Camouflage patterning modulates neural signatures of attention and decision-making.

Many animals rely on visual camouflage to avoid detection and increase their chances of survival. Edge disruption is commonly seen in the natural world, with animals evolving high-contrast markings that are incongruent with their real body outline in order to avoid recognition. While many studies have investigated how camouflage properties influence viewer performance and eye movement in predation search tasks, researchers in the field have yet to consider how camouflage may directly modulate visual attention and object processing. To examine how disruptive coloration modulates attention, we use a visual object recognition model to quantify object saliency. We determine if object saliency is predictive of human behavioural performance and subjective certainty, as well as neural signatures of attention and decision-making. We show that increasing edge disruption not only reduces detection and identification performance but is also associated with a dampening of neurophysiological signatures of attentional filtering. Increased self-reported certainty regarding decisions corresponds with neurophysiological signatures of evidence accumulation and decision-making. In summary, we have demonstrated a potential mechanism by which edge disruption increases the evolutionary fitness of animals by reducing the brain's ability to distinguish signal from noise, and hence to detect and identify the camouflaged animal.

Distinct Neural Signatures of Outcome Monitoring After Selection and Execution Errors.

Losing a point in tennis could result from poor shot selection or faulty stroke execution. To explore how the brain responds to these different types of errors, we examined feedback-locked EEG activity while participants completed a modified version of a standard three-armed bandit probabilistic reward task. Our task framed unrewarded outcomes as the result of either errors of selection or errors of execution. We examined whether amplitude of a medial frontal negativity (the feedback-related negativity [FRN]) was sensitive to the different forms of error attribution. Consistent with previous reports, selection errors elicited a large FRN relative to rewards, and amplitude of this signal correlated with behavioral adjustment after these errors. A different pattern was observed in response to execution errors. These outcomes produced a larger FRN, a frontocentral attenuation in activity preceding this component, and a subsequent enhanced error positivity in parietal sites. Notably, the only correlations with behavioral adjustment were with the early frontocentral attenuation and amplitude of the parietal signal; FRN differences between execution errors and rewarded trials did not correlate with subsequent changes in behavior. Our findings highlight distinct neural correlates of selection and execution error processing, providing insight into how the brain responds to the different classes of error that determine future action.

Emotional content overrides spatial attention.

Spatial attention is our capacity to attend to or ignore particular regions of our spatial environment. However, some classes of stimuli may be able to override our efforts to ignore them. Here we assessed the relationship between involuntary attentional capture with emotional images and spatial attention at early stages of perceptual processing. Multiple scenes of unpleasant and neutral content were displayed in rapid serial visual presentation (RSVP) streams that elicited the steady-state visual evoked potential (SSVEP), a neural marker of selective attention at early visual areas. In a spatial cueing task, participants were cued to covertly attend to RSVP streams presented at 4 and 6 Hz presentation rates in the left and right visual hemifields. The task was to detect square targets occasionally displayed within the image streams, responding only to those appearing on the cued side. The RSVP streams were always neutral pictures in one visual hemifield but would unpredictably switch from neutral to aversive content in the other visual hemifield. We found that SSVEP amplitude was consistently modulated by a change in emotional valence of image streams, regardless of whether the change in content occurred in the attended or unattended spatial location, reflecting an automatic sensory amplification for affective stimuli. The present data provide further evidence in support that emotional images can attract visual processing resources independently of spatial attention allocation, and are consistent with sustained sensory facilitation of early visual areas through re-entrant feedback projections from higher-order cortical areas involved in the extraction of affective information.

Affective Bias without Hemispheric Competition: Evidence for Independent Processing Resources in Each Cortical Hemisphere.

