Decoding Remapped Spatial Information in the Peri-Saccadic Period.

It has been suggested that, prior to a saccade, visual neurons predictively respond to stimuli that will fall in their receptive fields after completion of the saccade. This saccadic remapping process is thought to compensate for the shift of the visual world across the retina caused by eye movements. To map the timing of this predictive process in the brain, we recorded neural activity using electroencephalography (EEG) during a saccade task. Human participants (male and female) made saccades between two fixation points while covertly attending to oriented gratings briefly presented at various locations on the screen. Data recorded during trials in which participants maintained fixation were used to train classifiers on stimuli in different positions. Subsequently, data collected during saccade trials were used to test for the presence of remapped stimulus information at the post-saccadic retinotopic location in the peri-saccadic period, providing unique insight into when remapped information becomes available. We found that the stimulus could be decoded at the remapped location ∼180 ms post-stimulus onset, but only when the stimulus was presented 100-200 ms before saccade onset. Within this range, we found that the timing of remapping was dictated by stimulus onset rather than saccade onset. We conclude that presenting the stimulus immediately before the saccade allows for optimal integration of the corollary discharge signal with the incoming peripheral visual information, resulting in a remapping of activation to the relevant post-saccadic retinotopic neurons.Significance Statement Each eye movement leads to a shift of the visual world across the retina, such that the visual input before and after the eye movement do not match. Despite this, we perceive the visual world as stable. A predictive mechanism known as saccadic remapping is thought to contribute to this stability. We use a saccade task with time-resolved EEG decoding to obtain a fine-grained analysis of the temporal dynamics of the saccadic remapping process. Probing different stimulus-saccade latencies and an array of stimulus locations, we identify when remapped information becomes available in the visual cortex. We describe a critical window in which feedforward visual information and the preparatory motor signals interact to allow for predictive remapping of a stimulus.

Neural mechanisms of visual motion extrapolation.

Because neural processing takes time, the brain only has delayed access to sensory information. When localising moving objects this is problematic, as an object will have moved on by the time its position has been determined. Here, we consider predictive motion extrapolation as a fundamental delay-compensation strategy. From a population-coding perspective, we outline how extrapolation can be achieved by a forwards shift in the population-level activity distribution. We identify general mechanisms underlying such shifts, involving various asymmetries which facilitate the targeted 'enhancement' and/or 'dampening' of population-level activity. We classify these on the basis of their potential implementation (intra- vs inter-regional processes) and consider specific examples in different visual regions. We consider how motion extrapolation can be achieved during inter-regional signaling, and how asymmetric connectivity patterns which support extrapolation can emerge spontaneously from local synaptic learning rules. Finally, we consider how more abstract 'model-based' predictive strategies might be implemented. Overall, we present an integrative framework for understanding how the brain determines the real-time position of moving objects, despite neural delays.

Spike-timing dependent plasticity partially compensates for neural delays in a multi-layered network of motion-sensitive neurons.

The ability of the brain to represent the external world in real-time is impacted by the fact that neural processing takes time. Because neural delays accumulate as information progresses through the visual system, representations encoded at each hierarchical level are based upon input that is progressively outdated with respect to the external world. This 'representational lag' is particularly relevant to the task of localizing a moving object-because the object's location changes with time, neural representations of its location potentially lag behind its true location. Converging evidence suggests that the brain has evolved mechanisms that allow it to compensate for its inherent delays by extrapolating the position of moving objects along their trajectory. We have previously shown how spike-timing dependent plasticity (STDP) can achieve motion extrapolation in a two-layer, feedforward network of velocity-tuned neurons, by shifting the receptive fields of second layer neurons in the opposite direction to a moving stimulus. The current study extends this work by implementing two important changes to the network to bring it more into line with biology: we expanded the network to multiple layers to reflect the depth of the visual hierarchy, and we implemented more realistic synaptic time-courses. We investigate the accumulation of STDP-driven receptive field shifts across several layers, observing a velocity-dependent reduction in representational lag. These results highlight the role of STDP, operating purely along the feedforward pathway, as a developmental strategy for delay compensation.

