Cortical response field dynamics in cat visual cortex

Little is known about the "inverse" of the receptive field--the region of cortical space whose spatiotemporal pattern of electrical activity is influenced by a given sensory stimulus. We refer to this activated area as the cortical response field, the properties of which remain unexplored. Here, the dynamics of cortical response fields evoked in visual cortex by small, local drifting-oriented gratings were explored using voltage-sensitive dyes. We found that the cortical response field was often characterized by a plateau of activity, beyond the rim of which activity diminished quickly. Plateau rim location was largely independent of stimulus orientation. However, approximately 20 ms following plateau onset, 1-3 peaks emerged on it and were amplified for 25 ms. Spiking was limited to the peak zones, whose location strongly depended on stimulus orientation. Thus, alongside selective amplification of a spatially restricted suprathreshold response, wider activation to just below threshold encompasses all orientation domains within a well-defined retinotopic vicinity of the current stimulus, priming the cortex for processing of subsequent stimuli.     

Cortical specialization for processing first- and second-order motion

Distinct mechanisms underlying the visual perception of luminance-(first-order) and contrast-defined (second-order) motion havebeen proposed from electrophysiological, human psychophysicaland neurological studies; however a cortical specializationfor these mechanisms has proven elusive. Here human brain imaging combined with psychophysical methods was used to assess corticalspecializations for processing these two kinds of motion. Acommon stimulus construction was employed, controlling for differencesin spatial and temporal properties, psychophysical performanceand attention. Distinct cortical regions have been found preferentiallyprocessing either first- or second-order motion, both in occipitaland parietal lobes, producing the first physiological evidencein humans to support evidence from psychophysical studies, brainlesion sites and computational models. These results provideevidence for the idea that first-order motion is computed inV1 and second-order motion in later occipital visual areas,and additionally suggest a functional dissociation between thesetwo kinds of motion beyond the occipital lobe.     

Cortical substrates supporting visual search in humans.

Serial and parallel visual search tasks were presented to patients with focal lesions in dorsolateral frontal, lateral parietal, or temporal-parietal cortex. In the unilateral display conditions, search efficiency in all patient groups was similar to the normal control group for stimuli both on the ipsi- and on the contralesional side of the displays. In contrast, in the bilateral display conditions, all patient groups showed a marked delay in initiating search on the side contralateral to the lesion as compared to normal controls. This delay was more pronounced when attention demands on the ipsilateral side increased, either by making target-distractor discrimination more difficult (serial search task), or by increasing the number of ipsilateral distractor items. The contralateral deficit was evident in all patient groups, supporting the notion that dorsolateral frontal as well as posterior parietal and temporal-parietal cortex plays a critical role in visual spatial attention.     

High-frequency activity in human visual cortex is modulated by visual motion strength

A central goal in systems neuroscience is to understand how the brain encodes the intensity of sensory features. We used whole-head magnetoencephalography to investigate whether frequency-specific neuronal activity in the human visual cortex is systematically modulated by the intensity of an elementary sensory feature such as visual motion. Visual stimulation induced a tonic increase of neuronal activity at frequencies above 50 Hz. In order to define a functional frequency band of neuronal activity, we parametrically investigated which frequency band displays the strongest monotonic increase of responses with strength of visual motion. Consistently in all investigated subjects, this analysis resulted in a functional frequency band in the high gamma range from about 60 to 100 Hz in which activity reliably increased with visual motion strength. Using distributed source reconstruction, we found that this increase of high-frequency neuronal activity originates from several extrastriate cortical regions specialized in motion processing. We conclude that high-frequency activity in the human visual motion pathway may be relevant for encoding the intensity of visual motion signals.     

Cortical area MSTd combines visual cues to represent 3-D self-movement

As arboreal primates move through the jungle, they are immersed in visual motion that they must distinguish from the movement of predators and prey. We recorded dorsal medial superior temporal (MSTd) cortical neuronal responses to visual motion stimuli simulating self-movement and object motion. MSTd neurons encode the heading of simulated self-movement in three-dimensional (3-D) space. 3-D heading responses can be evoked either by the large patterns of visual motion in optic flow or by the visual object motion seen when an observer passes an earth-fixed landmark. Responses to naturalistically combined optic flow and object motion depend on their relative directions: an object moving as part of the optic flow field has little effect on neuronal responses. In contrast, an object moving separately from the optic flow field has large effects, decreasing the amplitude of the population response and shifting the population's heading estimate to match the direction of object motion as the object moves toward central vision. These effects parallel those seen in human heading perception with minimal effects of objects moving with the optic flow and substantial effects of objects violating the optic flow. We conclude that MSTd can contribute to navigation by supporting 3-D heading estimation, potentially switching from optic flow to object cues when a moving object passes in front of the observer.     

