Interhemispheric connections shape subjective experience of bistable motion

The right and left visual hemifields are represented in different cerebral hemispheres and are bound together by connections through the corpus callosum. Much has been learned on the functions of these connections from split-brain patients [1-4], but little is known about their contribution to conscious visual perception in healthy humans. We used diffusion tensor imaging and functional magnetic resonance imaging to investigate which callosal connections contribute to the subjective experience of a visual motion stimulus that requires interhemispheric integration. The "motion quartet" is an ambiguous version of apparent motion that leads to perceptions of either horizontal or vertical motion [5]. Interestingly, observers are more likely to perceive vertical than horizontal motion when the stimulus is presented centrally in the visual field [6]. This asymmetry has been attributed to the fact that, with central fixation, perception of horizontal motion requires integration across hemispheres whereas perception of vertical motion requires only intrahemispheric processing [7]. We are able to show that the microstructure of individually tracked callosal segments connecting motion-sensitive areas of the human MT/V5 complex (hMT/V5+; [8]) can predict the conscious perception of observers. Neither connections between primary visual cortex (V1) nor other surrounding callosal regions exhibit a similar relationship.     

Primary visual cortex activity along the apparent-motion trace reflects illusory perception

The illusion of apparent motion can be induced when visual stimuli are successively presented at different locations. It has been shown in previous studies that motion-sensitive regions in extrastriate cortex are relevant for the processing of apparent motion, but it is unclear whether primary visual cortex (V1) is also involved in the representation of the illusory motion path. We investigated, in human subjects, apparent-motion-related activity in patches of V1 representing locations along the path of illusory stimulus motion using functional magnetic resonance imaging. Here we show that apparent motion caused a blood-oxygenation-level-dependent response along the V1 representations of the apparent-motion path, including regions that were not directly activated by the apparent-motion-inducing stimuli. This response was unaltered when participants had to perform an attention-demanding task that diverted their attention away from the stimulus. With a bistable motion quartet, we confirmed that the activity was related to the conscious perception of movement. Our data suggest that V1 is part of the network that represents the illusory path of apparent motion. The activation in V1 can be explained either by lateral interactions within V1 or by feedback mechanisms from higher visual areas, especially the motion-sensitive human MT/V5 complex.     

The responses to illusory contours of neurons in cortex areas 17 and 18 of the cats

Responses to illusory contours (ICs) were sampled from neurons in cortical areas 17 and 18 of the anesthetized cats. For ICs sensitive cells, the differences of receptive field properties were compared when ICs and real contour stimuli were applied. Two hundred orientation or direction selective cells were studied. We find that about 42 percent of these cells were the ICs sensitive cells. Although their orientation or direction tuning curves to ICs bar and real bars were similar, the response modes (especially latency and time course) were different. The cells' responses to ICs were independent of the spatial phases of sinusoidal gratings, which composed the ICs. The cells' optimal spatial frequency to composing gratings the ICs was much higher than the one to moving gratings. Therefore, these cells really responded to the ICs rather than the line ends of composing gratings. For some kinds of velocity-tuning cells, the optimal velocity to moving ICs bar was much lower than the optimal velocity to moving     

A brain-derived metric for preferred kinetic stimuli

We here address the question of whether there is any correlation between subjective preference for simple configurations within a specific visual domain such as motion and strength of activity in visual areas in which that domain is emphasized. We prepared several distinctive patterns of dots in motion with various characteristics and asked humans to rate them according to their preference, before and while scanning the activity in their brains with functional magnetic resonance imaging. For simplicity, we restricted ourselves to motion in the fronto-parallel plane. Moving patterns produced activity in areas V1, V2, the V3 complex (V3, V3A, V3B) and V5, but only in areas V5, V3A/B and parietal cortex did the preferred kinetic patterns produce stronger activity when compared with the non-preferred ones. In addition, preferred patterns produced activity within field A1 of medial orbito-frontal cortex (mOFC), which is not otherwise activated by kinetic stimuli. Hence, for these areas, stronger neural activity correlated with subjective preference. We conclude that configurations of kinetic stimuli that are subjectively preferred correlate with stronger activity within early visual areas and within mOFC. This opens up the possibility of more detailed studies to relate subjective preferences to strength of activity in early visual areas and to relate activity in them to areas whose activity correlates with the subjective experience of beauty.     