We assessed the extent of neural competition for attentional processing resources in early visual cortex between foveally presented task stimuli and peripheral emotional distracter images. Task-relevant and distracting stimuli were shown in rapid serial visual presentation (RSVP) streams to elicit the steady-state visual evoked potential, which serves as an electrophysiological marker of attentional resource allocation in early visual cortex. A task-related RSVP stream of symbolic letters was presented centrally at 15 Hz while distracting RSVP streams were displayed at 4 or 6 Hz in the left and right visual hemifields. These image streams always had neutral content in one visual field and would unpredictably switch from neutral to unpleasant content in the opposite visual field. We found that the steady-state visual evoked potential amplitude was consistently modulated as a function of change in emotional valence in peripheral RSVPs, indicating sensory gain in response to distracting affective content. Importantly, the facilitated processing for emotional content shown in one visual hemifield was not paralleled by any perceptual costs in response to the task-related processing in the center or the neutral image stream in the other visual hemifield. Together, our data provide further evidence for sustained sensory facilitation in favor of emotional distracters. Furthermore, these results are in line with previous reports of a "different hemifield advantage" with low-level visual stimuli and are suggestive of independent processing resources in each cortical hemisphere that operate beyond low-level visual cues, that is, with complex images that impact early stages of visual processing via reentrant feedback loops from higher order processing areas.

Frontal theta brain activity varies as a function of surgical experience and task error.

OBJECTIVE: Investigations into surgical expertise have almost exclusively focused on overt behavioral characteristics with little consideration of the underlying neural processes. Recent advances in neuroimaging technologies, for example, wireless, wearable scalp-recorded electroencephalography (EEG), allow an insight into the neural processes governing performance. We used scalp-recorded EEG to examine whether surgical expertise and task performance could be differentiated according to an oscillatory brain activity signal known as frontal theta-a putative biomarker for cognitive control processes. DESIGN SETTING AND PARTICIPANTS: Behavioral and EEG data were acquired from dental surgery trainees with 1 year (n=25) and 4 years of experience (n=20) while they performed low and high difficulty drilling tasks on a virtual reality surgical simulator. EEG power in the 4-7 Hz range in frontal electrodes (indexing frontal theta) was examined as a function of experience, task difficulty and error rate. RESULTS: Frontal theta power was greater for novices relative to experts (p=0.001), but did not vary according to task difficulty (p=0.15) and there was no Experience × Difficulty interaction (p=0.87). Brain-behavior correlations revealed a significant negative relationship between frontal theta and error in the experienced group for the difficult task (r=-0.594, p=0.0058), but no such relationship emerged for novices. CONCLUSION: We find frontal theta power differentiates between surgical experiences but correlates only with error rates for experienced surgeons while performing difficult tasks. These results provide a novel perspective on the relationship between expertise and surgical performance.

Rapid sensory gain with emotional distracters precedes attentional deployment from a foreground task.

The steady-state visual evoked potential (SSVEP), an electrophysiological marker of attentional resource allocation, has recently been demonstrated to serve as a neural signature of emotional content extraction from a rapid serial visual presentation (RSVP). SSVEP amplitude was reduced for streams of emotional relative to neutral scenes passively viewed at 6 Hz (~167 ms per image), but it was enhanced for emotional relative to neutral scenes when viewed as 4 Hz RSVP (250 ms per image). Here, we investigated whether these seemingly contradictory observations may be related to different dynamics in the allocation of attentional resources as a consequence of stimulation frequency. To this end, we advanced our distraction paradigm by presenting a visual foreground task consisting of randomly moving squares flickering at 15 Hz superimposed on task-irrelevant RSVP streams shown at 6 or 4 Hz, which could unpredictably switch from neutral to unpleasant content during the trial or remained neutral. Critically, our findings demonstrate that affective distractors captured attentional resources more strongly than their neutral counterparts, irrespective of whether they were presented at 6 or 4 Hz rate. Moreover, the emotion-dependent attentional deployment from the foreground task was temporally preceded by sustained sensory facilitation in response to emotional background images. Together, present findings provide evidence for rapid sustained visual facilitation but a rather slow attentional bias in favor of emotional distractors in early visual areas.

Transcranial alternating current stimulation at 10 Hz modulates response bias in the Somatic Signal Detection Task.

