Learning to use an invisible visual signal for perception

How does the brain construct a percept from sensory signals? One approach to this fundamental question is to investigate perceptual learning as induced by exposure to statistical regularities in sensory signals [1-7]. Recent studies showed that exposure to novel correlations between sensory signals can cause a signal to have new perceptual effects [2, 3]. In those studies, however, the signals were clearly visible. The automaticity of the learning was therefore difficult to determine. Here we investigate whether learning of this sort, which causes new effects on appearance, can be low level and automatic by employing a visual signal whose perceptual consequences were made invisible-a vertical disparity gradient masked by other depth cues. This approach excluded high-level influences such as attention or consciousness. Our stimulus for probing perceptual appearance was a rotating cylinder. During exposure, we introduced a new contingency between the invisible signal and the rotation direction of the cylinder. When subsequently presenting an ambiguously rotating version of the cylinder, we found that the invisible signal influenced the perceived rotation direction. This demonstrates that perception can rapidly undergo "structure learning" by automatically picking up novel contingencies between sensory signals, thus automatically recruiting signals for novel uses during the construction of a percept.     

Interhemispheric switching mediates perceptual rivalry

BACKGROUND: Binocular rivalry refers to the alternating perceptual states that occur when the images seen by the two eyes are too different to be fused into a single percept. Logothetis and colleagues have challenged suggestions that this phenomenon occurs early in the visual pathway. They have shown that, in alert monkeys, neurons in the primary visual cortex continue to respond to their preferred stimulus despite the monkey reporting its absence. Moreover, they found that neural activity higher in the visual pathway is highly correlated with the monkey's reported percept. These and other findings suggest that the neural substrate of binocular rivalry must involve high levels, perhaps the same levels involved in reversible figure alternations. RESULTS: We present evidence that activation or disruption of a single hemisphere in human subjects affects the perceptual alternations of binocular rivalry. Unilateral caloric vestibular stimulation changed the ratio of time spent in each competing perceptual state. Transcranial magnetic stimulation applied to one hemisphere disrupted normal perceptual alternations when the stimulation was timed to occur at one phase of the perceptual switch, but not at the other. Furthermore, activation of a single hemisphere by caloric stimulation affected the perceptual alternations of a reversible figure, the Necker cube. CONCLUSIONS: Our findings suggest that interhemispheric switching mediates perceptual rivalry. Thus, competition for awareness in both binocular rivalry and reversible figures occurs between, rather than within, each hemisphere. This interhemispheric switch hypothesis has implications for understanding the neural mechanisms of conscious experience and also has clinical relevance as the rate of both types of perceptual rivalry is slow in bipolar disorder (manic depression).     

A 'sticky' interhemispheric switch in bipolar disorder?

Despite years of research into bipolar disorder (manic depression), its underlying pathophysiology remains elusive. It is widely acknowledged that the disorder is strongly heritable, but the genetics are complex with less than full concordance in monozygotic twins and at least four susceptibility loci identified. We propose that bipolar disorder is the result of a genetic propensity for slow interhemispheric switching mechanisms that become ''stuck'' in one or the other state. Because slow switches are also ''sticky'' when compared with fast switches, the clinical manifestations of bipolar disorder may be explained by hemispheric activation being ''stuck'' on the left (mania) or on the right (depression). Support for this ''sticky'' interhemispheric switching hypothesis stems from our recent observation that the rate of perceptual alternation in binocular rivalry is slow in euthymic subjects with bipolar disorder (n = 18, median = 0.27 Hz) compared with normal controls (n = 49, median = 0.60 Hz, p < 0.0005). We have presented evidence elsewhere that binocular rivalry is itself an interhemispheric switching phenomenon. The rivalry alternation rate (putative interhemispheric switch rate) is robust in a given individual, with a test-retest correlation of more than 0.8, making it suitable for genetic studies. The interhemispheric switch rate may provide a trait-dependent biological marker for bipolar disorder.     

