Features of the Retinotopic Representation in the Visual Wulst of a Laterally Eyed Bird,the Zebra Finch (Taeniopygia guttata)

The visual wulst of the zebra finch comprises at least two retinotopic maps of the contralateral eye. As yet, it is not known how much of the visual field is represented in the wulst neuronal maps, how the organization of the maps is related to the retinal architecture, and how information from the ipsilateral eye is involved in the activation of the wulst. Here, we have used autofluorescent flavoprotein imaging and classical anatomical methods to investigate such characteristics of the most posterior map of the multiple retinotopic representations. We found that the visual wulst can be activated by visual stimuli from a large part of the visual field of the contralateral eye. Horizontally, the visual field representation extended from -5° beyond the beak tip up to +125° laterally. Vertically, a small strip from -10° below to about +25° above the horizon activated the visual wulst. Although retinal ganglion cells had a much higher density around the fovea and along a strip extending from the fovea towards the beak tip, these areas were not overrepresented in the wulst map. The wulst area activated from the foveal region of the ipsilateral eye, overlapped substantially with the middle of the three contralaterally activated regions in the visual wulst, and partially with the other two. Visual wulst activity evoked by stimulation of the frontal visual field was stronger with contralateral than with binocular stimulation. This confirms earlier electrophysiological studies indicating an inhibitory influence of the activation of the ipsilateral eye on wulst activity elicited by stimulating the contralateral eye. The lack of a foveal overrepresentation suggests that identification of objects may not be the primary task of the zebra finch visual wulst. Instead, this brain area may be involved in the processing of visual information necessary for spatial orientation.     

Parkinson-Related Changes of Activation in Visuomotor Brain Regions during Perceived Forward Self-Motion

Radial expanding optic flow is a visual consequence of forward locomotion. Presented on screen, it generates illusionary forward self-motion, pointing at a close vision-gait interrelation. As particularly parkinsonian gait is vulnerable to external stimuli, effects of optic flow on motor-related cerebral circuitry were explored with functional magnetic resonance imaging in healthy controls (HC) and patients with Parkinson’s disease (PD). Fifteen HC and 22 PD patients, of which 7 experienced freezing of gait (FOG), watched wide-field flow, interruptions by narrowing or deceleration and equivalent control conditions with static dots. Statistical parametric mapping revealed that wide-field flow interruption evoked activation of the (pre-)supplementary motor area (SMA) in HC, which was decreased in PD. During wide-field flow, dorsal occipito-parietal activations were reduced in PD relative to HC, with stronger functional connectivity between right visual motion area V5, pre-SMA and cerebellum (in PD without FOG). Non-specific ‘changes’ in stimulus patterns activated dorsolateral fronto-parietal regions and the fusiform gyrus. This attention-associated network was stronger activated in HC than in PD. PD patients thus appeared compromised in recruiting medial frontal regions facilitating internally generated virtual locomotion when visual motion support falls away. Reduced dorsal visual and parietal activations during wide-field optic flow in PD were explained by impaired feedforward visual and visuomotor processing within a magnocellular (visual motion) functional chain. Compensation of impaired feedforward processing by distant fronto-cerebellar circuitry in PD is consistent with motor responses to visual motion stimuli being either too strong or too weak. The ‘change’-related activations pointed at covert (stimulus-driven) attention.     

Illusions of Visual Motion Elicited by Electrical Stimulation of Human MT Complex

Human cortical area MT⁺ (hMT⁺) is known to respond to visual motion stimuli, but its causal role in the conscious experience of motion remains largely unexplored. Studies in non-human primates demonstrate that altering activity in area MT can influence motion perception judgments, but animal studies are inherently limited in assessing subjective conscious experience. In the current study, we use functional magnetic resonance imaging (fMRI), intracranial electrocorticography (ECoG), and electrical brain stimulation (EBS) in three patients implanted with intracranial electrodes to address the role of area hMT⁺ in conscious visual motion perception. We show that in conscious human subjects, reproducible illusory motion can be elicited by electrical stimulation of hMT⁺. These visual motion percepts only occurred when the site of stimulation overlapped directly with the region of the brain that had increased fMRI and electrophysiological activity during moving compared to static visual stimuli in the same individual subjects. Electrical stimulation in neighboring regions failed to produce illusory motion. Our study provides evidence for the sufficient causal link between the hMT⁺ network and the human conscious experience of visual motion. It also suggests a clear spatial relationship between fMRI signal and ECoG activity in the human brain.     