Exploring the extent to which shared mechanisms contribute to motion-position illusions.

Motion-position illusions (MPIs) are visual motion illusions in which motion signals bias the perceived position of an object. Due to phenomenological similarities between these illusions, previous research has assumed that some are caused by common mechanisms. However, this assumption has yet to be directly tested. This study investigates this assumption by exploiting between-participant variations in illusion magnitude. During two sessions, 106 participants viewed the flash-lag effect, luminance flash-lag effect, Fröhlich effect, flash-drag effect, flash-grab effect, motion-induced position shift, twinkle-goes effect, and the flash-jump effect. For each effect, the magnitude of the illusion was reliable within participants, strongly correlating between sessions. When the pairwise correlations of averaged illusions magnitudes were explored, two clusters of statistically significant positively correlated illusions were identified. The first cluster comprised the flash-grab effect, motion-induced position shift, and twinkle-goes effect. The second cluster comprised the Fröhlich and flash-drag effect. The fact that within each of these two clusters, individual differences in illusion magnitude were correlated suggests that these clusters may reflect shared underlying mechanisms. An exploratory factor analysis provided additional evidence that these correlated clusters shared an underlying factor, with each cluster loading onto their own factor. Overall, our results reveal that, contrary to the prevailing perspective in the literature, while some motion-position illusions share processes, most of these illusions are unlikely to reflect any shared processes, instead implicating unique mechanisms.

Same but different: The latency of a shared expectation signal interacts with stimulus attributes.

Predictive coding theories assert that perceptual inference is a hierarchical process of belief updating, wherein the onset of unexpected sensory data causes so-called prediction error responses that calibrate erroneous inferences. Given the functionally specialised organisation of visual cortex, it is assumed that prediction error propagation interacts with the specific visual attribute violating an expectation. We sought to test this within the temporal domain by applying time-resolved decoding methods to electroencephalography (EEG) data evoked by contextual trajectory violations of either brightness, size, or orientation within a bound stimulus. We found that following ∼170 ms post stimulus onset, responses to both size violations and orientation violations were decodable from physically identical control trials in which no attributes were violated. These two violation types were then directly compared, with attribute-specific signalling being decoded from 265 ms. Temporal generalisation suggested that this dissociation was driven by latency shifts in shared expectation signalling between the two conditions. Using a novel temporal bias method, we then found that this shared signalling occurred earlier for size violations than orientation violations. To our knowledge, we are among the first to decode expectation violations in humans using EEG and have demonstrated a temporal dissociation in attribute-specific expectancy violations.

Visual Information Is Predictively Encoded in Occipital Alpha/Low-Beta Oscillations.

Hierarchical predictive coding networks are a general model of sensory processing in the brain. Under neural delays, these networks have been suggested to naturally generate oscillatory activity in approximately the α frequency range (∼8-12 Hz). This suggests that α oscillations, a prominent feature of EEG recordings, may be a spectral "fingerprint" of predictive sensory processing. Here, we probed this possibility by investigating whether oscillations over the visual cortex predictively encode visual information. Specifically, we examined whether their power carries information about the position of a moving stimulus, in a temporally predictive fashion. In two experiments (N = 32, 18 female; N = 34, 17 female), participants viewed an apparent-motion stimulus moving along a circular path while EEG was recorded. To investigate the encoding of stimulus-position information, we developed a method of deriving probabilistic spatial maps from oscillatory power estimates. With this method, we demonstrate that it is possible to reconstruct the trajectory of a moving stimulus from α/low-ß oscillations, tracking its position even across unexpected motion reversals. We also show that future position representations are activated in the absence of direct visual input, demonstrating that temporally predictive mechanisms manifest in α/ß band oscillations. In a second experiment, we replicate these findings and show that the encoding of information in this range is not driven by visual entrainment. By demonstrating that occipital α/ß oscillations carry stimulus-related information, in a temporally predictive fashion, we provide empirical evidence of these rhythms as a spectral "fingerprint" of hierarchical predictive processing in the human visual system.SIGNIFICANCE STATEMENT "Hierarchical predictive coding" is a general model of sensory information processing in the brain. When in silico predictive coding models are constrained by neural transmission delays, their activity naturally oscillates in roughly the α range (∼8-12 Hz). Using time-resolved EEG decoding, we show that neural rhythms in this approximate range (α/low-ß) over the human visual cortex predictively encode the position of a moving stimulus. From the amplitude of these oscillations, we are able to reconstruct the stimulus' trajectory, revealing signatures of temporally predictive processing. This provides direct neural evidence linking occipital α/ß rhythms to predictive visual processing, supporting the emerging view of such oscillations as a potential spectral "fingerprint" of hierarchical predictive processing in the human visual system.

