Inattentional aftereffects: The role of attention on the strength of the motion aftereffect.

The way that attention affects the processing of visual information is one of the most intriguing fields in the study of visual perception. One way to examine this interaction is by studying the way perceptual aftereffects are modulated by attention. In the present study, we have manipulated attention during adaptation to translational motion generated by coherently moving random dots, in order to investigate the effect of the distraction of attention on the strength of the peripheral dynamic motion aftereffect (MAE). A foveal rapid serial visual presentation task (RSVP) of varying difficulty was introduced during the adaptation period while the adaptation and test stimuli were presented peripherally. Furthermore, to examine the interaction between the physical characteristics of the stimulus and attention, we have manipulated the motion coherence level of the adaptation stimuli. Our results suggested that the removal of attention through an irrelevant task modulated the MAE's magnitude moderately and that such an effect depends on the stimulus strength. We also showed that the MAE still persists with subthreshold and unattended stimuli, suggesting that perhaps attention is not required for the complete development of the MAE.

Reading text works better than watching videos to improve acuity in a simulation of artificial vision.

Simulated artificial vision is used in visual prosthesis design to answer questions about device usability. We previously reported a striking increase in equivalent visual acuity with daily use of a simulation of artificial vision in an active task, reading sentences, that required high levels of subject engagement, but passive activities are more likely to dominate post-implant experience. Here, we investigated the longitudinal effects of a passive task, watching videos. Eight subjects used a simulation of a thalamic visual prosthesis with 1000 phosphenes to watch 23 episodes of classic American television in daily, 25-min sessions, for a period of 1 month with interspersed reading tests that quantified reading accuracy and reading speed. For reading accuracy, we found similar dynamics to the early part of the learning process in our previous report, here leading to an improvement in visual acuity of 0.15 ± 0.05 logMAR. For reading speed, however, no change was apparent by the end of training. We found that single reading sessions drove about twice the improvement in acuity of single video sessions despite being only half as long. We conclude that while passive viewing tasks may prove useful for post-implant rehabilitation, active tasks are likely to be preferable.

Invisible Adaptation: The Effect of Awareness on the Strength of the Motion Aftereffect.

The ability to process information despite the lack of perceptual awareness is one of the most fascinating aspects of the visual system. Such unconscious processing is often investigated using adaptation, where any presence of the former can be traced by its footprint on aftereffects following the latter. We have investigated the mechanisms of the motion aftereffect (MAE) using random dot displays of varying motion coherence as well as crowding to modulate both the physical as well as the perceptual strength of the adaptation stimulus. Perceptual strength was quantitatively measured as the performance in a forced-choice direction-discrimination task. A motion-nulling technique was used to quantitatively measure the strength of the MAE. We show that the strength of the dynamic MAE is independently influenced by both the physical stimulus strength as well as the subjective perceptual strength, with the effect of the former being more prominent than that of the latter. We further show that the MAE still persists under conditions of subthreshold perception. Our results suggest that perceptual awareness can influence the strength of visual processing, but the latter is not fully dependent on the former and can still take place at its partial or even total absence.

The Machine behind the Stage: A Neurobiological Approach toward Theoretical Issues of Sensory Perception.

The purpose of the present article is to try and give a brief, scientific perspective on several issues raised in the Philosophy of Perception literature. This perspective gives a central role to the brain mechanisms that underlie perception: a percept is something that emerges when the brain is activated in a certain way and thus all perceptual experiences (whether veridical, illusory, or hallucinatory) have a common cause behind them, namely a given brain-activation pattern. What distinguishes between different cases of perception is what has caused this activation pattern, i.e., something very separate and very different from the perceptual experience itself. It is argued that separating the perceptual event from its hypothetical content, a direct consequence of the way everyday language is structured, creates unnecessary ontological complications regarding the nature of the hypothetical 'object' of perception. A clear distinction between the physical properties of the real world on the one hand (e.g., wavelength reflectance), and the psychological properties of perceptual experiences on the other (e.g., color) is clearly made. Finally, although perception is a way of acquiring knowledge/information about the world, this acquisition should be considered as a cognitive process which is separate to and follows perception. Therefore, the latter should remain neutral with respect to the 'correctness' or 'truth' of the knowledge acquired.