Individual differences in perceptual switching rates; the role of occipital alpha and frontal theta band activity

Prolonged presentation of visually ambiguous figures leads to perceptual switching. Individual switching rates show great variability. The present study compares individuals with high versus low switching rates by investigating human scalp electroencephalogram and blink rates. Eight subjects viewed the Necker cube continuously and responded to perceptual switching by pressing a button. Frequent switchers showed characteristic occipital alpha and frontal theta band activity prior to a switch, whereas infrequent switchers did not. The alpha activity was specific to switching, the theta activity was generic to perceptual processing conditions. A negative correlation was observed between perceptual switching and blink rates. These results suggest that the ability to concentrate attentional effort on the task is responsible for the differences in perceptual switching rates     

Defining the V5/MT neuronal pool for perceptual decisions in a visual stereo-motion task

In the primate visual cortex, neurons signal differences in the appearance of objects with high precision. However, not all activated neurons contribute directly to perception. We defined the perceptual pool in extrastriate visual area V5/MT for a stereo-motion task, based on trial-by-trial co-variation between perceptual decisions and neuronal firing (choice probability (CP)). Macaque monkeys were trained to discriminate the direction of rotation of a cylinder, using the binocular depth between the moving dots that form its front and rear surfaces. We manipulated the activity of single neurons trial-to-trial by introducing task-irrelevant stimulus changes: dot motion in cylinders was aligned with neuronal preference on only half the trials, so that neurons were strongly activated with high firing rates on some trials and considerably less activated on others. We show that single neurons maintain high neurometric sensitivity for binocular depth in the face of substantial changes in firing rate. CP was correlated with neurometric sensitivity, not level of activation. In contrast, for individual neurons, the correlation between perceptual choice and neuronal activity may be fundamentally different when responding to different stimulus versions. Therefore, neuronal pools supporting sensory discrimination must be structured flexibly and independently for each stimulus configuration to be discriminated.This article is part of the themed issue ‘Vision in our three-dimensional world''.     

The Role of Task-Related Learned Representations in Explaining Asymmetries in Task Switching

Task switch costs often show an asymmetry, with switch costs being larger when switching from a difficult task to an easier task. This asymmetry has been explained by difficult tasks being represented more strongly and consequently requiring more inhibition prior to switching to the easier task. The present study shows that switch cost asymmetries observed in arithmetic tasks (addition vs. subtraction) do not depend on task difficulty: Switch costs of similar magnitudes were obtained when participants were presented with unsolvable pseudo-equations that did not differ in task difficulty. Further experiments showed that neither task switch costs nor switch cost asymmetries were due to perceptual factors (e.g., perceptual priming effects). These findings suggest that asymmetrical switch costs can be brought about by the association of some tasks with greater difficulty than others. Moreover, the finding that asymmetrical switch costs were observed (1) in the absence of a task switch proper and (2) without differences in task difficulty, suggests that present theories of task switch costs and switch cost asymmetries are in important ways incomplete and need to be modified.     

Fractal rotation isolates mechanisms for form-dependent motion in human vision

Here, we describe a motion stimulus in which the quality of rotation is fractal. This makes its motion unavailable to the translation-based motion analysis known to underlie much of our motion perception. In contrast, normal rotation can be extracted through the aggregation of the outputs of translational mechanisms. Neural adaptation of these translation-based motion mechanisms is thought to drive the motion after-effect, a phenomenon in which prolonged viewing of motion in one direction leads to a percept of motion in the opposite direction. We measured the motion after-effects induced in static and moving stimuli by fractal rotation. The after-effects found were an order of magnitude smaller than those elicited by normal rotation. Our findings suggest that the analysis of fractal rotation involves different neural processes than those for standard translational motion. Given that the percept of motion elicited by fractal rotation is a clear example of motion derived from form analysis, we propose that the extraction of fractal rotation may reflect the operation of a general mechanism for inferring motion from changes in form.     

The bacterial flagellar switch complex is getting more complex

The mechanism of function of the bacterial flagellar switch, which determines the direction of flagellar rotation and is essential for chemotaxis, has remained an enigma for many years. Here we show that the switch complex associates with the membrane-bound respiratory protein fumarate reductase (FRD). We provide evidence that FRD binds to preparations of isolated switch complexes, forms a 1:1 complex with the switch protein FliG, and that this interaction is required for both flagellar assembly and switching the direction of flagellar rotation. We further show that fumarate, known to be a clockwise/switch factor, affects the direction of flagellar rotation through FRD. These results not only uncover a new component important for switching and flagellar assembly, but they also reveal that FRD, an enzyme known to be primarily expressed and functional under anaerobic conditions in Escherichia coli, nonetheless, has important, unexpected functions under aerobic conditions.     