Position representations of moving objects align with real-time position in the early visual response.

When interacting with the dynamic world, the brain receives outdated sensory information, due to the time required for neural transmission and processing. In motion perception, the brain may overcome these fundamental delays through predictively encoding the position of moving objects using information from their past trajectories. In the present study, we evaluated this proposition using multivariate analysis of high temporal resolution electroencephalographic data. We tracked neural position representations of moving objects at different stages of visual processing, relative to the real-time position of the object. During early stimulus-evoked activity, position representations of moving objects were activated substantially earlier than the equivalent activity evoked by unpredictable flashes, aligning the earliest representations of moving stimuli with their real-time positions. These findings indicate that the predictability of straight trajectories enables full compensation for the neural delays accumulated early in stimulus processing, but that delays still accumulate across later stages of cortical processing.

The survival of animals depends on their ability to respond to different stimuli quickly and efficiently. From flies fluttering away when a swatter approaches, to deer running away at the sight of a lion to humans ducking to escape a looming punch, fast-paced reactions to harmful stimuli is what keep us (and other fauna) from getting injured or seriously maimed. This entire process is orchestrated by the nervous system, where cells called neurons carry signals from our senses to higher processing centres in the brain, allowing us to react appropriately. However, this relay process from the sensory organs to the brain accumulates delays: it takes time for signals to be transmitted from cell to cell, and also for the brain to process these signals. This means that the information received by our brains is usually outdated, which could lead to delayed responses. Experiments done in cats and monkeys have shown that the brain can compensate for these delays by predicting how objects might move in the immediate future, essentially extrapolating the trajectories of objects moving in a predictable manner. This might explain why rabbits run in an impulsive zigzag manner when trying to escape a predator: if they change direction often enough, the predator may not be able to predict where they are going next. Johnson et al. wanted to find out whether human brains can also compensate for delays in processing the movement of objects, and if so, at what point (early or late) in the processing pipeline the compensation occurs. To do this, they recorded the electrical activity of neurons using electroencephalography from volunteers who were presented with both static and moving stimuli. Electroencephalography or EEG records the average activity of neurons in a region of the brain over a period of time. The data showed that the volunteers' brains responded to moving stimuli significantly faster than to static stimuli in the same position on the screen, essentially being able to track the real-time position of the moving stimulus. Johnson et al. further analysed and compared the EEG recordings for moving versus static stimuli to demonstrate that compensation for processing delays occurred early on in the processing journey. Indeed, the compensation likely happens before the signal reaches a part of the brain called the visual cortex, which processes stimuli from sight. Any delays accrued beyond this point were not accommodated for. Johnson et al. clearly demonstrate that the human brain can work around its own shortcomings to allow us to perceive moving objects in real time. These findings start to explain, for example, how sportspersons are able to catch fast-moving balls and hit serves coming to them at speeds of approximately 200 kilometres per hour. The results also lay the foundation for studying processing delays in other senses, such as hearing and touch.