Opposite dependencies on visual motion coherence in human area MT+ and early visual cortex

In order to understand the relationship between brain activity and visual motion perception, knowledge of the cortical areas participating in signal processing alone is insufficient. Rather knowledge on how responses vary with the characteristics of visual motion is necessary. In this study, we measured whole brain activity using magnetoencephalography in humans discriminating the global motion direction of a random dot kinematogram whose strength was systematically varied by the percentage of coherently moving dot elements. Spectral analysis revealed 2 components correlating with motion coherence. A first component in the low-frequency domain ( approximately 3 Hz), linearly increasing with motion coherence, could be attributed to visual cortex including human area middle temporal (MT) +. A second component oscillating in the alpha frequency range and emerging after stimulus offset showed the inverse dependence on motion coherence and arose from early visual cortex. Based on these results, we first of all conclude that motion coherence is reflected in the population response of human extrastriate cortex. Second, we suggest that the occipital alpha activity represents a gating mechanism protecting visual motion integration in later cortical areas from disturbing upcoming signals.     

Cortical neuronal responses to optic flow are shaped by visual strategies for steering

We hypothesized that neuronal responses to virtual self-movement would be enhanced during steering tasks. We recorded the activity of medial superior temporal (MSTd) neurons in monkeys trained to steer a straight-ahead course, using optic flow. We found smaller optic flow responses during active steering than during the passive viewing of the same stimuli. Behavioral analysis showed that the monkeys had learned to steer using local motion cues. Retraining the monkeys to use the global pattern of optic flow reversed the effects of the active-steering task: active steering then evoked larger responses than passive viewing. We then compared the responses of neurons during active steering by local motion and by global patterns: Local motion trials promoted the use of local dot movement near the center of the stimulus by occluding the peripheral visual field midway through the trial. Global pattern trials promoted the use of radial pattern movement by occluding the central visual field midway through the trial. In this study, identical full-field optic-flow stimuli evoked larger responses in global-pattern trials than in local motion trials. We conclude that the selection of specific visual cues reflects strategies for active steering and alters MSTd neuronal responses to optic flow.     

The integration of colour and motion by the human visual brain

Objects in the visual scene are defined by different cues such as colour and motion. Through the integration of these cues the visual system is able to utilize different sources of information, thus enhancing its ability to discriminate objects from their backgrounds. In the following experiments, we investigate the neural mechanisms of cue integration in the human. We show, using functional magnetic resonance imaging (fMRI), that both colour and motion defined shapes activate the lateral occipital complex (LOC) and that shapes defined by both colour and motion simultaneously activate the anterior-ventral margins of this area more strongly than shapes defined by either cue alone. This suggests that colour and motion cues are integrated in the LOC and possibly a neighbouring, more anterior, region. We support this result using an fMR adaptation technique, demonstrating that a region of the LOC adapts on repeated presentations of a shape regardless of the cue that is used to define it and even if the cue is varied. This result raises the possibility that the LOC contains cue-invariant neurons that respond to shapes regardless of the cue that is used to define them. We propose that such neurons could integrate signals from different cues, making them more responsive to objects defined by more than one cue, thus increasing the ability of the observer to recognize them.     

The multiple roles of visual cortical areas MT/MST in remembering the direction of visual motion

Although the role of cortical areas MT and MST (MT/MST) in the processing of directional motion information is well established, little is known about the way these areas contribute to the execution of complex behavioral tasks requiring the use of such information. We tested monkeys with unilateral lesions of these areas on a visual working memory task in which motion signals not only had to be encoded, but also stored for brief periods of time and then retrieved. The monkeys compared the directions of motion of two random-dot stimuli, sample and test, separated by a temporal delay. By increasing the temporal delay and spatially separating the two stimuli, placing one in the affected visual field and the other in the intact visual field, we were able to assess the contribution of MT/MST to specific components of the task: encoding (sample), retention (delay) and encoding/retrieval/comparison (test). We found that the effects of MT/MST lesions on specific components depended upon the demands of the task and the nature of the visual motion stimuli. Whenever stimuli consisted of random dots moving in a broad range of directions, MT/MST lesions appeared to affect encoding. Furthermore, when the lesions affected encoding of the sample, retention of the direction of stimulus motion was also affected. However, when the stimulus was coherent and the emphasis of the task was on the comparison of small direction differences, the absence of MT/MST had major impact on the retrieval/comparison component of the task and not on encoding or storage.     

