Subcortical discrimination of unperceived objects during binocular rivalry

Rapid identification of behaviorally relevant objects is important for survival. In humans, the neural computations for visually discriminating complex objects involve inferior temporal cortex (IT). However, less detailed but faster form processing may also occur in a phylogenetically older subcortical visual system that terminates in the amygdala. We used binocular rivalry to present stimuli without conscious awareness, thereby eliminating the IT object representation and isolating subcortical visual input to the amygdala. Functional magnetic resonance imaging revealed significant brain activation in the left amygdala but not in object-selective IT in response to unperceived fearful faces compared to unperceived nonface objects. These findings indicate that, for certain behaviorally relevant stimuli, a high-level cortical representation in IT is not required for object discrimination in the amygdala.     

Hemispheric participation in the shaping of spatial-motor asymmetry in visual recognition in rats

Asymmetry of movement direction was found in Wistar rats at establishing of motor alimentary conditioned reflex to simultaneously presented visual stimuli. In the course of learning the asymmetry weakened on the whole, but some individuals retained right- or left side preference. The analysis of asymmetry change before and after unilateral cortical inactivation revealed a special role of right hemisphere influences for the formation of right-side preference and of the left hemisphere--for the choice of the left direction. The lack of asymmetry was observed at the presence of the influences from the left hemisphere cortex depressing ipsilateral nigro-striate system and activating the contralateral one. Influences of the cortex of both hemispheres reduce the absolute value of the asymmetry coefficient; the left hemisphere has a special significance for manifestation of temporal asymmetry parameters. Photic interference is a factor modulating the asymmetry. It reduces the right hemisphere activity more than that of the left one; it intensifies right hemisphere influences, contributes to the involvement of the transcallosal conduction channel in the formation of spatial-motor asymmetry.     

Category differences in brain activation studies: where do they come from?

Differences in the neural processing of six categories of pictorial stimuli (maps, body parts, objects, animals, famous faces and colours) were investigated using positron emission tomography. Stimuli were presented either with or without the written name of the picture, thereby creating a naming condition and a reading condition. As predicted, naming increased the demands on lexical processes. This was demonstrated by activation of the left temporal lobe in a posterior region associated with name retrieval in several previous studies. This lexical effect was common to all meaningful stimuli and no category-specific effects were observed for naming relative to reading. Nevertheless, category differences were found when naming and reading were considered together. Stimuli with greater visual complexity (animals, faces and maps) enhanced activation in the left extrastriate cortex. Furthermore, map recognition, which requires greater spatio-topographical processing, also activated the right occipito-parietal and parahippocampal cortices. These effects in the visuo-spatial regions emphasize inevitable differences in the perceptual properties of pictorial stimuli. In the semantic temporal regions, famous faces and objects enhanced activation in the left antero-lateral and postero-lateral cortices, respectively. In addition, we showed that the same posterior left temporal region is also activated by body parts. We conclude that category-specific brain activations depend more on differential processing at the perceptual and semantic levels rather than at the lexical retrieval level.     

Abnormal attentional modulation of retinotopic cortex in parietal patients with spatial neglect

Brain regions beyond visual cortex are thought to be responsible for attention-related modulation of visual processing [1, 2], but most evidence is indirect. Here, we applied functional magnetic resonance imaging (fMRI), including retinotopic mapping of visual areas, to patients with focal right-parietal lesions and left spatial neglect [3, 4]. When attentional load at fixation was minimal, retinotopic areas in right visual cortex showed preserved responses to task-irrelevant checkerboards in the contralateral left hemifield, analogously to left visual cortex for right-hemifield checkerboards, indicating a "symmetric" pattern in both hemispheres with respect to contralateral stimulation under these conditions. But when attentional load at fixation was increased, a functional asymmetry emerged for visual cortex, with contralateral responses in right visual areas being pathologically reduced (even eliminated for right V4/TEO), whereas left visual areas showed no such reduction in their contralateral response. These results reveal attention-dependent abnormalities in visual cortex after lesions in distant (parietal) regions. This may explain otherwise puzzling aspects of neglect [5, 6], as confirmed here by additional behavioral testing.     