Blurred Lines: Memory, Perceptions, and Consciousness: Commentary on "Consciousness as a Memory System" by Budson et al (2022).

In the previous issue, Budson, Richman, and Kensinger (2022) put forth the intriguing proposal that consciousness may have evolved from the episodic memory system. In addition to providing a possible evolutionary trajectory for consciousness, I believe that viewing consciousness as an extension of memory in this way is particularly useful for understanding some of the puzzling temporal complexities that are inherent to consciousness. For example, due to neural transmission delays, our conscious experience must necessarily lag the outside world, which creates a paradox for both conscious perception (Do we see the past, rather than the present?) and action (How can we make rapid decisions if it takes so long to become conscious of something?). These paradoxes can be elegantly solved by treating consciousness as a memory system. Finally, the proposal put forth by Budson and colleagues (2022) aligns with the emerging perspective that consciousness, like memory, represents a narrative time line of events rather than any single instant. However, I believe that this conceptualization can be further extended to include not only the past, but also the future. In this way, consciousness can be provocatively viewed as the remembered past, present, and future.

Decoding continuous variables from event-related potential (ERP) data with linear support vector regression using the Decision Decoding Toolbox (DDTBOX).

Multivariate classification analysis for event-related potential (ERP) data is a powerful tool for predicting cognitive variables. However, classification is often restricted to categorical variables and under-utilises continuous data, such as response times, response force, or subjective ratings. An alternative approach is support vector regression (SVR), which uses single-trial data to predict continuous variables of interest. In this tutorial-style paper, we demonstrate how SVR is implemented in the Decision Decoding Toolbox (DDTBOX). To illustrate in more detail how results depend on specific toolbox settings and data features, we report results from two simulation studies resembling real EEG data, and one real ERP-data set, in which we predicted continuous variables across a range of analysis parameters. Across all studies, we demonstrate that SVR is effective for analysis windows ranging from 2 to 100 ms, and relatively unaffected by temporal averaging. Prediction is still successful when only a small number of channels encode true information, and the analysis is robust to temporal jittering of the relevant information in the signal. Our results show that SVR as implemented in DDTBOX can reliably predict continuous, more nuanced variables, which may not be well-captured by classification analysis. In sum, we demonstrate that linear SVR is a powerful tool for the investigation of single-trial EEG data in relation to continuous variables, and we provide practical guidance for users.

Perception in real-time: predicting the present, reconstructing the past.

We feel that we perceive events in the environment as they unfold in real-time. However, this intuitive view of perception is impossible to implement in the nervous system due to biological constraints such as neural transmission delays. I propose a new way of thinking about real-time perception: at any given moment, instead of representing a single timepoint, perceptual mechanisms represent an entire timeline. On this timeline, predictive mechanisms predict ahead to compensate for delays in incoming sensory input, and reconstruction mechanisms retroactively revise perception when those predictions do not come true. This proposal integrates and extends previous work to address a crucial gap in our understanding of a fundamental aspect of our everyday life: the experience of perceiving the present.

Motion extrapolation in the flash-lag effect depends on perceived, rather than physical speed.

In the flash-lag effect (FLE), a flash in spatiotemporal alignment with a moving object is misperceived as lagging behind the moving object. One proposed explanation for this illusion is based on predictive motion extrapolation of trajectories. In this interpretation, the diverging effects of velocity on the perceived position of the moving object suggest that FLE might be based on the neural representation of perceived, rather than physical, velocity. By contrast, alternative explanations based on differential latency or temporal averaging would predict that the FLE does not rely on such a representation of perceived velocity. Here we examined whether the FLE is sensitive to illusory changes in perceived speed that result in changes to perceived velocity, while physical speed is constant. The perceived speed of the moving object was manipulated using revolving wedge stimuli with variable pattern textures (Experiment 1) and luminance contrast (Experiment 2). The motion extrapolation interpretation would predict that the changes in FLE magnitude should correspond to the changes in the perceived speed of the moving object. In the current study, two experiments demonstrated that perceived speed and FLE magnitude increased in the dynamic pattern relative to the static pattern conditions, and that the same effect was found in the low contrast compared to the high contrast conditions. These results showed that manipulations of texture and contrast that are known to alter judgments of perceived speed also modulate perceived position. We interpret this as a consequence of motion extrapolation mechanisms and discuss possible explanations for why we observed no cross-effect correlation.

