Noise in the brain: Transcranial random noise stimulation and perceptual noise act on a stochastic resonance-like mechanism.

Stochastic resonance (SR) is a phenomenon in which a certain amount of random noise added to a weak subthreshold stimulus can enhance signal detectability. It is unknown how external noise interacts with neural noise in producing an SR-like phenomenon and whether this interaction results in a modulation of either network efficiency or the efficiency of single neurons. Using random dot motion stimuli and noninvasive brain stimulation, we attempted to unveil the specific mechanism of action of the SR-like phenomenon in motion perception, if present. We aimed to determine whether signal integration efficiency changes with external noise (random dot numerosity) and how electrical transcranial random noise stimulation (tRNS) can affect the peak performance. The participants performed a coherent motion detection task in which the random dot numerosity varied, whereas the signal-to-noise ratio (SNR) remained constant. We applied placebo or tRNS with an amplitude of either 1 or 2 mA during task execution. We found peaks in participants' performance both in the case of placebo stimulation and in the case of 1-mA tRNS. In the latter case (i.e., with an additional noise source), the peak emerged at lower random dot numerosity levels than when no additional noise was added (placebo). No clear peak was observed with 2-mA tRNS. An equivalent noise (EN) analysis confirmed that SR arises from a modulation of the network efficiency underlying motion signal integration. These results indicate a joint contribution of external and neural noise (modulated by tRNS) in eliciting an SR-like phenomenon.

Probing the effect of the expected-speed violation illusion.

Motion perception is complex for the brain to process, involving interacting computations of distance, time, and speed. These computations can be biased by the context and the features of the perceived moving object, giving rise to several types of motion illusions. Recent research has shown that, in addition to object features and context, lifelong priors can bias attributes of perception. In the present work, we investigated if such long acquired expectations can bias speed perception. Using a two-interval forced-choice (2-IFC) task, we asked 160 participants in different experiments to judge which of two vehicles, one archetypically fast (e.g. a motorbike), and one comparatively slower (e.g. a bike), was faster. By varying the objective speeds of the two-vehicle types, and measuring the participants' point of subjective equality, we observed a consistent bias in participants' speed perception. Counterintuitively, in the first three experiments the speed of an archetypically slow vehicle had to be decreased relative to that of an archetypically fast vehicle, for the two to be judged as the same. Similarly, in the next three experiments, an archetypically fast vehicle's speed had to be increased relative to an archetypically slow vehicle's speed, for the two to be perceived as equal. Four additional control experiments replicated our results. We define this newly found bias as the expected-speed violation illusion (ESVI). We believe the ESVI as conceptually very similar to the size-weight illusion, and discuss it within the Bayesian framework of human perception.

Parietal tACS at beta frequency improves vision in a crowding regime.

Visual crowding is the inability to discriminate objects when presented with nearby flankers and sets a fundamental limit for conscious perception. Beta oscillations in the parietal cortex were found to be associated to crowding, with higher beta amplitude related to better crowding resilience. An open question is whether beta activity directly and selectively modulates crowding. We employed Transcranial Alternating Current Stimulation (tACS) in the beta band (18-Hz), in the alpha band (10-Hz) or in a sham regime, asking whether 18-Hz tACS would selectively improve the perception of crowded stimuli by increasing parietal beta activity. Resting-state electroencephalography (EEG) was measured before and after stimulation to test the influence of tACS on endogenous oscillations. Consistently with our predictions, we found that 18-Hz tACS, as compared to 10-Hz tACS and sham stimulation, reduced crowding. This improvement was found specifically in the contralateral visual hemifield and was accompanied by an increased amplitude of EEG beta oscillations, confirming an effect on endogenous brain rhythms. These results support a causal relationship between parietal beta oscillations and visual crowding and provide new insights into the precise oscillatory mechanisms involved in human vision.

Suppressive effects on motion discrimination induced by transient flankers are reduced by perceptual learning.