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.     

Asymmetry of the cerebral hemispheres during comparison of paired visual stimuli

In conditions of tachistoscopic presentation of visual stimuli, healthy (male and female) right-handed subjects carried out a paired comparison of the stimuli presented unilaterally and in the center of the visual field. In case of recognition of images of words and objects, the number of correct answers and motor reaction time usually did not significantly differ at two interstimuli intervals (1 and 10 s). In comparing images of faces, there also were no differences by the number of reactions, and the reaction time was less at the intervals of 1 s. The left hemisphere dominated at the identification of words and female faces, the right one--at the recognition of male faces. When the right visual field was stimulated images of various classes were recognized more differentially than at the stimulation of the left visual field. The male subjects had more prominent interhemispheric differences than the females. The increase of the interstimuli interval from 1 to 10 s brought to a weakening of the functional interhemispheric asymmetry and decreasing of the differences between the male and female subjects. The obtained data show that in the processes connected with short-time memory, functional interhemispheric asymmetry is basically formed at the initial stages of the information processing.     

Hemispheric asymetry of the evoked electrical activity of the cerebral cortex to letter and non-verbal stimuli]

Average evoked potentials (AEP) were recorded in practically healthy subjects to "meaningless" figures and letters, presented to different halves of the visual field. Analysis of the amplitudes of AEP late components to verbal and non-verbal stimuli reveals hemispheric asymmetry. A higher amplitude of the late positive evoked response (P300) to a "direct" stimulation both by verbal and non-verbal stimuli (in the contralateral field of vision) is recorded in the left hemisphere than in the right one. Similar stimulation of the right hemisphere does not reveal sucha difference. In the left hemisphere the P300 wave is of a clearly greater amplitude to a "direct" stimulation (contralateral visual field) than to an "indirect" one (ipsilateral visual field), regardless of the nature of the stimulus. No such difference is observed in the right hemisphere. The magnitude of the late negative wave (component N200) to non-verbal stimuli is greater in the right hemisphere both in response to "direct" and "indirect" stimulations. No intrahemispheric difference has been found in the amplitude of late evoked responses of the cerebral cortex to verbal and non-verbal stimuli.     

Human parieto-occipital visual cortex: lack of retinotopy and foveal magnification.

We studied visual representation in the parietal cortex by recording whole-scalp neuromagnetic responses to luminance stimuli of varying eccentricities. The stimuli were semicircles (5.5 degrees in radius) presented at horizontal eccentricities from 0 degree to 16 degrees, separately in the right and left hemifields. All stimuli evoked responses in the contralateral occipital and medial parietal areas. The waveforms and distributions of the occipital responses varied with stimulus side (left, right) and eccentricity, whereas the parietal responses were remarkably similar to all stimuli. The equivalent sources of the parietal signals clustered within 1 cm3 in the medial parieto-occipital sulcus and did not differ significantly between the stimuli. The strength of the parietal activation remained practically constant with increasing stimulus eccentricity, suggesting that the visual areas in the parieto-occipital sulcus lack the enhanced foveal representation typical of most other visual areas. This result strengthens our previous suggestion that the medial parieto-occipital sulcus is the human homologue of the monkey V6 complex, characterized by, for example, lack of retinotopy and the absence of relative foveal magnification.     

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.     

Neocortical connectivity during episodic memory formation

During the formation of new episodic memories, a rich array of perceptual information is bound together for long-term storage. However, the brain mechanisms by which sensory representations (such as colors, objects, or individuals) are selected for episodic encoding are currently unknown. We describe a functional magnetic resonance imaging experiment in which participants encoded the association between two classes of visual stimuli that elicit selective responses in the extrastriate visual cortex (faces and houses). Using connectivity analyses, we show that correlation in the hemodynamic signal between face- and place-sensitive voxels and the left dorsolateral prefrontal cortex is a reliable predictor of successful face-house binding. These data support the view that during episodic encoding, "top-down" control signals originating in the prefrontal cortex help determine which perceptual information is fated to be bound into the new episodic memory trace.     