The relationship between visual working memory and attention: retention of precise colour information in the absence of effects on perceptual selection

We examined the conditions under which a feature value in visual working memory (VWM) recruits visual attention to matching stimuli. Previous work has suggested that VWM supports two qualitatively different states of representation: an active state that interacts with perceptual selection and a passive (or accessory) state that does not. An alternative hypothesis is that VWM supports a single form of representation, with the precision of feature memory controlling whether or not the representation interacts with perceptual selection. The results of three experiments supported the dual-state hypothesis. We established conditions under which participants retained a relatively precise representation of a parcticular colour. If the colour was immediately task relevant, it reliably recruited attention to matching stimuli. However, if the colour was not immediately task relevant, it failed to interact with perceptual selection. Feature maintenance in VWM is not necessarily equivalent with feature-based attentional selection.     

Multi-timescale perceptual history resolves visual ambiguity

When visual input is inconclusive, does previous experience aid the visual system in attaining an accurate perceptual interpretation? Prolonged viewing of a visually ambiguous stimulus causes perception to alternate between conflicting interpretations. When viewed intermittently, however, ambiguous stimuli tend to evoke the same percept on many consecutive presentations. This perceptual stabilization has been suggested to reflect persistence of the most recent percept throughout the blank that separates two presentations. Here we show that the memory trace that causes stabilization reflects not just the latest percept, but perception during a much longer period. That is, the choice between competing percepts at stimulus reappearance is determined by an elaborate history of prior perception. Specifically, we demonstrate a seconds-long influence of the latest percept, as well as a more persistent influence based on the relative proportion of dominance during a preceding period of at least one minute. In case short-term perceptual history and long-term perceptual history are opposed (because perception has recently switched after prolonged stabilization), the long-term influence recovers after the effect of the latest percept has worn off, indicating independence between time scales. We accommodate these results by adding two positive adaptation terms, one with a short time constant and one with a long time constant, to a standard model of perceptual switching.     

Rapid Perceptual Switching of a Reversible Biological Figure

Certain visual stimuli can give rise to contradictory perceptions. In this paper we examine the temporal dynamics of perceptual reversals experienced with biological motion, comparing these dynamics to those observed with other ambiguous structure from motion (SFM) stimuli. In our first experiment, naïve observers monitored perceptual alternations with an ambiguous rotating walker, a figure that randomly alternates between walking in clockwise (CW) and counter-clockwise (CCW) directions. While the number of reported reversals varied between observers, the observed dynamics (distribution of dominance durations, CW/CCW proportions) were comparable to those experienced with an ambiguous kinetic depth cylinder. In a second experiment, we compared reversal profiles with rotating and standard point-light walkers (i.e. non-rotating). Over multiple test repetitions, three out of four observers experienced consistently shorter mean percept durations with the rotating walker, suggesting that the added rotational component may speed up reversal rates with biomotion. For both stimuli, the drift in alternation rate across trial and across repetition was minimal. In our final experiment, we investigated whether reversals with the rotating walker and a non-biological object with similar global dimensions (rotating cuboid) occur at random phases of the rotation cycle. We found evidence that some observers experience peaks in the distribution of response locations that are relatively stable across sessions. Using control data, we discuss the role of eye movements in the development of these reversal patterns, and the related role of exogenous stimulus characteristics. In summary, we have demonstrated that the temporal dynamics of reversal with biological motion are similar to other forms of ambiguous SFM. We conclude that perceptual switching with biological motion is a robust bistable phenomenon.     