The responses to illusory contours of neurons in cortex areas 17 and 18 of the cats

Responses to illusory contours (ICs) were sampled from neurons in cortical areas 17 and 18 of the anesthetized cats. For ICs sensitive cells, the differences of receptive field properties were compared when ICs and real contour stimuli were applied. Two hundred orientation or direction selective cells were studied. We find that about 42 percent of these cells were the ICs sensitive cells. Although their orientation or direction tuning curves to ICs bar and real bars were similar, the response modes (especially latency and time course) were different. The cells’ responses to ICs were independent of the spatial phases of sinusoidal gratings, which composed the ICs. The cells’ optimal spatial frequency to composing gratings the ICs was much higher than the one to moving gratings. Therefore, these cells really responded to the ICs rather than the line ends of composing gratings. For some kinds of velocity-tuning cells, the optimal velocity to moving ICs bar was much lower than the optimal velocity to moving bars. The present results demonstrate that some cells in areas 17 and 18 of cats have the ability to respond to ICs and have different response properties of the receptive fields to ICs and luminance boundaries via different neural mechanisms.     

Preparatory effects of distractor suppression: evidence from visual cortex

Spatial selective attention is the mechanism that facilitates the selection of relevant information over irrelevant information in the visual field. The current study investigated whether foreknowledge of the presence or absence of distractors surrounding an impending target stimulus results in preparatory changes in visual cortex. We cued the location of the target and the presence or absence of distractors surrounding the target while changes in blood oxygen level dependent (BOLD) signals were measured. In line with prior work, we found that top-down spatial attention resulted in an increased contralateral BOLD response, evoked by the cue throughout early visual cortex (areas V1, V2 and V3). In addition, cues indicating distractor presence evoked a substantial increase in the magnitude of the BOLD signal in visual area V3, but not in V2 or V1. This study shows that prior knowledge concerning the presence of a distractor results in enhanced attentional modulation of visual cortex, in visual areas where neuronal receptive fields are large enough to encompass both targets and distractors. We interpret these findings as evidence that top-down attentional control processes include active preparatory suppression mechanisms for irrelevant, distracting information in the visual scene.     

Compression and reflection of visually evoked cortical waves

Neuronal interactions between primary and secondary visual cortical areas are important for visual processing, but the spatiotemporal patterns of the interaction are not well understood. We used voltage-sensitive dye imaging to visualize neuronal activity in rat visual cortex and found visually evoked waves propagating from V1 to other visual areas. A primary wave originated in the monocular area of V1 and was "compressed" when propagating to V2. A reflected wave initiated after compression and propagated backward into V1. The compression occurred at the V1/V2 border, and local GABAA inhibition is important for the compression. The compression/reflection pattern provides a two-phase modulation: V1 is first depolarized by the primary wave, and then V1 and V2 are simultaneously depolarized by the reflected and primary waves, respectively. The compression/reflection pattern only occurred for evoked waves and not for spontaneous waves, suggesting that it is organized by an internal mechanism associated with visual processing.     

Reciprocal influence of cortical zones in normal cats and after creation of an experimental pathologic process

Acute experiments were conducted on non-anesthetized lystenon-restricted cats. The evoked potentials method was applied to the study of interaction of various cortical areas of one hemisphere and interhemispheric relations using stimuli of different modalities. It was shown that the action of somato-sensory areas on the visual cortex responses in the intact brain were mainly of a releasing character; these effects disappear in the formation of an experimental pathological focus. On the basis of the data obtained a conclusion was drawn on a possible participation of functional reorganization of the interhemispheric relations in the compensatory process mechanisms.     

Contrast invariance in the human lateral occipital complex depends on attention

The human visual system has a remarkable ability to successfully operate under a variety of challenging viewing conditions. For example, our object-recognition capabilities are largely unaffected by low-contrast (e.g., foggy) environments. The basis for this ability appears to be reflected in the neural responses in higher cortical visual areas that have been characterized as being invariant to changes in luminance contrast: neurons in these areas respond nearly equally to low-contrast as compared to high-contrast stimuli. This response pattern is fundamentally different than that observed in earlier visual areas such as primary visual cortex (V1), which is highly dependent on contrast. How this invariance is achieved in higher visual areas is largely unknown. We hypothesized that directed spatial attention is an important prerequisite of the contrast-invariant responses in higher visual areas and tested this with functional MRI (fMRI) while subjects directed their attention either toward or away from contrast-varying shape stimuli. We found that in the lateral occipital complex (LOC), a visual area important for processing shape information, attention changes the form of the contrast response function (CRF). By directing attention away from the shape stimuli, the CRF in the LOC was similar to that measured in V1. We describe a number of mechanisms that could account for this important function of attention.     