Ongoing, pre-stimulus oscillatory activity in the 8-13 Hz alpha range has been shown to correlate with both true and false reports of peri-threshold somatosensory stimuli. However, to directly test the role of such oscillatory activity in behaviour, it is necessary to manipulate it. Transcranial alternating current stimulation (tACS) offers a method of directly manipulating oscillatory brain activity using a sinusoidal current passed to the scalp. We tested whether alpha tACS would change somatosensory sensitivity or response bias in a signal detection task in order to test whether alpha oscillations have a causal role in behaviour. Active 10 Hz tACS or sham stimulation was applied using electrodes placed bilaterally at positions CP3 and CP4 of the 10-20 electrode placement system. Participants performed the Somatic Signal Detection Task (SSDT), in which they must detect brief somatosensory targets delivered at their detection threshold. These targets are sometimes accompanied by a light flash, which could also occur alone. Active tACS did not modulate sensitivity to targets but did modulate response criterion. Specifically, we found that active stimulation generally increased touch reporting rates, but particularly increased responding on light trials. Stimulation did not interact with the presence of touch, and thus increased both hits and false alarms. TACS stimulation increased reports of touch in a manner consistent with our observational reports, changing response bias, and consistent with a role for alpha activity in somatosensory detection.

Pre-stimulus alpha oscillations over somatosensory cortex predict tactile misperceptions.

Fluctuations of pre-stimulus oscillatory activity in the somatosensory alpha band (8-14Hz) observed using human EEG and MEG have been shown to influence the detection of supra- and peri-threshold somatosensory stimuli. However, some reports of touch occur even without a stimulus. We investigated the possibility that pre-stimulus alpha oscillations might also influence these false reports of touch - known as tactile misperceptions. We recorded EEG while participants performed the Somatic Signal Detection Task (SSDT), in which participants must detect brief, peri-threshold somatosensory targets. We found that pre-stimulus oscillatory power in the somatosensory alpha range exhibited a negative linear relationship with reporting of touch at electrode clusters over both contralateral and ipsilateral somatosensory regions. As pre-stimulus alpha power increased, the probability of reporting a touch declined; as it decreased, the probability of reporting a touch increased. This relationship was stronger on trials without a somatosensory stimulus than on trials with a somatosensory stimulus, although was present for both trial types. Spatio-temporal cluster-based permutation analysis also found that pre-stimulus alpha was lower on trials when touch was reported - irrespective of whether it was present - over contralateral and ipsilateral somatosensory cortices, as well as left frontocentral areas. We argue that alpha power may reflect changes in response criterion rather than sensitivity alone. Low alpha power relates to a low barrier to reporting a touch even when one is not present, while high alpha power is linked to less frequent reporting of touch overall.

Modulation of microsaccades by spatial frequency during object categorization.

The organization of visual processing into a coarse-to-fine information processing based on the spatial frequency properties of the input forms an important facet of the object recognition process. During visual object categorization tasks, microsaccades occur frequently. One potential functional role of these eye movements is to resolve high spatial frequency information. To assess this hypothesis, we examined the rate, amplitude and speed of microsaccades in an object categorization task in which participants viewed object and non-object images and classified them as showing either natural objects, man-made objects or non-objects. Images were presented unfiltered (broadband; BB) or filtered to contain only low (LSF) or high spatial frequency (HSF) information. This allowed us to examine whether microsaccades were modulated independently by the presence of a high-level feature - the presence of an object - and by low-level stimulus characteristics - spatial frequency. We found a bimodal distribution of saccades based on their amplitude, with a split between smaller and larger microsaccades at 0.4° of visual angle. The rate of larger saccades (⩾0.4°) was higher for objects than non-objects, and higher for objects with high spatial frequency content (HSF and BB objects) than for LSF objects. No effects were observed for smaller microsaccades (<0.4°). This is consistent with a role for larger microsaccades in resolving HSF information for object identification, and previous evidence that more microsaccades are directed towards informative image regions.

Accounting for microsaccadic artifacts in the EEG using independent component analysis and beamforming.