Schizotypal perceptual aberrations of time: correlation between score, behavior and brain activity

A fundamental trait of the human self is its continuum experience of space and time. Perceptual aberrations of this spatial and temporal continuity is a major characteristic of schizophrenia spectrum disturbances--including schizophrenia, schizotypal personality disorder and schizotypy. We have previously found the classical Perceptual Aberration Scale (PAS) scores, related to body and space, to be positively correlated with both behavior and temporo-parietal activation in healthy participants performing a task involving self-projection in space. However, not much is known about the relationship between temporal perceptual aberration, behavior and brain activity. To this aim, we composed a temporal Perceptual Aberration Scale (tPAS) similar to the traditional PAS. Testing on 170 participants suggested similar performance for PAS and tPAS. We then correlated tPAS and PAS scores to participants' performance and neural activity in a task of self-projection in time. tPAS scores correlated positively with reaction times across task conditions, as did PAS scores. Evoked potential mapping and electrical neuroimaging showed self-projection in time to recruit a network of brain regions at the left anterior temporal cortex, right temporo-parietal junction, and occipito-temporal cortex, and duration of activation in this network positively correlated with tPAS and PAS scores. These data demonstrate that schizotypal perceptual aberrations of both time and space, as reflected by tPAS and PAS scores, are positively correlated with performance and brain activation during self-projection in time in healthy individuals along the schizophrenia spectrum.     

Abacus Training Affects Math and Task Switching Abilities and Modulates Their Relationships in Chinese Children

Our previous work demonstrated that abacus-based mental calculation (AMC), a traditional Chinese calculation method, could help children improve their math abilities (e.g. basic arithmetical ability) and executive function (e.g. working memory). This study further examined the effects of long-term AMC training on math ability in visual-spatial domain and the task switching component of executive function. More importantly, this study investigated whether AMC training modulated the relationship between math abilities and task switching. The participants were seventy 7-year-old children who were randomly assigned into AMC and control groups at primary school entry. Children in AMC group received 2-hour AMC training every week since primary school entry. On the contrary, children in the control group had never received any AMC training. Math and task switching abilities were measured one year and three years respectively after AMC training began. The results showed that AMC children performed better than their peers on math abilities in arithmetical and visual-spatial domains. In addition, AMC group responded faster than control group in the switching task, while no group difference was found in switch cost. Most interestingly, group difference was present in the relationships between math abilities and switch cost. These results implied the effect of AMC training on math abilities as well as its relationship with executive function.     

Applying an Attentional Set to Perceived and Remembered Features

Previous research has examined our ability to attend selectively to particular features of perceptual objects, as well as our ability to switch from attending to one type of feature to another. This is usually done in the context of anticipatory attentional-set control, comparing the neural mechanisms involved as participants prepare to attend to the same stimulus feature as on the previous trial (“task-stay” trials) with those required as participants prepare to attend to a different stimulus feature to that previously attended (“task-switch” trials). We wanted to establish how participants maintain or switch attentional set retrospectively, as they attend to features of objects held in visual short-term memory (VSTM). We found that switching, relative to maintaining attentional set retrospectively, was associated with a performance cost, which can be reduced over time. This control process was mirrored by a large parietal and frontal amplitude difference in the event-related brain potentials (ERPs) and significant differences in global field power (GFP) between switch and stay trials. However, when taking into account the switch/stay GFP differences, thereby controlling for this difference in amplitude, we could not distinguish these trial types topographically. By contrast, we found clear topographic differences between preparing an anticipatory feature-based attentional set versus applying it retrospectively within VSTM. These complementary topographical and amplitude analyses suggested that anticipatory and retrospective set control recruited a qualitatively different configuration of underlying neural generators. In contrast, switch/stay differences were largely quantitative, with them differing primarily in terms of amplitude rather than topography.     

Beta,but Not Gamma,Band Oscillations Index Visual Form-Motion Integration

Electrophysiological oscillations in different frequency bands co-occur with perceptual, motor and cognitive processes but their function and respective contributions to these processes need further investigations. Here, we recorded MEG signals and seek for percept related modulations of alpha, beta and gamma band activity during a perceptual form/motion integration task. Participants reported their bound or unbound perception of ambiguously moving displays that could either be seen as a whole square-like shape moving along a Lissajou''s figure (bound percept) or as pairs of bars oscillating independently along cardinal axes (unbound percept). We found that beta (15–25 Hz), but not gamma (55–85 Hz) oscillations, index perceptual states at the individual and group level. The gamma band activity found in the occipital lobe, although significantly higher during visual stimulation than during base line, is similar in all perceptual states. Similarly, decreased alpha activity during visual stimulation is not different for the different percepts. Trial-by-trial classification of perceptual reports based on beta band oscillations was significant in most observers, further supporting the view that modulation of beta power reliably index perceptual integration of form/motion stimuli, even at the individual level.     