Crossmodal spatial influences of touch on extrastriate visual areas take current gaze direction into account

Recent results indicate that crossmodal interactions can affect activity in cortical regions traditionally regarded as "unimodal." Previously we found that combining touch on one hand with visual stimulation in the anatomically corresponding hemifield could boost responses in contralateral visual cortex. Here we manipulated which visual hemifield corresponded to the location of the stimulated hand, by changing gaze direction such that right-hand touch could now arise in either the left or right visual field. Crossmodal effects on visual cortex switched from one hemisphere to the other, depending on gaze direction, regardless of whether the hand was seen. This indicates that crossmodal influences of touch upon visual cortex depend on spatial alignment for the multimodal stimuli, with gaze posture taken into account.     

Optical imaging of nociception in primary somatosensory cortex of non-human primates

While the activation of primary somatosensory (SI) cortex during pain perception is consistently reported in functional imaging studies on normal subjects and chronic pain patients, the specific roles of SI, particularly the subregions within SI, in the processing of sensory aspects of pain are still largely unknown. Using optical imaging of intrinsic signal (OIS) and single unit electrophysiology, we studied cortical activation patterns within SI cortex (among Brodmann areas 3a, 3b and 1) and signal amplitude changes to various intensities of non-nociceptive, thermal nociceptive and mechanical nociceptive stimulation of individual distal finerpads in anesthetized squirrel monkeys. We have demonstrated that areas 3a and 1 are preferentially involved in the processing of nociceptive information while areas 3b and 1 are preferentially activated in the processing of non-nociceptive (touch) information. Nociceptive activations of individual fingerpad were organized topographically suggesting that nociceptive topographic map exits in areas 3a and 1. Signal amplitude was enhanced to increasing intensity of mechanical nociceptive stimuli in areas 3a, 3b and 1. Within area 1, nociceptive response co-localizes with the non-nociceptive response. Therefore, we hypothesize that nocicepitve information is area-specifically represented within SI cortex, in which nociceptive inputs are preferentially represented in areas 3a and 1 while non-nociceptive inputs are preferentially represented in areas 3b and 1.     

The Effect of 10 Hz Repetitive Transcranial Magnetic Stimulation of Posterior Parietal Cortex on Visual Attention

Repetitive transcranial magnetic stimulation (rTMS) of the posterior parietal cortex (PPC) at frequencies lower than 5 Hz transiently inhibits the stimulated area. In healthy participants, such a protocol can induce a transient attentional bias to the visual hemifield ipsilateral to the stimulated hemisphere. This bias might be due to a relatively less active stimulated hemisphere and a relatively more active unstimulated hemisphere. In a previous study, Jin and Hilgetag (2008) tried to switch the attention bias from the hemifield ipsilateral to the hemifield contralateral to the stimulated hemisphere by applying high frequency rTMS. High frequency rTMS has been shown to excite, rather than inhibit, the stimulated brain area. However, the bias to the ipsilateral hemifield was still present. The participants’ performance decreased when stimuli were presented in the hemifield contralateral to the stimulation site. In the present study we tested if this unexpected result was related to the fact that participants were passively resting during stimulation rather than performing a task. Using a fully crossed factorial design, we compared the effects of high frequency rTMS applied during a visual detection task and high frequency rTMS during passive rest on the subsequent offline performance in the same detection task. Our results were mixed. After sham stimulation, performance was better after rest than after task. After active 10 Hz rTMS, participants’ performance was overall better after task than after rest. However, this effect did not reach statistical significance. The comparison of performance after rTMS with task and performance after sham stimulation with task showed that 10 Hz stimulation significantly improved performance in the whole visual field. Thus, although we found a trend to better performance after rTMS with task than after rTMS during rest, we could not reject the hypothesis that high frequency rTMS with task and high frequency rTMS during rest equally affect performance.     