Role of the superior temporal region in human visual motion perception

While moving objects are usually seen using luminance (first-order) cues, humans can perceive the motion of objects via non-luminance (second-order) cues. Contrary to previous case reports, no physiological studies have elucidated distinct differences in the cortical regions involved in first- and second-order motion processes. We investigated brain responses related to these two types of motion perception in human subjects using 3 T functional magnetic resonance imaging and strictly controlled apparent motion stimulus pairs. Comparison of brain activation to moving versus static states of each motion stimulus isolated cortical activity related to each type of motion perception. We found a selective neural response to second-order motion stimulus in the anterior part of the superior temporal sulcus (STS) contralateral to stimulus presentation and cue-invariant activation of MT/V5+. No significant activation in the STS was observed by the first-order motion, even when its visibility was reduced to levels comparable to that of second-order motion. Furthermore, the STS demonstrated significant activation for highly visible motion stimulus with both first- and second-order attributes. The STS represents the cardinal structure for perception of second-order motions, although further studies are needed to elucidate the exact neural process occurring in this area.     

The brain structural hub of interhemispheric information integration for visual motion perception

We investigated the key anatomical structures mediating interhemispheric integration during the perception of apparent motion across the retinal midline. Previous studies of commissurotomized patients suggest that subcortical structures mediate interhemispheric transmission but the specific regions involved remain unclear. Here, we exploit interindividual variations in the propensity of normal subjects to perceive horizontal motion, in relation to vertical motion. We characterize these differences psychophysically using a Dynamic Dot Quartet (an ambiguous stimulus that induces illusory motion). We then tested for correlations between a tendency to perceive horizontal motion and fractional anisotropy (FA) (from structural diffusion tensor imaging), over subjects. FA is an indirect measure of the orientation and integrity of white matter tracts. Subjects who found it easy to perceive horizontal motion showed significantly higher FA values in the pulvinar. Furthermore, fiber tracking from an independently identified (subject-specific) visual motion area converged on the pulvinar nucleus. These results suggest that the pulvinar is an anatomical hub and may play a central role in interhemispheric integration.     

A comparison of visual and auditory motion processing in human cerebral cortex

Visual and auditory motion information can be used together to provide complementary information about the movement of objects. To investigate the neural substrates of such cross-modal integration, functional magnetic resonance imaging was used to assess brain activation while subjects performed separate visual and auditory motion discrimination tasks. Areas of unimodal activation included the primary and/or early sensory cortex for each modality plus additional sites extending toward parietal cortex. Areas conjointly activated by both tasks included lateral parietal cortex, lateral frontal cortex, anterior midline and anterior insular cortex. The parietal site encompassed distinct, but partially overlapping, zones of activation in or near the intraparietal sulcus (IPS). A subsequent task requiring an explicit cross-modal speed comparison revealed several foci of enhanced activity relative to the unimodal tasks. These included the IPS, anterior midline, and anterior insula but not frontal cortex. During the unimodal auditory motion task, portions of the dorsal visual motion system showed signals depressed below resting baseline. Thus, interactions between the two systems involved either enhancement or suppression depending on the stimuli present and the nature of the perceptual task. Together, these results identify human cortical regions involved in polysensory integration and the attentional selection of cross-modal motion information.     

Performance Dip in motor response induced by task-irrelevant weaker coherent visual motion signals

The Performance Dip is a newly characterized behavioral phenomenon, where, paradoxically, a weaker task-irrelevant visual stimulus causes larger disturbances on the accuracy of a main letter identification task than a stronger stimulus does. Understanding mechanisms of the Performance Dip may provide insight into unconsciousness behavior. Here, we investigated the generalization of the Performance Dip. Specifically, we tested whether the Performance Dip occurs in a motion-related Simon task, and if so, whether the Performance Dip involves the same brain region, that is, the dorsolateral prefrontal cortex (DLPFC), previously implicated in the Performance Dip, or the supplementary motor area (SMA) and pre-SMA, implicated in a motion-related Simon Task. Subjects made manual directional responses according to the color of stochastic moving dots while ignoring the global direction of moving dots, which could be either congruent or incongruent to the response appropriate to the main task. We found that weak incongruent task-irrelevant stimuli caused a Performance Dip, in which the SMA and pre-SMA, rather than DLPFC, played critical roles. Our results suggest a possible common brain mechanism across different neural circuits, in which weak, but not strong, task-irrelevant information is free from inhibition and intrudes into neural circuits relevant to the main task.     