Neuronal activity in the gamma-band range was long considered a marker of object representation. However, scalp-recorded EEG activity in this range is contaminated by a miniature saccade-related muscle artifact. Independent component analysis (ICA) has been proposed as a method of removal of such artifacts. Alternatively, beamforming, a source analysis method in which potential sources of activity across the whole brain are scanned independently through the use of adaptive spatial filters, offers a promising method of accounting for the artifact without relying on its explicit removal. We present here the application of ICA-based correction to a previously published dataset. Then, using beamforming, we examine the effect of ICA correction on the scalp-recorded EEG signal and the extent to which genuine activity is recoverable before and after ICA correction. We find that beamforming attributes much of the scalp-recorded gamma-band signal before correction to deep frontal sources, likely the eye muscles, which generate the artifact related to each miniature saccade. Beamforming confirms that what is removed by ICA is predominantly this artifactual signal, and that what remains after correction plausibly originates in the visual cortex. Thus, beamforming allows researchers to confirm whether their removal procedures successfully removed the artifact. Our results demonstrate that ICA-based correction brings about general improvements in signal-to-noise ratio suggesting it should be used along with, rather than be replaced by, beamforming.

Early and late effects of objecthood and spatial frequency on event-related potentials and gamma band activity.

BACKGROUND: The visual system may process spatial frequency information in a low-to-high, coarse-to-fine sequence. In particular, low and high spatial frequency information may be processed via different pathways during object recognition, with LSF information projected rapidly to frontal areas and HSF processed later in visual ventral areas. In an electroencephalographic study, we examined the time course of information processing for images filtered to contain different ranges of spatial frequencies. Participants viewed either high spatial frequency (HSF), low spatial frequency (LSF), or unfiltered, broadband (BB) images of objects or non-object textures, classifying them as showing either man-made or natural objects, or non-objects. Event-related potentials (ERPs) and evoked and total gamma band activity (eGBA and tGBA) recorded using the electroencephalogram were compared for object and non-object images across the different spatial frequency ranges. RESULTS: The visual P1 showed independent modulations by object and spatial frequency, while for the N1 these factors interacted. The P1 showed more positive amplitudes for objects than non-objects, and more positive amplitudes for BB than for HSF images, which in turn evoked more positive amplitudes than LSF images. The peak-to-peak N1 showed that the N1 was much reduced for BB non-objects relative to all other images, while HSF and LSF non-objects still elicited as negative an N1 as objects. In contrast, eGBA was influenced by spatial frequency and not objecthood, while tGBA showed a stronger response to objects than non-objects. CONCLUSIONS: Different pathways are involved in the processing of low and high spatial frequencies during object recognition, as reflected in interactions between objecthood and spatial frequency in the visual N1 component. Total gamma band seems to be related to a late, probably high-level representational process.

Attentional bias to affective faces and complex IAPS images in early visual cortex follows emotional cue extraction.

Emotionally arousing stimuli are known to rapidly draw the brain's processing resources, even when they are task-irrelevant. The steady-state visual evoked potential (SSVEP) response, a neural response to a flickering stimulus which effectively allows measurement of the processing resources devoted to that stimulus, has been used to examine this process of attentional shifting. Previous studies have used a task in which participants detected periods of coherent motion in flickering random dot kinematograms (RDKs) which generate an SSVEP, and found that task-irrelevant emotional stimuli withdraw more attentional resources from the task-relevant RDKs than task-irrelevant neutral stimuli. However, it is not clear whether the emotion-related differences in the SSVEP response are conditional on higher-level extraction of emotional cues as indexed by well-known event-related potential (ERPs) components (N170, early posterior negativity, EPN), or if affective bias in competition for visual attention resources is a consequence of a time-invariant shifting process. In the present study, we used two different types of emotional distractors - IAPS pictures and facial expressions - for which emotional cue extraction occurs at different speeds, being typically earlier for faces (at ~170ms, as indexed by the N170) than for IAPS images (~220-280ms, EPN). We found that emotional modulation of attentional resources as measured by the SSVEP occurred earlier for faces (around 180ms) than for IAPS pictures (around 550ms), after the extraction of emotional cues as indexed by visual ERP components. This is consistent with emotion related re-allocation of attentional resources occurring after emotional cue extraction rather than being linked to a time-fixed shifting process.