Multimodal spatial representations engaged in human parietal cortex during both saccadic and manual spatial orienting

BACKGROUND: Recent neuroimaging studies have found that several areas of the human brain, including parietal regions, can respond multimodally. But given single-cell evidence that responses in primate parietal cortex can be motor-related, some of the human multimodal activations might reflect convergent activation of potentially motor-related areas, rather than multimodal representations of space independent of motor factors. Here we crossed sensory stimulation of different modalities (vision or touch, in left or right hemifield) with spatially directed responses to such stimulation by different effector-systems (saccadic or manual). RESULTS: The fMRI results revealed representations of contralateral space in both the posterior part of the superior parietal gyrus and the anterior intraparietal sulcus that activated independently of both sensory modality and motor response. Multimodal saccade-related or manual-related activations were found, by contrast, in different regions of parietal cortex. CONCLUSIONS: Whereas some parietal regions have specific motor functions, others are engaged during the execution of movements to the contralateral hemifield irrespective of both input modality and the type of motor effector.     

Single cell responses from the optic tectum of the zebra finch (Taeniopygia guttata castanotis Gould)

Responses of neurons of the optic tectum, the prominent, highly laminated mesencephalic station of the tectofugal visual pathway in birds, to computer-generated and other visual stimuli were examined in zebra finches. Our study shows that the contralateral retina projects to the tectum in topographic order. The representation of the visual field is tilted against the horizon by 22°. The representation of the contralateral hemifield extends to the ipsilateral side by 15°. Most neurons have receptive fields with excitatory centres of different shapes and inhibitory surround. A new type of neuronal receptive field is described which has an excitatory centre and a surround which is movement sensitive and preferably excited by very small spots. The first type of neurons is mostly located in upper tectal layers, the latter only in deeper layers. Excitatory centre sizes increase with depth, and there is a tendency of smaller receptive fields in the foveal region. The representation of the frontal visual field does not show specializations which could be expected if it were used for fixation of grain during pecking. Our results are in accordance with previous behavioural experiments. Accepted: 30 April 1999     

Identification of Retinal Ganglion Cells and Their Projections Involved in Central Transmission of Information about Upward and Downward Image Motion

The direction of image motion is coded by direction-selective (DS) ganglion cells in the retina. Particularly, the ON DS ganglion cells project their axons specifically to terminal nuclei of the accessory optic system (AOS) responsible for optokinetic reflex (OKR). We recently generated a knock-in mouse in which SPIG1 (SPARC-related protein containing immunoglobulin domains 1)-expressing cells are visualized with GFP, and found that retinal ganglion cells projecting to the medial terminal nucleus (MTN), the principal nucleus of the AOS, are comprised of SPIG1⁺ and SPIG1⁻ ganglion cells distributed in distinct mosaic patterns in the retina. Here we examined light responses of these two subtypes of MTN-projecting cells by targeted electrophysiological recordings. SPIG1⁺ and SPIG1⁻ ganglion cells respond preferentially to upward motion and downward motion, respectively, in the visual field. The direction selectivity of SPIG1⁺ ganglion cells develops normally in dark-reared mice. The MTN neurons are activated by optokinetic stimuli only of the vertical motion as shown by Fos expression analysis. Combination of genetic labeling and conventional retrograde labeling revealed that axons of SPIG1⁺ and SPIG1⁻ ganglion cells project to the MTN via different pathways. The axon terminals of the two subtypes are organized into discrete clusters in the MTN. These results suggest that information about upward and downward image motion transmitted by distinct ON DS cells is separately processed in the MTN, if not independently. Our findings provide insights into the neural mechanisms of OKR, how information about the direction of image motion is deciphered by the AOS.     

