Edge-enhanced disruptive camouflage impairs shape discrimination

Disruptive colouration (DC) is a form of camouflage comprised of areas of pigmentation across a target’s surface that form false edges, which are said to impede detection by disguising the outline of the target. In nature, many species with DC also exhibit edge enhancement (EE); light areas have lighter edges and dark areas have darker edges. EE DC has been shown to undermine not only localisation but also identification of targets, even when they are not hidden (Sharman, Moncrieff, & Lovell, 2018). We use a novel task, where participants judge which “snake” is more “wiggly,” to measure shape discrimination performance for three colourations (uniform, DC, and EE DC) and two backgrounds (leafy and uniform). We show that EE DC impairs shape discrimination even when targets are not hidden in a textured background. We suggest that this mechanism may contribute to misidentification of EE DC targets

Dissociating the effect of disruptive colouration on localisation and identification of camouflaged targets

Disruptive camouflage features contrasting areas of pigmentation across the animals’ surface that form false edges which disguise the shape of the body and impede detection. In many taxa these false edges feature local contrast enhancement or edge enhancement, light areas have lighter edges and dark areas have darker edges. This additional quality is often overlooked in existing research. Here we ask whether disruptive camouflage can have benefits above and beyond concealing location. Using a novel paradigm, we dissociate the time courses of localisation and identification of a target in a single experiment. We measured the display times required for a stimulus to be located or identified (the critical duration). Targets featured either uniform, disruptive or edge enhanced disruptive colouration. Critical durations were longer for identifying targets with edge enhanced disruptive colouration camouflage even when presented against a contrasting background, such that all target types were located equally quickly. For the first time, we establish empirically that disruptive camouflage not only conceals location, but also disguises identity. This shows that this form of camouflage can be useful even when animals are not hidden. Our findings offer insights into how edge enhanced disruptive colouration undermines visual perception by disrupting object recognition

Luminance cues constrain chromatic blur discrimination in natural scene stimuli

Introducing blur into the color components of a natural scene has very little effect on its percept, whereas blur introduced into the luminance component is very noticeable. Here we quantify the dominance of luminance information in blur detection and examine a number of potential causes. We show that the interaction between chromatic and luminance information is not explained by reduced acuity or spatial resolution limitations for chromatic cues, the effective contrast of the luminance cue, or chromatic and achromatic statistical regularities in the images. Regardless of the quality of chromatic information, the visual system gives primacy to luminance signals when determining edge location. In natural viewing, luminance information appears to be specialized for detecting object boundaries while chromatic information may be used to determine surface properties

The role of motion and number of element locations in mirror symmetry perception

The human visual system has specialised mechanisms for encoding mirror-symmetry and for detecting symmetric motion-directions for objects that loom or recede from the observers. The contribution of motion to mirror-symmetry perception has never been investigated. Here we examine symmetry detection thresholds for stationary (static and dynamic flicker) and symmetrically moving patterns (inwards, outwards, random directions) with and without positional symmetry. We also measured motion detection and direction-discrimination thresholds for horizontal (left, right) and symmetrically moving patterns with and without positional symmetry. We found that symmetry detection thresholds were (a) significantly higher for static patterns, but there was no difference between the dynamic flicker and symmetrical motion conditions, and (b) higher than motion detection and direction-discrimination thresholds for horizontal or symmetrical motion, with or without positional symmetry. In addition, symmetrical motion was as easy to detect or discriminate as horizontal motion. We conclude that whilst symmetrical motion per se does not contribute to symmetry perception, limiting the lifetime of pattern elements does improve performance by increasing the number of element-locations as elements move from one location to the next. This may be explained by a temporal integration process in which weak, noisy symmetry signals are combined to produce a stronger signal