Decoding explicit and implicit representations of health and taste attributes of foods in the human brain.

Obesity has become a significant problem word-wide and is strongly linked to poor food choices. Even in healthy individuals, taste perceptions often drive dietary decisions more strongly than healthiness. This study tested whether health and taste representations can be directly decoded from brain activity, both when explicitly considered, and when implicitly processed for decision-making. We used multivariate support vector regression for event-related potentials (as measured by the electroencephalogram) to estimate a regression model predicting ratings of tastiness and healthiness for each participant, based on their neural activity occurring in the first second of food cue processing. In Experiment 1, 37 healthy participants viewed images of various foods and explicitly rated their tastiness and healthiness. In Experiment 2, 89 healthy participants completed a similar rating task, followed by an additional experimental phase, in which they indicated their desire to consume snack foods with no explicit instruction to consider tastiness or healthiness. In Experiment 1 both attributes could be decoded, with taste information being available earlier than health. In Experiment 2, both dimensions were also decodable, and their significant decoding preceded the decoding of decisions (i.e., desire to consume the food). However, in Experiment 2, health representations were decodable earlier than taste representations. These results suggest that health information is activated in the brain during the early stages of dietary decisions, which is promising for designing obesity interventions aimed at quickly activating health awareness.

Predictive Visual Motion Extrapolation Emerges Spontaneously and without Supervision at Each Layer of a Hierarchical Neural Network with Spike-Timing-Dependent Plasticity.

The fact that the transmission and processing of visual information in the brain takes time presents a problem for the accurate real-time localization of a moving object. One way this problem might be solved is extrapolation: using an object's past trajectory to predict its location in the present moment. Here, we investigate how a simulated in silico layered neural network might implement such extrapolation mechanisms, and how the necessary neural circuits might develop. We allowed an unsupervised hierarchical network of velocity-tuned neurons to learn its connectivity through spike-timing-dependent plasticity (STDP). We show that the temporal contingencies between the different neural populations that are activated by an object as it moves causes the receptive fields of higher-level neurons to shift in the direction opposite to their preferred direction of motion. The result is that neural populations spontaneously start to represent moving objects as being further along their trajectory than where they were physically detected. Because of the inherent delays of neural transmission, this effectively compensates for (part of) those delays by bringing the represented position of a moving object closer to its instantaneous position in the world. Finally, we show that this model accurately predicts the pattern of perceptual mislocalization that arises when human observers are required to localize a moving object relative to a flashed static object (the flash-lag effect; FLE).SIGNIFICANCE STATEMENT Our ability to track and respond to rapidly changing visual stimuli, such as a fast-moving tennis ball, indicates that the brain is capable of extrapolating the trajectory of a moving object to predict its current position, despite the delays that result from neural transmission. Here, we show how the neural circuits underlying this ability can be learned through spike-timing-dependent synaptic plasticity and that these circuits emerge spontaneously and without supervision. This demonstrates how the neural transmission delays can, in part, be compensated to implement the extrapolation mechanisms required to predict where a moving object is at the present moment.

The time-course of prediction formation and revision in human visual motion processing.