Central mechanisms of tactile shape perception

Studies show that while the cortical mechanisms of two-dimensional (2D) form and motion processing are similar in touch and vision, the mechanisms of three-dimensional (3D) shape processing are different. 2D form and motion are processed in areas 3b and 1 of SI cortex by neurons with receptive fields (RFs) composed of excitatory and inhibitory subregions. 3D shape is processed in area 2 and SII and relies on the integration of cutaneous and proprioceptive inputs. The RFs of SII neurons vary in size and shape with heterogeneous structures consisting of orientation-tuned fingerpads mixed with untuned excitatory or inhibitory fingerpads. Furthermore, the sensitivity of the neurons to cutaneous inputs changes with hand conformation. We hypothesize that these RFs are the kernels underlying tactile object recognition.     

Ten_m3 Regulates Eye-Specific Patterning in the Mammalian Visual Pathway and Is Required for Binocular Vision

Binocular vision requires an exquisite matching of projections from each eye to form a cohesive representation of the visual world. Eye-specific inputs are anatomically segregated, but in register in the visual thalamus, and overlap within the binocular region of primary visual cortex. Here, we show that the transmembrane protein Ten_m3 regulates the alignment of ipsilateral and contralateral projections. It is expressed in a gradient in the developing visual pathway, which is consistently highest in regions that represent dorsal visual field. Mice that lack Ten_m3 show profound abnormalities in mapping of ipsilateral, but not contralateral, projections, and exhibit pronounced deficits when performing visually mediated behavioural tasks. It is likely that the functional deficits arise from the interocular mismatch, because they are reversed by acute monocular inactivation. We conclude that Ten_m3 plays a key regulatory role in the development of aligned binocular maps, which are required for normal vision.     

A preference for contralateral stimuli in human object- and face-selective cortex

Visual input from the left and right visual fields is processed predominantly in the contralateral hemisphere. Here we investigated whether this preference for contralateral over ipsilateral stimuli is also found in high-level visual areas that are important for the recognition of objects and faces. Human subjects were scanned with functional magnetic resonance imaging (fMRI) while they viewed and attended faces, objects, scenes, and scrambled images in the left or right visual field. With our stimulation protocol, primary visual cortex responded only to contralateral stimuli. The contralateral preference was smaller in object- and face-selective regions, and it was smallest in the fusiform gyrus. Nevertheless, each region showed a significant preference for contralateral stimuli. These results indicate that sensitivity to stimulus position is present even in high-level ventral visual cortex.     

Delayed striate cortical activation during spatial attention

Recordings of event-related potentials (ERPs) and event-related magnetic fields (ERMFs) were combined with functional magnetic resonance imaging (fMRI) to study visual cortical activity in humans during spatial attention. While subjects attended selectively to stimulus arrays in one visual field, fMRI revealed stimulus-related activations in the contralateral primary visual cortex and in multiple extrastriate areas. ERP and ERMF recordings showed that attention did not affect the initial evoked response at 60-90 ms poststimulus that was localized to primary cortex, but a similarly localized late response at 140-250 ms was enhanced to attended stimuli. These findings provide evidence that the primary visual cortex participates in the selective processing of attended stimuli by means of delayed feedback from higher visual-cortical areas.     

Dynamic Changes in Brain Activations and Functional Connectivity during Affectively Different Tactile Stimuli

In the present study, we compared brain activations produced by pleasant, neutral and unpleasant touch, to the anterior lateral surface of lower leg of human subjects. It was found that several brain regions, including the contralateral primary somatosensory area (SI), bilateral secondary somatosensory area (SII), as well as contralateral middle and posterior insula cortex were commonly activated under the three touch conditions. In addition, pleasant and unpleasant touch conditions shared a few brain regions including the contralateral posterior parietal cortex (PPC) and bilateral premotor cortex (PMC). Unpleasant touch specifically activated a set of pain-related brain regions such as contralateral supplementary motor area (SMA) and dorsal parts of bilateral anterior cingulated cortex, etc. Brain regions specifically activated by pleasant touch comprised bilateral lateral orbitofrontal cortex (OFC), posterior cingulate cortex (PCC), medial prefrontal cortex (mPFC), intraparietal cortex and left dorsal lateral prefrontal cortex (DLPFC). Using a novel functional connectivity model based on graph theory, we showed that a series of brain regions related to affectively different touch had significant functional connectivity during the resting state. Furthermore, it was found that such a network can be modulated between affectively different touch conditions.     