We investigated spatial suppression of a drifting Gabor target of 0.5 c/° induced by adjacent and iso-oriented stationary Gabors (flankers) whose spatial frequency differed by ±1 and ±2 octaves to that of the drifting target. Stimuli (target and flankers) were presented for 33 ms. Results showed greater spatial suppression when the spatial frequency of the stationary but transient flanking Gabors was either equal or 1-2 octaves lower than when it was 1-2 octaves higher than the target's spatial frequency. This asymmetry was evident only for the drifting target, but not for the stationary target. In addition, we investigated whether perceptual learning (PL) reduced the spatial suppression induced by the flankers. We found that PL increased contrast sensitivity for the target, but only when it was reduced by the lateral masking flankers, and its effect did not transfer to an isolated drifting target of equal or higher spatial frequency. These results suggest that PL selectively affects suppressive interactions rather than contrast gain. We suggest that the suppressive effect of low spatial frequency flankers and the lack of suppression with high spatial frequency flankers may reflect two complementary phenomena: camouflage by the transient flankers (i.e., context) and breaking of camouflage by form-motion segmentation. Camouflage may result because both target and flankers activate the motion (magnocellular) system. Breaking of camouflage instead may occur when target and flankers' spatial frequency are more suitable for quasi-independent activation of the form system (by the flankers) and the motion system (by the target).

The role of crowding in contextual influences on contour integration.

Dakin and Baruch (2009) investigated how context influences contour integration, specifically reporting that near-perpendicular surrounding-elements reduced the exposure-duration observers required to localize and determine the shape of contours (compared to performance with randomly oriented surrounds) while near-parallel surrounds increased this time. Here, we ask if this effect might be a manifestation of visual crowding (the disruptive influence of "visual clutter" on object recognition). We first report that the effect generalizes to simple contour-localization (without explicit shape-discrimination) and influences tolerance to orientation jitter in the same way it affects threshold exposure-duration. We next directly examined the role of crowding by quantifying observers' local uncertainty (about the orientation of the elements that comprised our contours), showing that this largely accounts for the effects of context on global contour integration. These findings support the idea that context influences contour integration at a predominantly local stage of processing and that the local effects of crowding eventually influence downstream stages in the cortical processing of visual form.

Vision recovery with perceptual learning and non-invasive brain stimulation: Experimental set-ups and recent results, a review of the literature.

BACKGROUND: Vision is the sense which we rely on the most to interact with the environment and its integrity is fundamental for the quality of our life. However, around the globe, more than 1 billion people are affected by debilitating vision deficits. Therefore, finding a way to treat (or mitigate) them successfully is necessary. OBJECTIVE: This narrative review aims to examine options for innovative treatment of visual disorders (retinitis pigmentosa, macular degeneration, optic neuropathy, refractory disorders, hemianopia, amblyopia), especially with Perceptual Learning (PL) and Electrical Stimulation (ES). METHODS: ES and PL can enhance visual abilities in clinical populations, inducing plastic changes. We describe the experimental set-ups and discuss the results of studies using ES or PL or their combination in order to suggest, based on literature, which treatment is the best option for each clinical condition. RESULTS: Positive results were obtained using ES and PL to enhance visual functions. For example, repetitive transorbital Alternating Current Stimulation (rtACS) appeared as the most effective treatment for pre-chiasmatic disorders such as optic neuropathy. A combination of transcranial Direct Current Stimulation (tDCS) and visual training seems helpful for people with hemianopia, while transcranial Random Noise Stimulation (tRNS) makes visual training more efficient in people with amblyopia and mild myopia. CONCLUSIONS: This narrative review highlights the effect of different ES montages and PL in the treatment of visual disorders. Furthermore, new options for treatment are suggested. It is noteworthy to mention that, in some cases, unclear results emerged and others need to be more deeply investigated.

Effects of aging on interference control in selective attention and working memory.

Working memory decay in advanced age has been attributed to a concurrent decrease in the ability to control interference. The present study contrasted a form of interference control in selective attention that acts upon the perception of external stimuli (access) with another form that operates on internal representations in working memory (deletion), in order to determine both of their effects on working memory efficiency in younger and older adults. Additionally, we compared memory performance under these access and deletion functions to performance in their respective control conditions. The results indicated that memory accuracy improved in both age groups from the access functions, but that only young adults benefited from the deletion functions. In addition, intrusion effects in the deletion condition were larger in older than in younger adults. The ability to control the irrelevant perception- and memory-elicited interference did not decline in general with advancing age; rather, the control mechanisms that operate on internal memory representations declined specifically.