Task and spatial frequency modulations of object processing: an EEG study.

Visual object processing may follow a coarse-to-fine sequence imposed by fast processing of low spatial frequencies (LSF) and slow processing of high spatial frequencies (HSF). Objects can be categorized at varying levels of specificity: the superordinate (e.g. animal), the basic (e.g. dog), or the subordinate (e.g. Border Collie). We tested whether superordinate and more specific categorization depend on different spatial frequency ranges, and whether any such dependencies might be revealed by or influence signals recorded using EEG. We used event-related potentials (ERPs) and time-frequency (TF) analysis to examine the time course of object processing while participants performed either a grammatical gender-classification task (which generally forces basic-level categorization) or a living/non-living judgement (superordinate categorization) on everyday, real-life objects. Objects were filtered to contain only HSF or LSF. We found a greater positivity and greater negativity for HSF than for LSF pictures in the P1 and N1 respectively, but no effects of task on either component. A later, fronto-central negativity (N350) was more negative in the gender-classification task than the superordinate categorization task, which may indicate that this component relates to semantic or syntactic processing. We found no significant effects of task or spatial frequency on evoked or total gamma band responses. Our results demonstrate early differences in processing of HSF and LSF content that were not modulated by categorization task, with later responses reflecting such higher-level cognitive factors.

Low-level and high-level modulations of fixational saccades and high frequency oscillatory brain activity in a visual object classification task.

Until recently induced gamma-band activity (GBA) was considered a neural marker of cortical object representation. However, induced GBA in the electroencephalogram (EEG) is susceptible to artifacts caused by miniature fixational saccades. Recent studies have demonstrated that fixational saccades also reflect high-level representational processes. Do high-level as opposed to low-level factors influence fixational saccades? What is the effect of these factors on artifact-free GBA? To investigate this, we conducted separate eye tracking and EEG experiments using identical designs. Participants classified line drawings as objects or non-objects. To introduce low-level differences, contours were defined along different directions in cardinal color space: S-cone-isolating, intermediate isoluminant, or a full-color stimulus, the latter containing an additional achromatic component. Prior to the classification task, object discrimination thresholds were measured and stimuli were scaled to matching suprathreshold levels for each participant. In both experiments, behavioral performance was best for full-color stimuli and worst for S-cone isolating stimuli. Saccade rates 200-700 ms after stimulus onset were modulated independently by low and high-level factors, being higher for full-color stimuli than for S-cone isolating stimuli and higher for objects. Low-amplitude evoked GBA and total GBA were observed in very few conditions, showing that paradigms with isoluminant stimuli may not be ideal for eliciting such responses. We conclude that cortical loops involved in the processing of objects are preferentially excited by stimuli that contain achromatic information. Their activation can lead to relatively early exploratory eye movements even for foveally-presented stimuli.

Time course of information processing in visual and haptic object classification.

Vision identifies objects rapidly and efficiently. In contrast, object recognition by touch is much slower. Furthermore, haptics usually serially accumulates information from different parts of objects, whereas vision typically processes object information in parallel. Is haptic object identification slower simply due to sequential information acquisition and the resulting memory load or due to more fundamental processing differences between the senses? To compare the time course of visual and haptic object recognition, we slowed visual processing using a novel, restricted viewing technique. In an electroencephalographic (EEG) experiment, participants discriminated familiar, nameable from unfamiliar, unnamable objects both visually and haptically. Analyses focused on the evoked and total fronto-central theta-band (5-7 Hz; a marker of working memory) and the occipital upper alpha-band (10-12 Hz; a marker of perceptual processing) locked to the onset of classification. Decreases in total upper alpha-band activity for haptic identification of objects indicate a likely processing role of multisensory extrastriate areas. Long-latency modulations of alpha-band activity differentiated between familiar and unfamiliar objects in haptics but not in vision. In contrast, theta-band activity showed a general increase over time for the slowed-down visual recognition task only. We conclude that haptic object recognition relies on common representations with vision but also that there are fundamental differences between the senses that do not merely arise from differences in their speed of processing.