Dynamic spatial coding within the dorsal frontoparietal network during a visual search task

To what extent are the left and right visual hemifields spatially coded in the dorsal frontoparietal attention network? In many experiments with neglect patients, the left hemisphere shows a contralateral hemifield preference, whereas the right hemisphere represents both hemifields. This pattern of spatial coding is often used to explain the right-hemispheric dominance of lesions causing hemispatial neglect. However, pathophysiological mechanisms of hemispatial neglect are controversial because recent experiments on healthy subjects produced conflicting results regarding the spatial coding of visual hemifields. We used an fMRI paradigm that allowed us to distinguish two attentional subprocesses during a visual search task. Either within the left or right hemifield subjects first attended to stationary locations (spatial orienting) and then shifted their attentional focus to search for a target line. Dynamic changes in spatial coding of the left and right hemifields were observed within subregions of the dorsal front-parietal network: During stationary spatial orienting, we found the well-known spatial pattern described above, with a bilateral hemifield representation in the right hemisphere and a contralateral preference in the left hemisphere. However, during search, the right hemisphere had a contralateral preference and the left hemisphere equally represented both hemifields. This finding leads to novel perspectives regarding models of visuospatial attention and hemispatial neglect.     

Movement in the normal visual hemifield induces a percept in the 'blind' hemifield of a human hemianope.

We have investigated visual responses to moving stimuli presented to the normal hemifield of a hemianope, GY, who exhibits residual visual function in his right, ''blind'' hemifield. Preliminary experiments established that his perception of moving stimuli localized in his ''blind'' hemifield is retained when a similar stimulus is presented simultaneously in the normal hemifield. In response to a grating stimulus moving horizontally towards fixation in the non-foveal region of the normal, left hemifield, he perceives in addition to a normal motion percept in the left hemifield, a sensation of movement localized in the right hemifield. Qualitatively, this latter is indistinguishable from responses elicited by direct stimulation localized within his ''blind'' hemifield by moving stimuli. We have investigated the characteristics of the mechanisms which induce the ''blind'' field component of GY''s responses to stimulation of the normal hemifield. We show that GY''s sensitivity for detection of movement localized within his ''blind'' hemifield is dependent on the direction of movement, the contrast and the velocity of a grating presented to the normal hemifield. No induced effects were recorded in response to colour or to non-moving, flickering stimuli. We examine the possible contribution of scattered light to our observations, and eliminate this factor by consideration of our experimental results. We discuss the neural mechanisms which may be involved in this response.     

‘Vector white noise’: a technique for mapping the motion receptive fields of direction-selective visual neurons

A technique is described and tested for mapping the sensitivities and preferred directions of motion at different locations within the receptive fields of direction-selective motion-detecting visual neurons. The procedure is to record the responses to a number of visual stimuli, each stimulus presentation consisting of a set of short, randomly-oriented, moving bars arranged in a square grid. Each bar moves perpendicularly to its long axis. The vector describing the sensitivity and preferred direction of motion at each grid location is obtained as a sum of the unit vectors defining the directions of motion of the bars in each of the stimuli at that location, weighted by the strengths of the corresponding responses. The resulting vector field specifies the optimum flow field for the neuron. The advantage of this technique over the conventional approach of probing the receptive field sequentially at each grid location is that the parallel nature of the stimulus is sensitive to nonlinear interactions (such as shunting inhibition for mutual facilitation) between different regions of the visual field. The technique is used to determine accurately the motion receptive fields of direction-selective motion detecting neurons in the optic lobes of insects. It is potentially applicable to motion-sensitive neurons with highly structured receptive fields, such as those in the optic tectum of the pigeon or in area MST of the monkey.     