Dyslexia and the magnocellular-parvocellular coactivaton hypothesis.

Previous studies showed that the lateral masking of a fast-moving low spatial frequency (SF) target was strong when exerted by static flankers of lower or equal to the target SF and absent when flankers' SF was higher than the target's one. These masking and unmasking effects have been interpreted as due to Magnocellular-Magnocellular (M-M) inhibition and Parvocellular-on-Magnocellular (P-M) disinhibitory coactivation, respectively. Based on the hypothesis that the balance between the two systems is perturbed in Developmental Dyslexia (DD), we asked whether dyslexic children (DDs) behaved differently than Typically Developing children (TDs) in conditions of lateral masking. DDs and TDs performed a motion discrimination task, of a .5c/deg Gabor target moving at 16 deg/sec, either isolated or flanked by static Gabors with a SF of .125, .5 or 2 c/deg (Experiment 1). As a control, they also performed a contrast detection task of a static target, either isolated or flanked (Experiment 2). DDs did not perform any different from TDs with either a static target or an isolated moving target of low spatial frequency, thus suggesting efficient feedforward Magnocellular (M) and Parvocellular (P) processing. Also, DDs showed similar contrast thresholds to TDs in the M-M inhibition condition. Conversely, DDs did not recover from lateral masking in the M-P coactivation condition. In addition, their performance in this condition negatively correlated with non-words accuracy, supporting the suggestion that an inefficient Magno-Parvo coactivation may possibly be associated to both higher visual suppression and reduced perceptual stability during reading.

Perceptual learning improves visual functions in patients with albinistic bilateral amblyopia: A pilot study.

BACKGROUND: Several visual functions are impaired in patients with oculocutaneous albinism (OCA) associated to albinistic bilateral amblyopia (ABA). OBJECTIVE: In this study, we aimed at exploring whether perceptual learning (PL) can improve visual functions in albinism. METHOD: Six patients and six normal sighted controls, were trained in a contrast detection task with lateral masking. Participants were asked to choose which of the two intervals contained a foveally presented low-contrast Gabor patch. Targets were presented between higher contrast collinear flankers with equal spatial frequency. When increasing target-to-flanker distance, lateral interactions effect normally switches from inhibition to facilitation, up to no effect. RESULTS: Our findings showed that before PL, only controls showed facilitation. After PL, results suggest that facilitatory lateral interactions are found both in controls as well as in albino patients. These results suggest that PL could induce higher processing efficiency at early cortical level. Moreover, PL positive effect seems to transfer to higher-level visual functions, but results were not very consistent among tasks (visual acuity, contrast sensitivity function, hyperacuity and foveal crowding). CONCLUSIONS: Although a small sample size was tested, our findings suggest a rehabilitative potential of PL in improving visual functions in albinism.

Improving motion detection via anodal transcranial direct current stimulation.

BACKGROUND: To study motion perception, a stimulus consisting of a field of small, moving dots is often used. Generally, some of the dots coherently move in the same direction (signal) while the rest move randomly (noise). A percept of global coherent motion (CM) results when many different local motion signals are combined. CM computation is a complex process that requires the integrity of the middle-temporal area (MT/V5) and there is evidence that increasing the number of dots presented in the stimulus makes such computation more efficient. OBJECTIVE: In this study, we explored whether anodal direct current stimulation (tDCS) over MT/V5 would increase individual performance in a CM task at a low signal-to-noise ratio (SNR, i.e. low percentage of coherent dots) and with a target consisting of a large number of moving dots (high dot numerosity, e.g. >250 dots) with respect to low dot numerosity (<60 dots), indicating that tDCS favour the integration of local motion signal into a single global percept (global motion). METHOD: Participants were asked to perform a CM detection task (two-interval forced-choice, 2IFC) while they received anodal, cathodal, or sham stimulation on three different days. RESULTS: Our findings showed no effect of cathodal tDCS with respect to the sham condition. Instead, anodal tDCS improves performance, but mostly when dot numerosity is high (>400 dots) to promote efficient global motion processing. CONCLUSIONS: The present study suggests that tDCS may be used under appropriate stimulus conditions (low SNR and high dot numerosity) to boost the global motion processing efficiency, and may be useful to empower clinical protocols to treat visual deficits.