Cue combination of colour and luminance in edge detection

Much is known about visual processing of chromatic and luminance information. However, less is known about how these two signals are combined. This thesis has three aims to investigate how colour and luminance are combined in edge detection. 1) To determine whether presenting colour and luminance information together improves performance in tasks such as edge localisation and blur detection. 2) To investigate how the visual system resolves conflicts between colour and luminance edge information. 3) To explore whether colour and luminance edge information is always combined in the same way. It is well known that the perception of chromatic blur can be constrained by sharp luminance information in natural scenes. The first set of experiments (Chapter 3) quantifies this effect and demonstrates that it cannot be explained by poorer acuity in processing chromatic information, higher contrast of luminance information or differences in the statistical structure of colour and luminance information in natural scenes. It is therefore proposed that there is a neural mechanism that actively promotes luminance information. Chapter 4 and Experiments 5.1 and 5.3 aimed to investigate whether the presence of both chromatic and luminance information improves edge localisation performance. Participant performance in a Vernier acuity (alignment) task was compared to predictions from three models; ‘winner takes all’, unweighted averaging and maximum likelihood estimation (a form of weighted averaging). Despite several attempts to differentiate the models we failed to increase the differences in model predictions sufficiently and it was not possible to determine whether edge localisation was enhanced by the presence of both cues. In Experiment 5.4 we investigated how edges are localised when colour and luminance cues conflict, using the method of adjustment. Maximum likelihood estimation was used to make predictions based on measurements of each cue in isolation. These predictions were then compared to observed data. It was found that, whilst maximum likelihood estimation captured the pattern of the data, it consistently over-estimated the weight of the luminance component. It is suggested that chromatic information may be weighted more heavily than predicted as it is more useful for detecting object boundaries in natural scenes. In Chapter 6 a novel approach, perturbation discrimination, was used to investigate how the spatial arrangement of chromatic and luminance cues, and the type of chromatic and luminance information, can affect cue combination. Perturbation discrimination requires participants to select the grating stimulus that contains spatial perturbation. If one cue dominated over the other it was expected that this would be reflected by masking and increased perturbation detection thresholds. We compared perturbation thresholds for chromatic and luminance defined line and square-wave gratings in isolation and when presented with a mask of the other channel and other grating type. For example, the perturbation threshold for a luminance line target alone was compared to the threshold for a luminance line target presented with a chromatic square-wave target. The introduction of line masks caused masking for both combinations. Introduction of an achromatic square-wave mask had no effect on perturbation thresholds for chromatic line targets. However, the introduction of a chromatic square-wave mask to luminance line targets improved perturbation discrimination performance. This suggests that the perceived location of the chromatic edges is determined by the location of the luminance lines. Finally, in Chapter 7, we investigated whether chromatic blur is constrained by luminance information in bipartite edges. Earlier in the thesis we demonstrated that luminance information constrains chromatic blur in natural scenes, but also that chromatic information has more influence than expected when colour and luminance edges conflict. This difference may be due to differences in the stimuli or due to differences in the task. The luminance masking effect found using natural scenes was replicated using bipartite edges. Therefore, the finding that luminance constrains chromatic blur is not limited to natural scene stimuli. This suggests that colour and luminance are combined differently for blur discrimination tasks and edge localisation tasks. Overall we can see that luminance often dominates in edge perception tasks. For blur discrimination this seems to be because the mechanisms differ. For edge localisation it might be simply that luminance cues are often higher contrast and, when this is equated, chromatic cues are actually a good indicator of edge location

The relationship between visual discomfort and cortical excitability in cone-opponent stimuli

Increased colour contrast can induce visual discomfort, but there is little research on the effect of hue. Colour is processed via one or more information streams or channels. We hypothesized that hues which activate more than one channel would induce greater visual discomfort, as they will demand greater neural resources. Normally-sighted young observers made discomfort judgments of isoluminant stimuli of varying hue and contrast whilst EEG was recorded. As predicted, stimuli recruiting more than one channel were more uncomfortable, and this increased with contrast. Uncomfortable stimuli showed increased N2 event-related potentials and decreased alpha-band oscillations, potentially indicating increased neural excitability. This is evidence that increased neural responses are related to visual discomfort for chromatic stimuli. Furthermore, it suggests that the origins of visual discomfort are in early visual areas, when colour is represented in a cone-opponent space, rather than later areas where colour representation is determined by perceptual similarity

Temporal dynamics of mirror-symmetry perception

Recent studies have suggested that temporal dynamics rather than symmetrical motion-direction contribute to mirror-symmetry perception. Here we investigate temporal aspects of symmetry perception and implicitly, its temporal flexibility and limitations, by examining how symmetrical pattern elements are combined over time. Stimuli were dynamic dot-patterns consisting of either an on-going alternation of two images (sustained stimulus presentation) or just two images each presented once (transient stimulus presentation) containing different amounts of symmetry about the vertical axis. We varied the presentation duration of the two images under five temporal-arrangement conditions: (1)‘whole patterns’ in which a symmetric pattern alternated with a noise pattern; (2)‘delayed halves’ – the halves of the symmetric and noise patterns were presented with temporal delay; (3)‘matched-pairs’ – two alternating images each containing equal amounts of symmetrical matched-pairs; (4)‘delayed matched-pairs’ – the same as arrangement 3, but with matched-pairs presented with delay; (5)‘static’ – both images presented simultaneously as one. We found increased sensitivity in sustained compared to transient stimulus presentations and with synchronous compared to delayed matched-pairs stimuli. For the delayed conditions, sensitivity decreased gradually with longer image durations (>60ms), prominently for the transient stimulus presentations. We conclude that correlations across-the-symmetry-midline can be integrated over time (~120ms) and symmetry mechanisms can tolerate temporal delays between symmetric dot-pairs of up to ~60ms