Cortical visual areas in monkeys: location, topography, connections, columns, plasticity and cortical dynamics

The visual system is constantly challenged to organize the retinal pattern of stimulation into coherent percepts. This task is achieved by the cortical visual system, which is composed by topographically organized analytic areas and by synthetic areas of the temporal lobe that have more holistic processing. Additional visual areas of the parietal lobe are related to motion perception and visuomotor control. V1 and V2 represent the entire visual field. MT represents only the binocular field, and V4 only the central 30 degrees-40 degrees. The parietal areas represent more of the periphery. For any eccentricity, the receptive field grows at each step of processing, more at anterior areas in the temporal lobe. Minimal point image size increases towards the temporal lobe, but remains fairly constant toward the parietal lobe. Patterns of projection show asymmetries. Central V2 and V4 project mainly to the temporal lobe, while peripherals V2 (more than 30 degrees) and V4 (more than 10 degrees) also project to the parietal lobe. Visual information that arrives at V1 projects to V2, MT and PO, which then project to other areas. Local lateral propagation and recursive loops corroborate to perceptual completion and filling in. Priority connections to temporal, parietal and parieto-temporal cortices help construct crude early representations of objects, trajectories and movements.     

Auditory and Visual Interhemispheric Communication in Musicians and Non-Musicians

The corpus callosum (CC) is a brain structure composed of axon fibres linking the right and left hemispheres. Musical training is associated with larger midsagittal cross-sectional area of the CC, suggesting that interhemispheric communication may be faster in musicians. Here we compared interhemispheric transmission times (ITTs) for musicians and non-musicians. ITT was measured by comparing simple reaction times to stimuli presented to the same hemisphere that controlled a button-press response (uncrossed reaction time), or to the contralateral hemisphere (crossed reaction time). Both visual and auditory stimuli were tested. We predicted that the crossed-uncrossed difference (CUD) for musicians would be smaller than for non-musicians as a result of faster interhemispheric transfer times. We did not expect a difference in CUDs between the visual and auditory modalities for either musicians or non-musicians, as previous work indicates that interhemispheric transfer may happen through the genu of the CC, which contains motor fibres rather than sensory fibres. There were no significant differences in CUDs between musicians and non-musicians. However, auditory CUDs were significantly smaller than visual CUDs. Although this auditory-visual difference was larger in musicians than non-musicians, the interaction between modality and musical training was not significant. Therefore, although musical training does not significantly affect ITT, the crossing of auditory information between hemispheres appears to be faster than visual information, perhaps because subcortical pathways play a greater role for auditory interhemispheric transfer.     

Interhemispheric connections of ocular-dominance columns in cats with binocular vision impairment

We have investigated the interhemispheric connections of areas 17 and 18 in cats with impaired binocular vision (monocular deprivation, uni- and bilateral strabismus). Monosynaptic neuronal connections were studied using microionophoretic injections of horseradish peroxidase in the single cortical columns and analsys of spatial distribution of retrogradely labelled callosal cells was performed. In the cases of monocular deprivation and strabismus, the spatial asymmetry and eye-specificity of interhemispheric connections are retained. Quantitative changes of connections are more pronounced in strabismic cats. In cats with binocular vision impairments, as well as in control ones, the width of callosal-recipient zone is larger than of the callosal cells zone. This may indicate that interhemispheric connections are non-reciprocal in the areas of cortex that are more distant from the projection of vertical meridian of visual field. We expect that there should be morpho-functional in the cells that are providing connections in opposite directions.     

Correction of Distortion in Flattened Representations of the Cortical Surface Allows Prediction of V1-V3 Functional Organization from Anatomy

Several domains of neuroscience offer map-like models that link location on the cortical surface to properties of sensory representation. Within cortical visual areas V1, V2, and V3, algebraic transformations can relate position in the visual field to the retinotopic representation on the flattened cortical sheet. A limit to the practical application of this structure-function model is that the cortex, while topologically a two-dimensional surface, is curved. Flattening of the curved surface to a plane unavoidably introduces local geometric distortions that are not accounted for in idealized models. Here, we show that this limitation is overcome by correcting the geometric distortion induced by cortical flattening. We use a mass-spring-damper simulation to create a registration between functional MRI retinotopic mapping data of visual areas V1, V2, and V3 and an algebraic model of retinotopy. This registration is then applied to the flattened cortical surface anatomy to create an anatomical template that is linked to the algebraic retinotopic model. This registered cortical template can be used to accurately predict the location and retinotopic organization of these early visual areas from cortical anatomy alone. Moreover, we show that prediction accuracy remains when extrapolating beyond the range of data used to inform the model, indicating that the registration reflects the retinotopic organization of visual cortex. We provide code for the mass-spring-damper technique, which has general utility for the registration of cortical structure and function beyond the visual cortex.     