The Effect of Alpha tACS on the Temporal Resolution of Visual Perception.

We experience the world around us as a smooth and continuous flow. However, there is growing evidence that the stream of sensory inputs is not elaborated in an analog way but is instead organized in discrete or quasi-discrete temporal processing windows. These discrete windows are suggested to depend on rhythmic neural activity in the alpha (and theta) frequency bands, which in turn reflect changes in neural activity within, and coupling between, cortical areas. In the present study, we investigated a possible causal link between oscillatory brain activity in the alpha range (8-12 Hz) and the temporal resolution of visual perception, which determines whether sequential stimuli are perceived as distinct entities or combined into a single representation. To this aim, we employed a two-flash fusion task while participants received focal transcranial alternating current stimulation (tACS) in extra-striate visual regions including V5/MT of the right hemisphere. Our findings show that 10-Hz tACS, as opposed to a placebo (sham tACS), reduces the temporal resolution of perception, inducing participants to integrate the two stimuli into a unique percept more often. This pattern was observed only in the contralateral visual hemifield, providing further support for a specific effect of alpha tACS. The present findings corroborate the idea of a causal link between temporal windows of integration/segregation and oscillatory alpha activity in V5/MT and extra-striate visual regions. They also stimulate future research on possible ways to shape the temporal resolution of human vision in an individualized manner.

The motion aftereffect reloaded.

The motion aftereffect is a robust illusion of visual motion resulting from exposure to a moving pattern. There is a widely accepted explanation of it in terms of changes in the response of cortical direction-selective neurons. Research has distinguished several variants of the effect. Converging recent evidence from different experimental techniques (psychophysics, single-unit recording, brain imaging, transcranial magnetic stimulation, visual evoked potentials and magnetoencephalography) reveals that adaptation is not confined to one or even two cortical areas, but occurs at multiple levels of processing involved in visual motion analysis. A tentative motion-processing framework is described, based on motion aftereffect research. Recent ideas on the function of adaptation see it as a form of gain control that maximises the efficiency of information transmission at multiple levels of the visual pathway.

Audiovisual bounce-inducing effect: attention alone does not explain why the discs are bouncing.

Two discs moving from opposite points in space, overlapping and stopping at the other disc's starting point, can be seen as either bouncing or streaming through each other. With silent displays, observers report the discs as streaming, whereas if a sound is played when the discs touch each other, observers report the discs as bouncing. The origin of the switch from streaming to bouncing response is not known yet. The sound either shifts perception toward that of an impact-elastic event (i.e., a bounce) or subtracts the attention that is necessary to perceive the discs as streaming. We used either impact-similar (abrupt amplitude attack, gradual decay) or impact-dissimilar sounds (gradual amplitude attack, abrupt decay) and found that the first sounds induce the bouncing response, whereas the latter, although as distracting as the first, render streaming and bouncing responses equally frequent at most. We interpret the audiovisual bouncing effect as resulting from attention subtraction, which raises the number of bounce responses in comparison with silent displays, and from perception, which further increments the number of bounce responses and turns the response into a strong bounce response.

Perceived shrinkage of motion paths.

We show that human observers strongly underestimate a linear or circular trajectory that a luminous spot follows in the dark. At slow speeds, observers are relatively accurate, but, as the speed increases, the size of the path is progressively underestimated, by up to 35%. The underestimation imposes little memory load and does not require tracking of the trajectory. Most importantly, we found that underestimation occurred only when successive motion vectors changed in direction. This suggests a perceptual rather than representational origin of the illusion, related to vector-sum integration over time of neural motion signals in different directions.

A new "tilt" illusion reveals the relation between border ownership and border binding.