The switching dynamics of the bacterial flagellar motor

Many bacteria are propelled by flagellar motors that stochastically switch between the clockwise and counterclockwise rotation direction. Although the switching dynamics is one of their most important characteristics, the mechanisms that control it are poorly understood. We present a statistical–mechanical model of the bacterial flagellar motor. At its heart is the assumption that the rotor protein complex, which is connected to the flagellum, can exist in two conformational states and that switching between these states depends on the interactions with the stator proteins, which drive the rotor. This couples switching to rotation, making the switch sensitive to torque and speed. Another key element is that after a switch, it takes time for the load to build up, due to conformational transitions of the flagellum. This slow relaxation dynamics of the filament leads, in combination with the load dependence of the switching frequency, to a characteristic switching time, as recently observed. Hence, our model predicts that the switching dynamics is not only controlled by the chemotaxis‐signaling network, but also by mechanical feedback of the flagellum.     

Fumarate or a fumarate metabolite restores switching ability to rotating flagella of bacterial envelopes.

Flagella of cytoplasm-free envelopes of Escherichia coli or Salmonella typhimurium can rotate in either the counterclockwise or clockwise direction, but they never switch from one direction of rotation to another. Exogenous fumarate, in the intracellular presence of the chemotaxis protein CheY, restored switching ability to envelopes, with a concomitant increase in clockwise rotation. An increase in clockwise rotation was also observed after fumarate was added to partially lysed cells of E. coli, but the proportion of switching cells remained unchanged.     

Biased Recognition of Facial Affect in Patients with Major Depressive Disorder Reflects Clinical State

Cognitive theories of depression posit that perception is negatively biased in depressive disorder. Previous studies have provided empirical evidence for this notion, but left open the question whether the negative perceptual bias reflects a stable trait or the current depressive state. Here we investigated the stability of negatively biased perception over time. Emotion perception was examined in patients with major depressive disorder (MDD) and healthy control participants in two experiments. In the first experiment subjective biases in the recognition of facial emotional expressions were assessed. Participants were presented with faces that were morphed between sad and neutral and happy expressions and had to decide whether the face was sad or happy. The second experiment assessed automatic emotion processing by measuring the potency of emotional faces to gain access to awareness using interocular suppression. A follow-up investigation using the same tests was performed three months later. In the emotion recognition task, patients with major depression showed a shift in the criterion for the differentiation between sad and happy faces: In comparison to healthy controls, patients with MDD required a greater intensity of the happy expression to recognize a face as happy. After three months, this negative perceptual bias was reduced in comparison to the control group. The reduction in negative perceptual bias correlated with the reduction of depressive symptoms. In contrast to previous work, we found no evidence for preferential access to awareness of sad vs. happy faces. Taken together, our results indicate that MDD-related perceptual biases in emotion recognition reflect the current clinical state rather than a stable depressive trait.     

Opposite influence of perceptual memory on initial and prolonged perception of sensory ambiguity

Observers continually make unconscious inferences about the state of the world based on ambiguous sensory information. This process of perceptual decision-making may be optimized by learning from experience. We investigated the influence of previous perceptual experience on the interpretation of ambiguous visual information. Observers were pre-exposed to a perceptually stabilized sequence of an ambiguous structure-from-motion stimulus by means of intermittent presentation. At the subsequent re-appearance of the same ambiguous stimulus perception was initially biased toward the previously stabilized perceptual interpretation. However, prolonged viewing revealed a bias toward the alternative perceptual interpretation. The prevalence of the alternative percept during ongoing viewing was largely due to increased durations of this percept, as there was no reliable decrease in the durations of the pre-exposed percept. Moreover, the duration of the alternative percept was modulated by the specific characteristics of the pre-exposure, whereas the durations of the pre-exposed percept were not. The increase in duration of the alternative percept was larger when the pre-exposure had lasted longer and was larger after ambiguous pre-exposure than after unambiguous pre-exposure. Using a binocular rivalry stimulus we found analogous perceptual biases, while pre-exposure did not affect eye-bias. We conclude that previously perceived interpretations dominate at the onset of ambiguous sensory information, whereas alternative interpretations dominate prolonged viewing. Thus, at first instance ambiguous information seems to be judged using familiar percepts, while re-evaluation later on allows for alternative interpretations.