Early category-specific cortical activation revealed by visual stimulus inversion

Visual categorization may already start within the first 100-ms after stimulus onset, in contrast with the long-held view that during this early stage all complex stimuli are processed equally and that category-specific cortical activation occurs only at later stages. The neural basis of this proposed early stage of high-level analysis is however poorly understood. To address this question we used magnetoencephalography and anatomically-constrained distributed source modeling to monitor brain activity with millisecond-resolution while subjects performed an orientation task on the upright and upside-down presented images of three different stimulus categories: faces, houses and bodies. Significant inversion effects were found for all three stimulus categories between 70-100-ms after picture onset with a highly category-specific cortical distribution. Differential responses between upright and inverted faces were found in well-established face-selective areas of the inferior occipital cortex and right fusiform gyrus. In addition, early category-specific inversion effects were found well beyond visual areas. Our results provide the first direct evidence that category-specific processing in high-level category-sensitive cortical areas already takes place within the first 100-ms of visual processing, significantly earlier than previously thought, and suggests the existence of fast category-specific neocortical routes in the human brain.     

Model approach to neurological variants of visuo-spatial neglect

Neglect is a neurological disorder of spatial attention with reduced awareness of visual stimuli in the hemifield contralateral to an acute temporo-parietal lesion mainly of the right hemisphere. There is a close association of multisensory orientation centers (MSO) and vestibular tonus imbalance. A lesion of the dominant right MSO causes a left-sided neglect due to a lack of ipsilateral activation of the visual cortex, which is further enhanced by increasing inhibition from the contralateral visual cortex. The nondominant MSO in the left hemisphere might be involved in the manifestation of the less frequent and more transient right-sided neglect and in the plastic mechanisms of gradual recovery from left-sided neglect or extinction. There is evidence that a vestibular tonus inbalance due to peripheral or central vestibular pathway lesions may also induce a neglect. In a first model approach using an attractor network and assuming that there is only one MSO in the right hemisphere, it is possible to simulate attentional shifts into a visual hemifield and to induce a neglect. The neural network model consists of four layers of neurons: retina, MSO, visual cortex V1, and superior colliculus. The superior colliculus layer is modeled as a recurrent attractor network with one inhibitory interneuron and synaptic weights chosen to implement a winner-take-all network that centers the hill of activity on the strongest input. We are well aware of the simplifications used in the conceptual drawings and the computational model, but nevertheless hope that they will serve as an inspiration for further modeling and clinical studies.     

Imagine Jane and identify John: face identity aftereffects induced by imagined faces

It is not known whether prolonged exposure to perceived and imagined complex visual images produces similar shifts in subsequent perception through selective adaptation. This question is important because a positive finding would suggest that perception and imagery of visual stimuli are mediated by shared neural networks. In this study, we used a selective adaptation procedure designed to induce high-level face-identity aftereffects--a phenomenon in which extended exposure to a particular face facilitates recognition of subsequent faces with opposite features while impairing recognition of all other faces. We report here that adaptation to either real or imagined faces produces a similar shift in perception and that identity boundaries represented in real and imagined faces are equivalent. Together, our results show that identity information contained in imagined and real faces produce similar behavioral outcomes. Our findings of high-level visual aftereffects induced by imagined stimuli can be taken as evidence for the involvement of shared neural networks that mediate perception and imagery of complex visual stimuli.     

Neural Correlates of the Perception for Novel Objects

Perception of novel objects is of enormous importance in our lives. People have to perceive or understand novel objects when seeing an original painting, admiring an unconventional construction, and using an inventive device. However, very little is known about neural mechanisms underlying the perception for novel objects. Perception of novel objects relies on the integration of unusual features of novel objects in order to identify what such objects are. In the present study, functional Magnetic Resonance Imaging (MRI) was employed to investigate neural correlates of perception of novel objects. The neuroimaging data on participants engaged in novel object viewing versus ordinary object viewing revealed that perception of novel objects involves significant activation in the left precuneus (Brodmann area 7) and the right visual cortex. The results suggest that the left precuneus is associated with the integration of unusual features of novel objects, while the right visual cortex is sensitive to the detection of such features. Our findings highlight the left precuneus as a crucial component of the neural circuitry underlying perception of novel objects.     