The "association field" models of contour detection predict specific spatial conditions for linking or grouping neighboring elements into smooth contours. We previously suggested that the "association field" model may account for perceptual binding of near-collinear luminance edges of same contrast polarity and their consequent unification into a unique contrast border with illusory tilt. This approach is now developed into a new version of the tilt illusion, the seesaw illusion, in which the contrast border is perceived as inverting concave-convex illusory curvature when background luminance is inverted, indicating that contrast polarity must be incorporated into the notion of "association field" to account for the seesaw illusion. We found that although tile-edge segmentation into alternating black-white segments produces conflicting local tilts, the illusion remains, up to 16 arcmin edge distance. This occurs at extreme background luminance for long segments (where only congruent edge segments of higher contrast bind, the others being perceptually assimilated into the background surface) and, when segments are too short for their orientation to be detected, at all background luminance values except that equidistant from black and white stripes. Our findings provide further confirmation that these are striking border ownership phenomena, demonstrating that figure/ground organization precedes perceptual binding of edges through association fields.

Separate motion-detecting mechanisms for first- and second-order patterns revealed by rapid forms of visual motion priming and motion aftereffect.

Fast adaptation biases the perceived motion direction of a subsequently presented ambiguous test pattern (R. Kanai & F. A. Verstraten, 2005). Depending on both the duration of the adapting stimulus (ranging from tens to hundreds of milliseconds) and the duration of the adaptation-test blank interval, the perceived direction of an ambiguous test pattern can be biased towards the same or the opposite direction of the adaptation pattern, resulting in rapid forms of motion priming or motion aftereffect respectively. These findings were obtained employing drifting luminance gratings. Many studies have shown that first-order motion (luminance-defined) and second-order motion (contrast-defined) stimuli are processed by separate mechanisms. We assessed whether these effects also exist within the second-order motion domain. Results show that fast adaptation to second-order motion biases the perceived direction of a subsequently presented second-order ambiguous test pattern with similar time courses to that obtained for first-order motion. To assess whether a single mechanism could account for these results, we ran a cross-order adaptation condition. Results showed little or no transfer between the two motion cues and probes, suggesting a degree of separation between the neural substrates subserving fast adaptation of first- and second-order motion.

Contextual influences in the peripheral retina of patients with macular degeneration.

Macular degeneration (MD) is the leading cause of low vision in the elderly population worldwide. In case of complete bilateral loss of central vision, MD patients start to show a preferred retinal region for fixation (PRL). Previous literature has reported functional changes that are connected with the emergence of the PRL. In this paper, we question whether the PRL undergoes a use-dependent cortical reorganization that alters the range of spatial lateral interactions between low-level filters. We asked whether there is a modulation of the excitatory/inhibitory lateral interactions or whether contextual influences are well accounted for by the same law that describes the integration response in normal viewers. In a group of 13 MD patients and 7 age-matched controls, we probed contextual influences by measuring the contrast threshold for a vertical target Gabor, flanked by two collinear high-contrast Gabors. Contextual influences of the collinear flankers were indicated by the changes in contrast threshold obtained at different target-to-flanker distances (λs) relative to the baseline orthogonal condition. Results showed that MDs had higher thresholds in the baseline condition and functional impairment in the identification tasks. Moreover, at the shortest λ, we found facilitatory rather than inhibitory contextual influence. No difference was found between the PRL and a symmetrical retinal position (non-PRL). By pulling together data from MD and controls we showed that in the periphery this inversion occurs when the target threshold approach the flankers' contrast (about 1:3 ratio) and that for patients it does occur in both the PRL and a symmetrical retinal position (non-PRL). We conclude that contrary to previous interpretations, this modulation doesn't seem to reflect use-dependent cortical reorganization but rather, it might result from a reduction of contrast gain for the target that promotes target-flankers grouping.

Developmental dyslexia: A deficit in magnocellular-parvocellular co-activation, not simply in pure magnocellular activation.