The physiology and psychophysics of the color-form relationship: a review.

The relationship between color and form has been a long standing issue in visual science. A picture of functional segregation and topographic clustering emerges from anatomical and electrophysiological studies in animals, as well as by brain imaging studies in human. However, one of the many roles of chromatic information is to support form perception, and in some cases it can do so in a way superior to achromatic (luminance) information. This occurs both at an early, contour-detection stage, as well as in late, higher stages involving spatial integration and the perception of global shapes. Pure chromatic contrast can also support several visual illusions related to form-perception. On the other hand, form seems a necessary prerequisite for the computation and assignment of color across space, and there are several respects in which the color of an object can be influenced by its form. Evidently, color and form are mutually dependent. Electrophysiological studies have revealed neurons in the visual brain able to signal contours determined by pure chromatic contrast, the spatial tuning of which is similar to that of neurons carrying luminance information. It seems that, especially at an early stage, form is processed by several, independent systems that interact with each other, each one having different tuning characteristics in color space. At later processing stages, mechanisms able to combine information coming from different sources emerge. A clear interaction between color and form is manifested by the fact that color-form contingencies can be observed in various perceptual phenomena such as adaptation aftereffects and illusions. Such an interaction suggests a possible early binding between these two attributes, something that has been verified by both electrophysiological and fMRI studies.

Independent sources of anisotropy in visual orientation representation: a visual and a cognitive oblique effect.

The representation of visual orientation is more accurate for cardinal orientations compared to oblique, and this anisotropy has been hypothesized to reflect a low-level visual process (visual, "class 1" oblique effect). The reproduction of directional and orientation information also leads to a mean error away from cardinal orientations or directions. This anisotropy has been hypothesized to reflect a high-level cognitive process of space categorization (cognitive, "class 2," oblique effect). This space categorization process would be more prominent when the visual representation of orientation degrades such as in the case of working memory with increasing cognitive load, leading to increasing magnitude of the "class 2" oblique effect, while the "class 1" oblique effect would remain unchanged. Two experiments were performed in which an array of orientation stimuli (1-4 items) was presented and then subjects had to realign a probe stimulus within the previously presented array. In the first experiment, the delay between stimulus presentation and probe varied, while in the second experiment, the stimulus presentation time varied. The variable error was larger for oblique compared to cardinal orientations in both experiments reproducing the visual "class 1" oblique effect. The mean error also reproduced the tendency away from cardinal and toward the oblique orientations in both experiments (cognitive "class 2" oblique effect). The accuracy or the reproduced orientation degraded (increasing variable error) and the cognitive "class 2" oblique effect increased with increasing memory load (number of items) in both experiments and presentation time in the second experiment. In contrast, the visual "class 1" oblique effect was not significantly modulated by any one of these experimental factors. These results confirmed the theoretical predictions for the two anisotropies in visual orientation reproduction and provided support for models proposing the categorization of orientation in visual working memory.

Asynchrony in visual consciousness and the possible involvement of attention.

When subjects are asked to perceptually bind rapidly alternating color and motion stimuli, the pairings they report are different from the ones actually occurring in physical reality. A possible explanation for this misbinding is that the time necessary for perception is different for different visual attributes. Such an explanation is in logical harmony with the fact that the visual brain is characterized by different, functionally specialized systems, with different processing times for each; this type of organization naturally leads to different perceptual times for the corresponding attributes. In the present review, the experimental findings supporting perceptual asynchrony are presented, together with the original theoretical explanation behind the phenomenon and its implication for visual consciousness. Alternative theoretical views and additional experimental facts concerning perceptual misbinding are also reviewed, with a particular emphasis given to the role of attention. With few exceptions, most theories converge on the idea that the observed misbinding reflects a difference in perception times, which is in turn due to differences in neuronal processing times for different attributes within the brain. These processing time differences have been attributed to several different factors, attention included, with the possibility of co-existence between them.