Establishing the real-time position of a moving object poses a challenge to the visual system due to neural processing delays. While sensory information is travelling through the visual hierarchy, the object continues moving and information about its position becomes outdated. By extrapolating the position of a moving object along its trajectory, predictive mechanisms might effectively decrease the processing time associated with these objects. Here, we use time-resolved decoding of electroencephalographic (EEG) data from an apparent motion paradigm to demonstrate the interaction of two separate predictive mechanisms. First, we reveal predictive latency advantages for position representations as soon as the second object in an apparent motion sequence - even before the stimulus contains any physical motion energy. This is consistent with the existence of omni-directional, within-layer waves of sub-threshold activity that bring neurons coding for adjacent positions closer to their firing threshold, thereby reducing the processing time of the second stimulus in one of those positions. Second, we show that an additional direction-specific latency advantage emerges from the third sequence position onward, once the direction of the apparent motion stimulus is uniquely determined. Because the receptive fields of early visual areas are too small to encompass sequential apparent motion positions (as evidenced by the lack of latency modulation for the second stimulus position), this latency advantage most likely arises from descending predictions from higher to lower visual areas through feedback connections. Finally, we reveal that the same predictive activation that facilitates the processing of the object in its expected position needs to be overcome when the object's trajectory unexpectedly reverses, causing an additional latency disadvantage for stimuli that violate predictions. Altogether, our results suggest that two complementary mechanisms interact to form and revise predictions in visual motion processing, modulating the latencies of neural position representations at different levels of visual processing.

Predictive activation of sensory representations as a source of evidence in perceptual decision-making.

Our brains can represent expected future states of our sensory environment. Recent work has shown that, when we expect a specific stimulus to appear at a specific time, we can predictively generate neural representations of that stimulus even before it is physically presented. These observations raise two exciting questions: Are pre-activated sensory representations used for perceptual decision-making? And, do we transiently perceive an expected stimulus that does not actually appear? To address these questions, we propose that pre-activated neural representations provide sensory evidence that is used for perceptual decision-making. This can be understood within the framework of the Diffusion Decision Model as an early accumulation of decision evidence in favour of the expected percept. Our proposal makes novel predictions relating to expectation effects on neural markers of decision evidence accumulation, and also provides an explanation for why we sometimes perceive stimuli that are expected, but do not appear.

Visual mismatch responses index surprise signalling but not expectation suppression.

The ability to distinguish between commonplace and unusual sensory events is critical for efficient learning and adaptive behaviour. This has been investigated using oddball designs in which sequences of often-appearing (i.e., expected) stimuli are interspersed with rare (i.e., surprising) deviants. Resulting differences in electrophysiological responses following surprising compared to expected stimuli are known as visual mismatch responses (VMRs). VMRs are thought to index co-occurring contributions of stimulus repetition effects, expectation suppression (that occurs when one's expectations are fulfilled), and expectation violation (i.e., surprise) responses; however, these different effects have been conflated in existing oddball designs. To better isolate and quantify effects of expectation suppression and surprise, we adapted an oddball design based on Fast Periodic Visual Stimulation (FPVS) that controls for stimulus repetition effects. We recorded electroencephalography (EEG) while participants (N = 48) viewed stimulation sequences in which a single face identity was periodically presented at 6 Hz. Critically, one of two different face identities (termed oddballs) appeared as every 7th image throughout the sequence. The presentation probabilities of each oddball image within a sequence varied between 10 and 90%, such that participants could form expectations about which oddball face identity was more likely to appear within each sequence. We also included 'expectation neutral' 50% probability sequences, whereby consistently biased expectations would not be formed for either oddball face identity. We found that VMRs indexed surprise responses, and effects of expectation suppression were absent. That is, ERPs were more negative-going at occipitoparietal electrodes for surprising compared to neutral oddballs, but did not differ between expected and neutral oddballs. Surprising oddball-evoked ERPs were also highly similar across the 10-40% appearance probability conditions. Our findings indicate that VMRs which are not accounted for by repetition effects are best described as an all-or-none surprise response, rather than a minimisation of prediction error responses associated with expectation suppression.

Motion extrapolation in the High-Phi illusion: Analogous but dissociable effects on perceived position and perceived motion.