The magnocellular deficit theory of dyslexia suggests a selective impairment in contrast detection of stimuli involving pure magnocellular response (e.g. Gabor patches of 0.5 c/deg, 30 Hz, low contrast). An alternative hypothesis is that, dyslexia may be associated with a reduction of typical facilitation that normal readers present for stimuli relying on low-level magno-parvo co-activation, relative to stimuli eliciting pure magno activation. According to this hypothesis, any advantage in contrast sensitivity, produced by either decreasing stimuli temporal frequency (from 30 to 10 Hz, Experiment 1) or using static stimuli of increasing spatial frequency (from 0.5 to 4 c/deg, Experiment 2), would be ascribed to the coexisting responses of the magnocellular and parvocellular systems. In the control group, this advantage in contrast sensitivity was found for a 0.5 c/deg Gabor (either static or flickering at 10 Hz) and for a static Gabor of 4 c/deg. In contrast to magnocellular deficit theory predictions, dyslexic individuals showed no deficit in the unmixed magnocellular response. However, they showed no advantage when the relative weight between magnocellular and parvocellular inputs was thrown off balance in favor of the latter. These results suggest that in order to interpret low-level visual deficits in dyslexia, it is worth considering that fast, feedforward low-frequency representations of spatial structures may result from the coexisting responses of two systems. Our results suggest that in dyslexia, the relative contribution of these two systems in visual processing is perturbed, and that this may have detrimental consequences in word processing, both within the parafovea and the fovea during fixation.

Testing the visual field of children and adults with Rarebit: The role of task repetition on sensitivity.

Rarebit is a simple and user-friendly perimetry that tests the visual field by using tiny supra-threshold dot stimuli. It appears to be especially useful for examining the visual field of children who are under 12 years of age. However, previous data showed that the number of errors was higher in children than adults. We ask whether the different number of errors in these two groups depended on task learning and whether it may be accounted for by sensitivity differences or a response bias. Thirty-one children between 9 and 12 years of age and thirty-nine adults were tested three times with Rarebit perimetry. A bias-free sensitivity index, d', rather than the simple hit rate, revealed a group difference that remained after extensive task repetition. Indeed, d' increased with task learning in a similar way in the two groups so that group difference remained after practice. The response bias differed in the two groups, being conservative in the older group (criterion C >0) and liberal in the younger (criterion C < 0). Both biases disappeared with task learning in the third session, suggesting that response bias cannot account for the group difference in sensitivity after practice. When bias-free measures of sensitivity are used and task learning effects are minimized, Rarebit perimetry may be a more valuable method than simple mean hit rate (MHR) to enlighten sensitivity differences in the visual field assessment within the pediatric population.

Priming of first- and second-order motion: mechanisms and neural substrates.

Priming for luminance-modulated (first-order) motion has been shown to rely on the functional integrity of visual area V5/MT [Campana, G., Cowey, A., & Walsh, V. (2002). Priming of motion direction and area V5/MT: A test of perceptual memory. Cerebral Cortex, 12, 663-669; Campana, G., Cowey, A., & Walsh, V. (2006). Visual area V5/MT remembers "what" but not "where". Cerebral Cortex, 16, 1766-1770]. The high retinotopical organization of this area would predict that direction priming is sensitive to spatial position. In order to test this hypothesis, and to see whether a similar priming mechanism also exists with second-order motion, we tested motion direction priming and its interaction with spatial position with both first- and second-order motion. Indeed, whereas a number of studies have pinpointed the specific mechanisms and neural substrates for these two kinds of motion perception that appear to be (partially) non-overlapping (i.e., Lu, Z. L., & Sperling, G. (2001). Three-systems theory of human visual motion perception: Review and update. Journal of the Optical Society of America A, 18, 2331-2370; Vaina, L. M., & Soloviev, S. (2004). First-order and second-order motion: Neurological evidence for neuroanatomically distinct systems. Progress in Brain Research, 144, 197-212), the mechanisms and neural substrates mediating implicit memory for first- and second-order motion are still unknown. Our results indicate that priming for motion direction occurs not only with first-order but also with second-order motion. Priming for motion direction is position-sensitive both with first- and second-order motion, suggesting for both processes a locus of representation where retinotopicity is still maintained, that is within the V5/MT complex but earlier than MST. Cross-order motion priming also exists but is not sensitive to spatial position, suggesting that the locus where processing of first- and second-order motion converge is situated in MST or beyond.