Binocular rivalry alternations and their relation to visual adaptation.

When different stimuli are presented dichoptically, perception alternates between the two in a stochastic manner. After a long-lasting and rigorous debate, there is growing consensus that this phenomenon, known as binocular rivalry (BR), is the result of a dynamic competition occurring at multiple levels of the visual hierarchy. The role of low- and high-level adaptation mechanisms in controlling these perceptual alternations has been a key issue in the rivalry literature. Both types of adaptation are dispersed throughout the visual system and have an equally influential, or even causal, role in determining perception. Such an explanation of BR is also in accordance with the relationship between the latter and attention. However, an overall explanation of this intriguing perceptual phenomenon needs to also include noise as an equally fundamental process involved in the stochastic resonance of perceptual bistability.

The effect of language on visual contrast sensitivity.

Embodied cognition and perceptual symbol theories assume that higher cognition interacts with and is grounded in perception and action. Recent experiments have shown that language processing interacts with perceptual processing in various ways, indicating that linguistic representations have a strong perceptual character. In the present study, we have used signal detection theory to investigate whether the comprehension of written sentences, implying either horizontal or vertical orientation, could improve the participants' visual sensitivity for discriminating between horizontal or vertical square-wave gratings and noise. We tested this prediction by conducting one main and one follow-up experiment. Our results indicate that language can, indeed, affect perception at such a low level of the visual process and thus provide further support for the embodied theories of cognition.

The effects of categorical and linguistic adaptation on binocular rivalry initial dominance.

Binocular rivalry (BR) is a phenomenon in which visual perception alternates between two different monocular stimuli. There has been a long debate regarding its nature, with a special emphasis on whether low- or high-level mechanisms are involved. Prior adaptation to one of the two monocular stimuli is known to affect initial dominance in the subsequent dichoptic presentation. In the present work, we have used three different types of adaptation in order to investigate how each one affects initial dominance during BR. In the first adaptation type, adapting to a stimulus identical to the one used during rivalry has led to its consequent suppression, verifying previous findings. The binocular presentation which we have used excludes the possibility of eye-adaptation, suggesting that it is the specific stimulus that the brain adapts to. In the second adaptation type, we find suppression effects following adaptation to stimuli belonging to the same category (face or house) but are different from the specific ones used in the following BR presentation. In the final adaptation type, in which the words "face" or "house" are used as adaptors, no statistically significant effect was found. These results suggest that perceptual selection can be directly influenced by the prior presentation of visual stimuli different to the ones used during BR, and thus support a higher-level, cognitive influence on the latter.

Brain activation and the locus of visual awareness.

A major problem in visual neuroscience is to distinguish neuronal activity which is directly related to the conscious percept. The word "directly" is used here as opposed to an indirect relationship, as is for example the case with activity in the retina, which is produced by a stimulus in the outside world and will eventually lead to the perception of this stimulus. As for the word "related", it is used to mean activity which creates the perceptual experience or, even more extremely, activity that is the perceptual experience. The distinction between the two (is vs. creates) is not straightforward, although there might be some differences between them. Philosophers would argue that they have a different phenomenology, the percept existing only for the perceiving person, whereas the underlying neuronal activation exists for all to observe. One could go on to argue that it is actually not the neuronal activation that "everybody" observes, but each one observes his own percept of it, which is also unique and subjective. Still, the content of this percept and the one of the original stimulus are quite different. The purpose of the present review is not to dig deep into such philosophical issues, but rather to give an overview of neuroscientific approaches trying to locate the neural correlate of conscious perception.

Feature binding in the feedback layers of area V2.