Non-Bayesian contour synthesis

Recent research has witnessed an explosive increase in models that treat percepts as optimal probabilistic inference. The ubiquity of partial camouflage and occlusion in natural scenes, and the demonstrated capacity of the visual system to synthesize coherent contours and surfaces from fragmented image data, has inspired numerous attempts to model visual interpolation processes as rational inference. Here, we report striking new forms of visual interpolation that generate highly improbable percepts. We present motion displays depicting simple occlusion sequences that elicit vivid percepts of illusory contours (ICs) in displays for which they play no necessary explanatory role. These ICs define a second, redundant occluding surface, even though all of the image data can be fully explained by an occluding surface that is clearly visible. The formation of ICs in these images therefore entails an extraordinarily improbable co-occurrence of two occluding surfaces that arise from the same local occlusion events. The perceived strength of the ICs depends on simple low-level image properties, which suggests that they emerge as the outputs of mechanisms that automatically synthesize contours from the pattern of occlusion and disocclusion of local contour segments. These percepts challenge attempts to model visual interpolation as a form of rational inference and suggest the need to consider a broader space of computational problems and/or implementation level constraints to understand their genesis.     

Implications on visual apperception: Energy, duration, structure and synchronization

Although primary visual cortex (V1 or striate) activity per se is not sufficient for visual apperception (normal conscious visual experiences and conscious functions such as detection, discrimination, and recognition), the same is also true for extrastriate visual areas (such as V2, V3, V4/V8/VO, V5/M5/MST, IT, and GF). In the lack of V1 area, visual signals can still reach several extrastriate parts but appear incapable of generating normal conscious visual experiences. It is scarcely emphasized in the scientific literature that conscious perceptions and representations must have also essential energetic conditions. These energetic conditions are achieved by spatiotemporal networks of dynamic mitochondrial distributions inside neurons. However, the highest density of neurons in neocortex (number of neurons per degree of visual angle) devoted to representing the visual field is found in retinotopic V1. It means that the highest mitochondrial (energetic) activity can be achieved in mitochondrial cytochrome oxidase-rich V1 areas. Thus, V1 bear the highest energy allocation for visual representation.In addition, the conscious perceptions also demand structural conditions, presence of adequate duration of information representation, and synchronized neural processes and/or ‘interactive hierarchical structuralism.’ For visual apperception, various visual areas are involved depending on context such as stimulus characteristics such as color, form/shape, motion, and other features. Here, we focus primarily on V1 where specific mitochondrial-rich retinotopic structures are found; we will concisely discuss V2 where smaller riches of these structures are found. We also point out that residual brain states are not fully reflected in active neural patterns after visual perception. Namely, after visual perception, subliminal residual states are not being reflected in passive neural recording techniques, but require active stimulation to be revealed.     

Do Mirror Glasses Have the Same Effect on Brain Activity as a Mirror Box? Evidence from a Functional Magnetic Resonance Imaging Study with Healthy Subjects

Since its original proposal, mirror therapy has been established as a successful neurorehabilitative intervention in several neurological disorders to recover motor function or to relieve pain. Mirror therapy seems to operate by reactivating the contralesional representation of the non-mirrored limb in primary motor- and somatosensory cortex. However, mirror boxes have some limitations which prompted the use of additional mirror visual feedback devices. The present study evaluated the utility of mirror glasses compared to a mirror box. We also tested the hypothesis that increased interhemispheric communication between the motor hand areas is the mechanism by which mirror visual feedback recruits the representation of the non-mirrored limb. Therefore, mirror illusion capacity and brain activations were measured in a within-subject design during both mirror visual feedback conditions in counterbalanced order with 20 healthy subjects inside a magnetic resonance imaging scanner. Furthermore, we analyzed task-dependent functional connectivity between motor hand representations using psychophysiological interaction analysis during both mirror tasks. Neither the subjective quality of mirror illusions nor the patterns of functional brain activation differed between the mirror tasks. The sensorimotor representation of the non-mirrored hand was recruited in both mirror tasks. However, a significant increase in interhemispheric connectivity between the hand areas was only observed in the mirror glasses condition, suggesting different mechanisms for the recruitment of the representation of the non-mirrored hand in the two mirror tasks. We conclude that the mirror glasses might be a promising alternative to the mirror box, as they induce similar patterns of brain activation. Moreover, the mirror glasses can be easy applied in therapy and research. We want to emphasize that the neuronal mechanisms for the recruitment of the affected limb representation might differ depending on conceptual differences between MVF devices. However, our findings need to be validated within specific patient groups.     