A neural model of motion processing and visual navigation by cortical area MST

Cells in the dorsal medial superior temporal cortex (MSTd) process optic flow generated by self-motion during visually guided navigation. A neural model shows how interactions between well-known neural mechanisms (log polar cortical magnification, Gaussian motion-sensitive receptive fields, spatial pooling of motion-sensitive signals and subtractive extraretinal eye movement signals) lead to emergent properties that quantitatively simulate neurophysiological data about MSTd cell properties and psychophysical data about human navigation. Model cells match MSTd neuron responses to optic flow stimuli placed in different parts of the visual field, including position invariance, tuning curves, preferred spiral directions, direction reversals, average response curves and preferred locations for stimulus motion centers. The model shows how the preferred motion direction of the most active MSTd cells can explain human judgments of self-motion direction (heading), without using complex heading templates. The model explains when extraretinal eye movement signals are needed for accurate heading perception, and when retinal input is sufficient, and how heading judgments depend on scene layouts and rotation rates.     

Synchrony dynamics in monkey V1 predict success in visual detection

Behavioral measures such as expectancy and attention have been associated with the strength of synchronous neural activity. On this basis, it is hypothesized that synchronous activity affects our ability to detect and recognize visual objects. To investigate the role of synchronous activity in visual perception, we studied the magnitude and precision of correlated activity, before and after stimulus presentation within the visual cortex (V1), in relation to a monkey's performance in a figure-ground discrimination task. We show that during the period of stimulus presentation a transition in synchronized activity occurs that is characterized by a reduction of the correlation peak height and width. Before stimulus onset, broad peak correlations are observed that change towards thin peak correlations after stimulus onset, due to a specific decrease of low-frequency components. The magnitude of the transition in correlated activity is larger, i.e. a stronger desynchronization occurs, when the animal perceives the stimulus correctly than when the animal fails to detect the stimulus. These results therefore show that a transition in synchronous firing is important for the detection of sensory stimuli. We hypothesize that the transition in synchrony reflects a change from loose and global neuronal interactions towards a finer temporal and spatial scale of neuronal interactions, and that such a change in neuronal interactions is required for figure-ground discrimination.     

Selective attention increases the dependency of cortical responses on visual motion coherence in man

Attention improves visual discrimination and consequently allows to discern stimuli with low signal-to-noise ratios that otherwise would remain undetected. We used magnetoencephalography (MEG) to test whether neuromagnetic responses recorded from occipito-temporal cortex, reflecting the size of visual motion signals embedded in noise (motion coherence), would mirror the perceptual changes induced by attention. Attention directed to a given hemifield increased and decreased the coherence modulation of the MEG response over contralateral and ipsilateral visual cortex, respectively, indicating a change in the neuronal signal-to-noise ratio at the population level.     

Training-induced recovery of visual motion perception after extrastriate cortical damage in the adult cat

Unilateral ibotenic acid lesions of the lateral suprasylvian (LS) cortex severely impair the ability of cats to integrate local motion signals (measured as direction range thresholds) and to extract motion signals from noise (measured as motion signal thresholds) in their contra-lesional visual hemifields. These deficits were found up to several months after the lesions and were limited to thresholds measured with random-dot stimuli, while contrast sensitivity for discriminating the direction of motion of sine-wave gratings remained unaffected. Our goal was to determine whether deficits of complex motion perception could recover and whether the recovery was spontaneous or required retraining. In each cat, a single location in the impaired visual hemifield was selected for visual retraining, which required the animals to discriminate motion direction using random-dot stimuli in which the range of dot directions was varied. Fifteen to 40 days of intensive retraining led to a gradual, complete recovery of motion integration. The recovery was stimulus specific since it did not transfer from direction range to motion signal thresholds, and it was largely restricted to the visual field locations retrained. Delaying the onset of retraining by several days to several months had no significant impact on the extent or rate of recovery. Once recovery was achieved, performance remained stable over a period of several months. These results suggest that recovery of complex visual motion perception after lesions of extrastriate visual cortex is an active process that requires extensive, stimulus- and retinotopically-specific visual retraining.     

Immunocytochemical localization of pro-opiomelanocortin neurons in human brain areas subserving stimulation analgesia

The distribution of pro-opiomelanocortin (beta-endorphin, adrenocorticotropic hormone, and 16-K) neurons and fiber projections was evaluated immunocytochemically in 50-mu thick cryostat sections of human diencephalon and midbrain. Specific attention was focused upon regions in which deep brain stimulation has been most effective in the relief of selected chronic pain syndromes. This study revealed a remarkable, nearly point-to-point correlation between clinically effective stimulation sites and the distribution of pro-opiomelanocortin fibers in the human brain. Of particular interest was the dense innervation of the periventricular stratum along the third ventricle, the parafascicular centromedian region of the thalamus, and the periaqueductal gray matter of the midbrain. This study provides anatomical support for the hypothesis that beta-endorphin-containing neuronal systems may contribute to stimulation analgesia in the human.