Emotion Separation Is Completed Early and It Depends on Visual Field Presentation

It is now apparent that the visual system reacts to stimuli very fast, with many brain areas activated within 100 ms. It is, however, unclear how much detail is extracted about stimulus properties in the early stages of visual processing. Here, using magnetoencephalography we show that the visual system separates different facial expressions of emotion well within 100 ms after image onset, and that this separation is processed differently depending on where in the visual field the stimulus is presented. Seven right-handed males participated in a face affect recognition experiment in which they viewed happy, fearful and neutral faces. Blocks of images were shown either at the center or in one of the four quadrants of the visual field. For centrally presented faces, the emotions were separated fast, first in the right superior temporal sulcus (STS; 35–48 ms), followed by the right amygdala (57–64 ms) and medial pre-frontal cortex (83–96 ms). For faces presented in the periphery, the emotions were separated first in the ipsilateral amygdala and contralateral STS. We conclude that amygdala and STS likely play a different role in early visual processing, recruiting distinct neural networks for action: the amygdala alerts sub-cortical centers for appropriate autonomic system response for fight or flight decisions, while the STS facilitates more cognitive appraisal of situations and links appropriate cortical sites together. It is then likely that different problems may arise when either network fails to initiate or function properly.     

Sensory competition in the face processing areas of the human brain

The concurrent presentation of multiple stimuli in the visual field may trigger mutually suppressive interactions throughout the ventral visual stream. While several studies have been performed on sensory competition effects among non-face stimuli relatively little is known about the interactions in the human brain for multiple face stimuli. In the present study we analyzed the neuronal basis of sensory competition in an event-related functional magnetic resonance imaging (fMRI) study using multiple face stimuli. We varied the ratio of faces and phase-noise images within a composite display with a constant number of peripheral stimuli, thereby manipulating the competitive interactions between faces. For contralaterally presented stimuli we observed strong competition effects in the fusiform face area (FFA) bilaterally and in the right lateral occipital area (LOC), but not in the occipital face area (OFA), suggesting their different roles in sensory competition. When we increased the spatial distance among pairs of faces the magnitude of suppressive interactions was reduced in the FFA. Surprisingly, the magnitude of competition depended on the visual hemifield of the stimuli: ipsilateral stimulation reduced the competition effects somewhat in the right LOC while it increased them in the left LOC. This suggests a left hemifield dominance of sensory competition. Our results support the sensory competition theory in the processing of multiple faces and suggests that sensory competition occurs in several cortical areas in both cerebral hemispheres.     

Transcranial Direct Current Stimulation of the Dorsolateral Prefrontal Cortex Modulates Repetition Suppression to Unfamiliar Faces: An ERP Study

Repeated visual processing of an unfamiliar face suppresses neural activity in face-specific areas of the occipito-temporal cortex. This "repetition suppression" (RS) is a primitive mechanism involved in learning of unfamiliar faces, which can be detected through amplitude reduction of the N170 event-related potential (ERP). The dorsolateral prefrontal cortex (DLPFC) exerts top-down influence on early visual processing. However, its contribution to N170 RS and learning of unfamiliar faces remains unclear. Transcranial direct current stimulation (tDCS) transiently increases or decreases cortical excitability, as a function of polarity. We hypothesized that DLPFC excitability modulation by tDCS would cause polarity-dependent modulations of N170 RS during encoding of unfamiliar faces. tDCS-induced N170 RS enhancement would improve long-term recognition reaction time (RT) and/or accuracy rates, whereas N170 RS impairment would compromise recognition ability. Participants underwent three tDCS conditions in random order at ∼72 hour intervals: right anodal/left cathodal, right cathodal/left anodal and sham. Immediately following tDCS conditions, an EEG was recorded during encoding of unfamiliar faces for assessment of P100 and N170 visual ERPs. The P3a component was analyzed to detect prefrontal function modulation. Recognition tasks were administered ∼72 hours following encoding. Results indicate the right anodal/left cathodal condition facilitated N170 RS and induced larger P3a amplitudes, leading to faster recognition RT. Conversely, the right cathodal/left anodal condition caused N170 amplitude and RTs to increase, and a delay in P3a latency. These data demonstrate that DLPFC excitability modulation can influence early visual encoding of unfamiliar faces, highlighting the importance of DLPFC in basic learning mechanisms.     