Distractor-target interactions during directed visual attention

The spatial extent of directed visual attention (DVA) was examined in a series of experiments using precuing in a suprathreshold luminance detection (reaction time) paradigm. Previous findings (Hughes, H. C. and Zimba, L. D. J. Exp. Psychol.; Human Percept Perf., 1985, 11, 409-430) indicated that, in an empty visual field, the effects of DVA were primarily manifest as a uniform elevation of response times to all probe targets in the hemifield contralateral to the observer's expectancy. The present experiments were designed to determine whether increased spatial selectivity could be found when luminous markers indicated the exact location of the expected visual target. To maintain equivalent states of adaptation in both hemifields, luminous markers were also present at the same location in the contralateral hemifield. In general, hemifield effects were again obtained, but with two notable exceptions. First, marking locations in the unattended hemifield produced a local increase (enhanced interference) in RTs above the level characteristic of other locations within that hemifield. Second, when multiple locations were indicated with identical luminous markers, graded costs were obtained in both hemifields. However, scaling the markers according to estimates of cortical magnification factor (M) substantially reduced the slope of these inhibitory gradients, and the results once again approached those characteristic of an unstructured visual field. The findings suggest that when attention is directed to a marked location along the horizontal meridian, a transition in performance typically occurs at the vertical meridian. In addition, irrelevant stimuli some distance from the attentional focus interfere with detection times to unexpected targets that appear in the same vicinity. This interference may relate to an enhanced susceptibility to spatial interactions between the distractors and target away from the attentional focus. The interference appears to extend over a constant area of visual cortex, since it is reduced when the markers are M-scaled.     

A Bio-Inspired,Motion-Based Analysis of Crowd Behavior Attributes Relevance to Motion Transparency,Velocity Gradients,and Motion Patterns

The analysis of motion crowds is concerned with the detection of potential hazards for individuals of the crowd. Existing methods analyze the statistics of pixel motion to classify non-dangerous or dangerous behavior, to detect outlier motions, or to estimate the mean throughput of people for an image region. We suggest a biologically inspired model for the analysis of motion crowds that extracts motion features indicative for potential dangers in crowd behavior. Our model consists of stages for motion detection, integration, and pattern detection that model functions of the primate primary visual cortex area (V1), the middle temporal area (MT), and the medial superior temporal area (MST), respectively. This model allows for the processing of motion transparency, the appearance of multiple motions in the same visual region, in addition to processing opaque motion. We suggest that motion transparency helps to identify “danger zones” in motion crowds. For instance, motion transparency occurs in small exit passages during evacuation. However, motion transparency occurs also for non-dangerous crowd behavior when people move in opposite directions organized into separate lanes. Our analysis suggests: The combination of motion transparency and a slow motion speed can be used for labeling of candidate regions that contain dangerous behavior. In addition, locally detected decelerations or negative speed gradients of motions are a precursor of danger in crowd behavior as are globally detected motion patterns that show a contraction toward a single point. In sum, motion transparency, image speeds, motion patterns, and speed gradients extracted from visual motion in videos are important features to describe the behavioral state of a motion crowd.     

Selectivity to Translational Egomotion in Human Brain Motion Areas

The optic flow generated when a person moves through the environment can be locally decomposed into several basic components, including radial, circular, translational and spiral motion. Since their analysis plays an important part in the visual perception and control of locomotion and posture it is likely that some brain regions in the primate dorsal visual pathway are specialized to distinguish among them. The aim of this study is to explore the sensitivity to different types of egomotion-compatible visual stimulations in the human motion-sensitive regions of the brain. Event-related fMRI experiments, 3D motion and wide-field stimulation, functional localizers and brain mapping methods were used to study the sensitivity of six distinct motion areas (V6, MT, MST+, V3A, CSv and an Intra-Parietal Sulcus motion [IPSmot] region) to different types of optic flow stimuli. Results show that only areas V6, MST+ and IPSmot are specialized in distinguishing among the various types of flow patterns, with a high response for the translational flow which was maximum in V6 and IPSmot and less marked in MST+. Given that during egomotion the translational optic flow conveys differential information about the near and far external objects, areas V6 and IPSmot likely process visual egomotion signals to extract information about the relative distance of objects with respect to the observer. Since area V6 is also involved in distinguishing object-motion from self-motion, it could provide information about location in space of moving and static objects during self-motion, particularly in a dynamically unstable environment.