye may predict MS risk in ON patients. No standardization currently exists and no direct comparisons in ON and MS between at least 5 different commercially available mf-VEP systems have so far been published. Despite these limitations, mf-VEP is a promising new tool of diagnostic and prognostic value of mf-VEP in ON and MS.     

Spatial attention in area V4 is mediated by circuits in primary visual cortex

The ability to covertly select visual stimuli in our environment based on their behavioral relevance is an important skill. Stimulus selection has been studied experimentally, at the single neuron as well as at the population level, by recording from the visual cortex of subjects performing attention-demanding tasks, but studies at the local circuit level are lacking. We conducted simulations of a primary visual cortex (V1) model to provide insight into the local circuit computation underlying stimulus selection in V4.Two small oriented rectangular bars were placed at different locations in the 4 by 4 degree visual field represented by the V1 model, such that they activated different V1 neurons but such that they were both inside the classical receptive field (CRF) of the same V4 neuron. The biased competition framework [Desimone, R., & Duncan, J. (1995). Neural mechanisms of selective visual attention. Annual Review of Neuroscience, 18, 193–222] makes predictions for the response of V4 neurons and the modulation thereof by spatial and feature attention. In our simulation of the V1 network, we obtained results consistent with these predictions for V4 when the model had long-range excitatory projections targeting inhibitory neurons and when spatial attention was mediated by a spatially restricted projection that either inhibited the inhibitory neurons or excited the excitatory neurons. Although it is not clear whether attention effects measured in V4 neurons are generated mostly by local circuits within V4, our simulations suggest that spatial attention at a resolution less than the size of the CRF of a V4 neuron is inherited from upstream areas like V1 and relies on circuits mediating surround suppression at the single neuron level. Furthermore, the model displayed global oscillations in the alpha frequency range (around 10 Hz), whose coherence was highest in the absence of visual stimulation, which is consistent with electroencephalograms recorded in humans. By contrast, when a stimulus was presented the alpha oscillation sped up and became less coherent, whereas at the single column level (40–480 cells) transient beta/gamma oscillations were observed with a frequency between 25 and 50 Hz.     

Pushing attention to one side: Force field adaptation alters neural correlates of orienting and disengagement of spatial attention

Sensorimotor adaptation to wedge prisms can alter the balance of attention between left and right space in healthy adults, and improve symptoms of spatial neglect after stroke. Here we asked whether the orienting of spatial attention to visual stimuli is affected by a different form of sensorimotor adaptation that involves physical perturbations of arm movement, rather than distortion of visual feedback. Healthy participants performed a cued discrimination task before and after they made reaching movements to a central target. A velocity‐dependent force field pushed the hand aside during each reach, and required participants to apply compensatory forces toward the opposite side. We used event‐related potentials (ERPs) to determine whether electroencephalography (EEG) responses reflecting orienting (cue‐locked N1) and disengagement (target‐locked P1) of spatial attention are affected by adaptation to force fields. After adaptation, the cue‐locked N1 was relatively larger for stimuli presented in the hemispace corresponding to the direction of compensatory hand force. P1 amplitudes evoked by invalidly cued targets presented on the opposite side were reduced. This suggests that force field adaptation boosted attentional orienting responses toward the side of hand forces, and impeded attentional disengagement from that side, mimicking previously reported effects of prism adaptation. Thus, remapping between motor commands and intended movement direction is sufficient to bias ERPs, reflecting changes in the orienting of spatial attention in the absence of visuo‐spatial distortion or visuo‐proprioceptive mismatch. Findings are relevant to theories of how sensorimotor adaptation can modulate attention, and may open new avenues for treatment of spatial neglect.