Hemispheric interaction in man during perception of visual stimuli]

Averaged evoked potentials (AEP) to verbal (letters) and nonverbal (random shapes) stimuli exposed in the left and right visual fields were registered in healthy subjects with normal vision. Analysis of the later AEP latencies pointed to asymmetry in the temporal parameters of the interhemispheric interaction. The late AEP latency is shorter in the right hemisphere than in the left hemisphere. The difference is more pronounced in responses to nonverbal stimuli. The earlier development of the evoked potential in the right hemisphere (or the later one in the left hemisphere) accounts for the interhemispheric difference in the temporal parameters of the late AEP components. Comparison of the latency of the component P300 to verbal and nonverbal stimuli presented in the ipsilateral or the contralateral visual fields reveals a transfer of the results of the cortical processing of visual information in the course of interhemispheric interaction.     

Behavioral and neural lateralization of vision in courtship singing of the zebra finch

Along with human speech and language processing, birdsong has been one of the best-characterized model systems for understanding the relationship of lateralization of brain function to behavior. Lateralization of song production has been extensively characterized, and lateralization of song perception has begun to be studied. Here we have begun to examine whether behavior and brain function are lateralized in relation to communicative aspects of singing, as well. In order to monitor central brain function, we assayed the levels of several activity dependent immediate early genes after directed courtship singing. Consistent with a lateralization of visual processing during communication, there were higher levels of expression of both egr-1 and c-fos in the left optic tectum after directed singing. Because input from the eyes to the brain is almost completely contralateral in birds, these results suggest that visual input from the right eye should be favored during normal singing to females. Consistent with this, we further found that males sang more when they could use only their right eye compared to when they could use only their left eye. Normal levels of singing, though, required free use of both eyes to view the female. These results suggest that there is a preference for visual processing by the right eye and left brain hemisphere during courtship singing. This may reflect a proposed specialization of the avian left hemisphere in sustaining attention on stimuli toward which a motor response is planned.     

Genetic variation in the serotonin transporter modulates neural system-wide response to fearful faces

A distributed, serotonergically innervated neural system comprising extrastriate cortex, amygdala and ventral prefrontal cortex is critical for identification of socially relevant emotive stimuli. The extent to which a genetic variation of serotonin transporter gene 5-HTTLPR impacts functional connectivity between the amygdala and the other components of this neural system remains little examined. In our study, neural activity was measured using event-related functional magnetic resonance imaging in 29 right-handed, white Caucasian healthy subjects as they viewed mild or prototypical fearful and neutral facial expressions. 5-HTTLPR genotype was classified as homozygous for the short allele ( S/S ), homozygous for the long allele ( L/L ) or heterozygous ( S/L ). S/S showed greater activity than L/L within right fusiform gyrus (FG) to prototypically fearful faces. To these fearful faces, S/S more than other genotype subgroups showed significantly greater positive functional connectivity between right amygdala and FG and between right FG and right ventrolateral prefrontal cortex (VLPFC). There was a positive association between measure of psychoticism and degree of functional connectivity between right FG and right VLPFC in response to prototypically fearful faces. Our data are the first to show that genotypic variation in 5-HTTLPR modulates both the amplitude within and the functional connectivity between different components of the visual object-processing neural system to emotionally salient stimuli. These effects may underlie the vulnerability to mood and anxiety disorders potentially triggered by socially salient, emotional cues in individuals with the S allele of 5-HTTLPR.