Shifting attention within memory representations involves early visual areas

Prior studies have shown that spatial attention modulates early visual cortex retinotopically, resulting in enhanced processing of external perceptual representations. However, it is not clear whether the same visual areas are modulated when attention is focused on, and shifted within a working memory representation. In the current fMRI study participants were asked to memorize an array containing four stimuli. After a delay, participants were presented with a verbal cue instructing them to actively maintain the location of one of the stimuli in working memory. Additionally, on a number of trials a second verbal cue instructed participants to switch attention to the location of another stimulus within the memorized representation. Results of the study showed that changes in the BOLD pattern closely followed the locus of attention within the working memory representation. A decrease in BOLD-activity (V1-V3) was observed at ROIs coding a memory location when participants switched away from this location, whereas an increase was observed when participants switched towards this location. Continuous increased activity was obtained at the memorized location when participants did not switch. This study shows that shifting attention within memory representations activates the earliest parts of visual cortex (including V1) in a retinotopic fashion. We conclude that even in the absence of visual stimulation, early visual areas support shifting of attention within memorized representations, similar to when attention is shifted in the outside world. The relationship between visual working memory and visual mental imagery is discussed in light of the current findings.     

Interrelations of the cerebral hemispheres in the cat in early stages of ontogeny

Multiple recording of transcallosal responses (TCRs) from different cortex areas has been carried out by means of acute experiments with immobilized and anesthetized kittens at the age of 1 to 30 days after birth. Homotopical TCRs in kittens at the age of 2-15 days appear earlier, are presented wider and reveal features of a greater maturity configuration and of amplitudinal-temporal parameters in association zone (parietal and sensorimotor) in comparison with projection zones (somatosensory, visual and auditory). Interhemispheric interrelations in association cortex of kittens are carried out not only by means of callosal but extracallosal system. In the course of animal developing in the parietal cortex the drain of the surface-positive oscillation moves from V to III layer and the drain of the surface-negative deviation remains at the level of II-III layers. The late component is registered up to the depth of III-IV layers, having the drain in I-II layers. In sensorimotor cortex the surface-negative oscillation has the drain in I-II layers, surface-positive--in III and V--VI layers. The interhemispheric asymmetry emerging from the moment of responses appearance is peculiar to TCRs of projection and association zones. In the first month of the postnatal development the asymmetry of positive and negative TCR oscillation amplitude has an individual character in sensorimotor cortex and a specific one--in parietal. The temporal parameters of TCR in association areas of the left hemisphere cortex are significantly shorter than of the right one. The data given testify to the possibility of interhemispheric interrelation realization and the presence of interhemispheric asymmetry in cat's brain on the early stages of postnatal ontogenesis.     

Seeing invisible motion: a human FMRI study

A common view about visual consciousness is that it could arise when and where activity reaches some higher level of processing along the cortical hierarchy. Reports showing that activity in striate cortex can be dissociated from awareness , whereas the latter modulates activity in higher areas , point in this direction. In the specific case of visual motion, a central, "perceptual" role has been assigned to area V5: several human and monkey studies have shown V5 activity to correlate with the motion percept. Here we show that activity in this and other higher cortical areas can be also dissociated from perception and follow the physical stimulus instead. The motion information in a peripheral grating modulated fMRI responses, despite being invisible to human volunteers: under crowding conditions , areas V3A, V5, and parietal cortex still showed increased activity when the grating was moving compared to when it was flickering. We conclude that stimulus-specific activation of higher cortical areas does not necessarily result in awareness of the underlying stimulus.     

Responses to lightness variations in early human visual cortex

Lightness is the apparent reflectance of a surface, and it depends not only on the actual luminance of the surface but also on the context in which the surface is viewed [1-10]. The cortical mechanisms of lightness processing are largely unknown, and the role of early cortical areas is still a matter of debate [11-17]. We studied the cortical responses to lightness variations in early stages of the human visual system with functional magnetic resonance imaging (fMRI) while observers were performing a demanding fixation task. The set of dynamically presented visual stimuli included the rectangular version of the classic Craik-O'Brien stimulus [3, 18, 19] and a variant that led to a weaker lightness effect, as well as a pattern with actual luminance variations. We found that the cortical activity in retinotopic areas, including the primary visual cortex (V1), is correlated with context-dependent lightness variations.