A range of visual illusions, including the much-studied flash-lag effect, demonstrate that neural signals coding for motion and position interact in the visual system. One interpretation of these illusions is that they are the consequence of motion extrapolation mechanisms in the early visual system. Here, we study the recently reported High-Phi illusion to investigate whether it might be caused by the same underlying mechanisms. In the High-Phi illusion, a rotating texture is abruptly replaced by a new, uncorrelated texture. This leads to the percept of a large illusory jump, which can be forward or backward depending on the duration of the initial motion sequence (the inducer). To investigate whether this motion illusion also leads to illusions of perceived position, in three experiments we asked observers to localize briefly flashed targets presented concurrently with the new texture. Our results replicate the original finding of perceived forward and backward jumps, and reveal an illusion of perceived position. Like the observed effects on illusory motion, these position shifts could be forward or backward, depending on the duration of the inducer: brief inducers caused forward mislocalization, and longer inducers caused backward mislocalization. Additionally, we found that both jumps and mislocalizations scaled in magnitude with the speed of the inducer. Interestingly, forward position shifts were observed at shorter inducer durations than forward jumps. We interpret our results as an interaction of extrapolation and correction-for-extrapolation, and discuss possible mechanisms in the early visual system that might carry out these computations.

Decoding the Temporal Dynamics of Covert Spatial Attention Using Multivariate EEG Analysis: Contributions of Raw Amplitude and Alpha Power.

Attention can be oriented in space covertly without the need of eye movements. We used multivariate pattern classification analyses (MVPA) to investigate whether the time course of the deployment of covert spatial attention leading up to the observer's perceptual decision can be decoded from both EEG alpha power and raw activity traces. Decoding attention from these signals can help determine whether raw EEG signals and alpha power reflect the same or distinct features of attentional selection. Using a classical cueing task, we showed that the orientation of covert spatial attention can be decoded by both signals. However, raw activity and alpha power may reflect different features of spatial attention, with alpha power more associated with the orientation of covert attention in space and raw activity with the influence of attention on perceptual processes.

The SSVEP tracks attention, not consciousness, during perceptual filling-in.

Research on the neural basis of conscious perception has almost exclusively shown that becoming aware of a stimulus leads to increased neural responses. By designing a novel form of perceptual filling-in (PFI) overlaid with a dynamic texture display, we frequency-tagged multiple disappearing targets as well as their surroundings. We show that in a PFI paradigm, the disappearance of a stimulus and subjective invisibility is associated with increases in neural activity, as measured with steady-state visually evoked potentials (SSVEPs), in electroencephalography (EEG). We also find that this increase correlates with alpha-band activity, a well-established neural measure of attention. These findings cast doubt on the direct relationship previously reported between the strength of neural activity and conscious perception, at least when measured with current tools, such as the SSVEP. Instead, we conclude that SSVEP strength more closely measures changes in attention.

The role of hue in visual search for texture differences: Implications for camouflage design.

The purpose of camouflage is to be inconspicuous against a given background. Colour is an important component of camouflage, and the task of designing a single camouflage pattern for use against multiple different backgrounds is particularly challenging. As it is impossible to match the colour gamut of each background exactly, the question arises which colours from the different backgrounds should be incorporated in a camouflage pattern to achieve optimal concealment. Here, we used a visual search paradigm to address this question. Observers searched multi-coloured continuous textures for target regions defined by either the presence or absence of additional hues. Targets could be either a combination of five hues against a four-hued background ("patches"), or a combination of four hues against a five-hued background ("holes"). In Experiment 1, a search asymmetry was observed for the different targets, as observers were less accurate and slower at detecting holes than patches. Additionally, we observed a linear separability effect: search for a target was more difficult when the hue that defined the target was within the gamut of distractor colours (e.g. orange amongst reds and yellows). In Experiment 2, we further investigated "hole" targets designed for two different backgrounds and found that optimal concealment against both backgrounds was achieved by including intermediate colours that represented a compromise between the common colours and the unique colours of each background. The findings provide insights into how search asymmetries can be extended to complex texture properties and help inform the design process of camouflage for multiple backgrounds.