Cue combination of colour and luminance in edge detection

Much is known about visual processing of chromatic and luminance information. However, less is known about how these two signals are combined. This thesis has three aims to investigate how colour and luminance are combined in edge detection. 1) To determine whether presenting colour and luminance information together improves performance in tasks such as edge localisation and blur detection. 2) To investigate how the visual system resolves conflicts between colour and luminance edge information. 3) To explore whether colour and luminance edge information is always combined in the same way. It is well known that the perception of chromatic blur can be constrained by sharp luminance information in natural scenes. The first set of experiments (Chapter 3) quantifies this effect and demonstrates that it cannot be explained by poorer acuity in processing chromatic information, higher contrast of luminance information or differences in the statistical structure of colour and luminance information in natural scenes. It is therefore proposed that there is a neural mechanism that actively promotes luminance information. Chapter 4 and Experiments 5.1 and 5.3 aimed to investigate whether the presence of both chromatic and luminance information improves edge localisation performance. Participant performance in a Vernier acuity (alignment) task was compared to predictions from three models; ‘winner takes all’, unweighted averaging and maximum likelihood estimation (a form of weighted averaging). Despite several attempts to differentiate the models we failed to increase the differences in model predictions sufficiently and it was not possible to determine whether edge localisation was enhanced by the presence of both cues. In Experiment 5.4 we investigated how edges are localised when colour and luminance cues conflict, using the method of adjustment. Maximum likelihood estimation was used to make predictions based on measurements of each cue in isolation. These predictions were then compared to observed data. It was found that, whilst maximum likelihood estimation captured the pattern of the data, it consistently over-estimated the weight of the luminance component. It is suggested that chromatic information may be weighted more heavily than predicted as it is more useful for detecting object boundaries in natural scenes. In Chapter 6 a novel approach, perturbation discrimination, was used to investigate how the spatial arrangement of chromatic and luminance cues, and the type of chromatic and luminance information, can affect cue combination. Perturbation discrimination requires participants to select the grating stimulus that contains spatial perturbation. If one cue dominated over the other it was expected that this would be reflected by masking and increased perturbation detection thresholds. We compared perturbation thresholds for chromatic and luminance defined line and square-wave gratings in isolation and when presented with a mask of the other channel and other grating type. For example, the perturbation threshold for a luminance line target alone was compared to the threshold for a luminance line target presented with a chromatic square-wave target. The introduction of line masks caused masking for both combinations. Introduction of an achromatic square-wave mask had no effect on perturbation thresholds for chromatic line targets. However, the introduction of a chromatic square-wave mask to luminance line targets improved perturbation discrimination performance. This suggests that the perceived location of the chromatic edges is determined by the location of the luminance lines. Finally, in Chapter 7, we investigated whether chromatic blur is constrained by luminance information in bipartite edges. Earlier in the thesis we demonstrated that luminance information constrains chromatic blur in natural scenes, but also that chromatic information has more influence than expected when colour and luminance edges conflict. This difference may be due to differences in the stimuli or due to differences in the task. The luminance masking effect found using natural scenes was replicated using bipartite edges. Therefore, the finding that luminance constrains chromatic blur is not limited to natural scene stimuli. This suggests that colour and luminance are combined differently for blur discrimination tasks and edge localisation tasks. Overall we can see that luminance often dominates in edge perception tasks. For blur discrimination this seems to be because the mechanisms differ. For edge localisation it might be simply that luminance cues are often higher contrast and, when this is equated, chromatic cues are actually a good indicator of edge location

Cue combination of conflicting color and luminance edges

Abrupt changes in the color or luminance of a visual image potentially indicate object boundaries. Here, we consider how these cues to the visual “edge” location are combined when they conflict. We measured the extent to which localization of a compound edge can be predicted from a simple maximum likelihood estimation model using the reliability of chromatic (L−M) and luminance signals alone. Maximum likelihood estimation accurately predicted thepatternof results across a range of contrasts. Predictions consistently overestimated the relative influence of the luminance cue; although L−M is often considered a poor cue for localization, it was used more than expected. This need not indicate that the visual system is suboptimal but that its priors about which cue is moreusefulare not flat. This may be because, although strong changes in chromaticity typically represent object boundaries, changes in luminance can be caused by either a boundary or a shadow

Edge enhancement improves disruptive camouflage by emphasising false edges and creating pictorial relief

Disruptive colouration is a visual camouflage composed of false edges and boundaries. Many disruptively camouflaged animals feature enhanced edges; light patches are surrounded by a lighter outline and/or a dark patches are surrounded by a darker outline. This camouflage is particularly common in amphibians, reptiles and lepidopterans. We explored the role that this pattern has in creating effective camouflage. In a visual search task utilising an ultra-large display area mimicking search tasks that might be found in nature, edge enhanced disruptive camouflage increases crypsis, even on substrates that do not provide an obvious visual match. Specifically, edge enhanced camouflage is effective on backgrounds both with and without shadows; i.e. this is not solely due to background matching of the dark edge enhancement element with the shadows. Furthermore, when the dark component of the edge enhancement is omitted the camouflage still provided better crypsis than control patterns without edge enhancement. This kind of edge enhancement improved camouflage on all background types. Lastly, we show that edge enhancement can create a perception of multiple surfaces. We conclude that edge enhancement increases the effectiveness of disruptive camouflage through mechanisms that may include the improved disruption of the object outline by implying pictorial relief