The visual features of an object are processed by multiple, functionally specialized areas of cerebral cortex. When several objects are seen simultaneously, what mechanism preserves the association of features that belong to a single item? We address this question-known as the "binding problem"-by examining combinatorial feature selectivity of neurons in area V2. In recording from anesthetized macaques, we estimate that dual selectivity for chromatic and spatiotemporal attributes is 50% more common (27% vs. 18% sampling frequency) in superficial and deep layer neurons receiving feedback connections from higher areas, compared with layer 4-3 neurons relaying ascending signals. The operation of feedback pathways is thought to mediate attentional modulation of activity that may achieve binding through acting to select one single object for higher representation and filtering out competing objects. We propose that dual-selective neurons perform a "bridging" function, mediating the transfer of feedback-induced bias between feature dimensions. The bias can be propagated through V2 output neurons (of unitary selectivity) to higher levels of specialized processing and so promote selection of the target object's representation among multiple feature maps. The bridging function would thus act to unify the outcome of parallel, object-selective processes taking place along segregated visual pathways.

Motion processing, directional selectivity, and conscious visual perception in the human brain.

Several human and monkey studies have demonstrated a close relationship between motion perception and activation of area V5, leading to the general view that activity in this area correlates with the subjective experience of motion. In the present study, we investigate whether the responses of this area are still governed by the motion percept when the latter is in conflict with the reality of the physical visual stimulation. We simultaneously presented two different, specially designed random-dot kinematograms, one to each eye. These stimuli either both had a single direction of motion and worked in synergy, or had opposite motion directions and thus cancelled each other out perceptually. In this way, we were able to pit the visual stimulus (one vs. two stimulating directions) against the reported perception (directional motion vs. motion noise) of human volunteers during fMRI experiments. We found that a strong motion stimulus that is weakly perceived is more effective in activating V5 (as well as V3) than a weaker motion stimulus, which is nevertheless robustly perceived. Thus, contrary to the prevailing view of perception being the correlate of activity in higher visual areas, we show here that activity is instead dominated by the properties of the physical stimulus, raising the question of whether there is a subpopulation of cells in V5 whose activity is critical for generating the motion percept. In addition, our results provide the first robust evidence for the presence of directionally selective neuronal populations in human prestriate cortex.

fMRI and its interpretations: an illustration on directional selectivity in area V5/MT.

fMRI is a tool to study brain function noninvasively that can reliably identify sites of neural involvement for a given task. However, to what extent can fMRI signals be related to measures obtained in electrophysiology? Can the blood-oxygen-level-dependent signal be interpreted as spatially pooled spiking activity? Here we combine knowledge from neurovascular coupling, functional imaging and neurophysiology to discuss whether fMRI has succeeded in demonstrating one of the most established functional properties in the visual brain, namely directional selectivity in the motion-processing region V5/MT+. We also discuss differences of fMRI and electrophysiology in their sensitivity to distinct physiological processes. We conclude that fMRI constitutes a complement, not a poor-resolution substitute, to invasive techniques, and that it deserves interpretations that acknowledge its stand as a separate signal.

Effect of background colors on the tuning of color-selective cells in monkey area V4.

When objects are viewed in different illuminants, their color does not change or changes little in spite of significant changes in the wavelength composition of the light reflected from them. In previous studies, we have addressed the physiology underlying this color constancy by recording from cells in areas V1, V2, and V4 of the anesthetized monkey. Truly color-coded cells, ones that respond to a patch of a given color irrespective of the wavelength composition of the light reflected from it, were only found in area V4. In the present study, we have used a different approach to test the responses of V4 cells in both anesthetized and awake behaving monkeys. Stimuli of different colors, embedded within a Mondrian-type multicolored background, were used to identify the chromatic selectivity of neurons. The illumination of the background was then varied, and the tuning of V4 neurons was tested again for each background illumination. With anesthetized monkeys, the psychophysical effect of changing background illumination was inferred from our own experience, whereas in the awake behaving animal, it was directly reported by the monkey. We found that the majority of V4 neurons shifted their color-tuning profile with each change in the background illumination: each time the color of the background on the computer screen was changed so as to simulate a change in illumination, cells shifted their color-tuning function in the direction of the chromaticity component that had been increased. A similar shift was also observed in colored match-to-sample psychometric functions of both human and monkey. The shift in monkey psychometric functions was quantitatively equivalent to the shift in the responses of the corresponding population of cells. We conclude that neurons in area V4 exhibit the property of color constancy and that their response properties are thus able to reflect color perception.

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.