An advantage for active versus passive aperture-viewing in visual object recognition.

In aperture viewing the field-of-view is restricted, such that only a small part of an image is visible, enforcing serial exploration of different regions of an object in order to successfully recognise it. Previous studies have used either active control or passive observation of the viewing aperture, but have not contrasted the two modes. Active viewing has previously been shown to confer an advantage in visual object recognition. We displayed objects through a small moveable aperture and tested whether people's ability to identify the images as familiar or novel objects was influenced by how the window location was controlled. Participants recognised objects faster when they actively controlled the window using their finger on a touch-screen, as opposed to passively observing the moving window. There was no difference between passively viewing again one's own window movement as generated in a previous block of trials versus viewing window movements that had been generated by other participants. These results contrast with those from comparable studies of haptic object recognition, which have found a benefit for passive over active stimulus exploration, but accord with findings of an advantage of active viewing in visual object recognition.

The effects of temporal delay and orientation on haptic object recognition.

We examined the effects of interstimulus interval (ISI) and orientation changes on the haptic recognition of novel objects, using a sequential shape-matching task. The stimuli consisted of 36 wedge-shaped plastic objects that varied along two shape dimensions (hole/bump and dip/ridge). Two objects were presented at either the same orientation or a different orientation, separated by either a short (3-sec) ISI or a long (15-sec) ISI. In separate conditions, ISI was blocked or randomly intermixed. Participants ignored orientation changes and matched on shape alone. Although performance was better in the mixed condition, there were no other differences between conditions. There was no decline in performance at the long ISI. There were similar, marginally significant benefits to same-orientation matching for short and long ISIs. The results suggest that the perceptual object representations activated from haptic inputs are both stable, being maintained for at least 15 sec, and orientation sensitive.

Size-sensitive perceptual representations underlie visual and haptic object recognition.

A variety of similarities between visual and haptic object recognition suggests that the two modalities may share common representations. However, it is unclear whether such common representations preserve low-level perceptual features or whether transfer between vision and haptics is mediated by high-level, abstract representations. Two experiments used a sequential shape-matching task to examine the effects of size changes on unimodal and crossmodal visual and haptic object recognition. Participants felt or saw 3D plastic models of familiar objects. The two objects presented on a trial were either the same size or different sizes and were the same shape or different but similar shapes. Participants were told to ignore size changes and to match on shape alone. In Experiment 1, size changes on same-shape trials impaired performance similarly for both visual-to-visual and haptic-to-haptic shape matching. In Experiment 2, size changes impaired performance on both visual-to-haptic and haptic-to-visual shape matching and there was no interaction between the cost of size changes and direction of transfer. Together the unimodal and crossmodal matching results suggest that the same, size-specific perceptual representations underlie both visual and haptic object recognition, and indicate that crossmodal memory for objects must be at least partly based on common perceptual representations.

Do left and right matter for haptic recognition of familiar objects?

Two experiments were carried out to examine the effects of dominant right versus non-dominant left exploration hand and left versus right object orientation on haptic recognition of familiar objects. In experiment 1, participants named 48 familiar objects in two blocks. There was no dominant-hand advantage to naming objects haptically and there was no interaction between exploration hand and object orientation. Furthermore, priming of naming was not reduced by changes of either object orientation or exploration hand. To test whether these results were attributable to a failure to encode object orientation and exploration hand, experiment 2 replicated experiment 1 except that the unexpected task in the second block was to decide whether either exploration hand or object orientation had changed relative to the initial naming block. Performance on both tasks was above chance, demonstrating that this information had been encoded into long-term haptic representations following the initial block of naming. Thus when identifying familiar objects, the haptic processing system can achieve object constancy efficiently across hand changes and object-orientation changes, although this information is often stored even when it is task-irrelevant.