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.

Is countershading camouflage robust to lighting change due to weather?

Countershading is a pattern of coloration thought to have evolved in order to implement camouflage. By adopting a pattern of coloration that makes the surface facing towards the sun darker and the surface facing away from the sun lighter, the overall amount of light reflected off an animal can be made more uniformly bright. Countershading could hence contribute to visual camouflage by increasing background matching or reducing cues to shape. However, the usefulness of countershading is constrained by a particular pattern delivering 'optimal' camouflage only for very specific lighting conditions. In this study, we test the robustness of countershading camouflage to lighting change due to weather, using human participants as a 'generic' predator. In a simulated three-dimensional environment, we constructed an array of simple leaf-shaped items and a single ellipsoidal target 'prey'. We set these items in two light environments: strongly directional 'sunny' and more diffuse 'cloudy'. The target object was given the optimal pattern of countershading for one of these two environment types or displayed a uniform pattern. By measuring detection time and accuracy, we explored whether and how target detection depended on the match between the pattern of coloration on the target object and scene lighting. Detection times were longest when the countershading was appropriate to the illumination; incorrectly camouflaged targets were detected with a similar pattern of speed and accuracy to uniformly coloured targets. We conclude that structural changes in light environment, such as caused by differences in weather, do change the effectiveness of countershading camouflage.

Stacking Chairs: Local Sense and Global Nonsense.

We report a confusing stimulus which demonstrates the power of local interpretation of three-dimensional structure to disrupt a coherent global perception.

Establishing the behavioural limits for countershaded camouflage.

Countershading is a ubiquitous patterning of animals whereby the side that typically faces the highest illumination is darker. When tuned to specific lighting conditions and body orientation with respect to the light field, countershading minimizes the gradient of light the body reflects by counterbalancing shadowing due to illumination, and has therefore classically been thought of as an adaptation for visual camouflage. However, whether and how crypsis degrades when body orientation with respect to the light field is non-optimal has never been studied. We tested the behavioural limits on body orientation for countershading to deliver effective visual camouflage. We asked human participants to detect a countershaded target in a simulated three-dimensional environment. The target was optimally coloured for crypsis in a reference orientation and was displayed at different orientations. Search performance dramatically improved for deviations beyond 15 degrees. Detection time was significantly shorter and accuracy significantly higher than when the target orientation matched the countershading pattern. This work demonstrates the importance of maintaining body orientation appropriate for the displayed camouflage pattern, suggesting a possible selective pressure for animals to orient themselves appropriately to enhance crypsis.

Three-Dimensional Camouflage: Exploiting Photons to Conceal Form.

Many animals have a gradation of body color, termed "countershading," where the areas that are typically exposed to more light are darker. One hypothesis is that this patterning enhances visual camouflage by making the retinal image of the animal match that of the background, a fundamentally two-dimensional theory. More controversially, countershading may also obliterate cues to three-dimensional (3D) shape delivered by shading. Despite relying on distinct cognitive mechanisms, these two potential functions hitherto have been amalgamated in the literature. It has previously not been possible to validate either hypothesis empirically, because there has been no general theory of optimal countershading that allows quantitative predictions to be made about the many environmental parameters involved. Here we unpack the logical distinction between using countershading for background matching and using it to obliterate 3D shape. We use computational modeling to determine the optimal coloration for the camouflage of 3D shape. Our model of 3D concealment is derived from the physics of light and informed by perceptual psychology: we simulate a 3D world that incorporates naturalistic lighting environments. The model allows us to predict countershading coloration for terrestrial environments, for any body shape and a wide range of ecologically relevant parameters. The approach can be generalized to any light distribution, including those underwater.

Changes in Women's Facial Skin Color over the Ovulatory Cycle are Not Detectable by the Human Visual System.

Human ovulation is not advertised, as it is in several primate species, by conspicuous sexual swellings. However, there is increasing evidence that the attractiveness of women's body odor, voice, and facial appearance peak during the fertile phase of their ovulatory cycle. Cycle effects on facial attractiveness may be underpinned by changes in facial skin color, but it is not clear if skin color varies cyclically in humans or if any changes are detectable. To test these questions we photographed women daily for at least one cycle. Changes in facial skin redness and luminance were then quantified by mapping the digital images to human long, medium, and shortwave visual receptors. We find cyclic variation in skin redness, but not luminance. Redness decreases rapidly after menstrual onset, increases in the days before ovulation, and remains high through the luteal phase. However, we also show that this variation is unlikely to be detectable by the human visual system. We conclude that changes in skin color are not responsible for the effects of the ovulatory cycle on women's attractiveness.

Orientation to the sun by animals and its interaction with crypsis.

Orientation with respect to the sun has been observed in a wide range of species and has generally been interpreted in terms of thermoregulation and/or ultraviolet (UV) protection. For countershaded animals, orientation with respect to the sun may also result from the pressure to exploit the gradient of coloration optimally to enhance crypsis.Here, we use computational modelling to predict the optimal countershading pattern for an oriented body. We assess how camouflage performance declines as orientation varies using a computational model that incorporates realistic lighting environments.Once an optimal countershading pattern for crypsis has been chosen, we determine separately how UV protection/irradiation and solar thermal inflow fluctuate with orientation.We show that body orientations that could optimally use countershading to enhance crypsis are very similar to those that allow optimal solar heat inflow and UV protection.Our findings suggest that crypsis has been overlooked as a selective pressure on orientation and that new experiments should be designed to tease apart the respective roles of these different selective pressures. We propose potential experiments that could achieve this.

Egg-laying substrate selection for optimal camouflage by quail.

Camouflage is conferred by background matching and disruption, which are both affected by microhabitat. However, microhabitat selection that enhances camouflage has only been demonstrated in species with discrete phenotypic morphs. For most animals, phenotypic variation is continuous; here we explore whether such individuals can select microhabitats to best exploit camouflage. We use substrate selection in a ground-nesting bird (Japanese quail, Coturnix japonica). For such species, threat from visual predators is high and egg appearance shows strong between-female variation. In quail, variation in appearance is particularly obvious in the amount of dark maculation on the light-colored shell. When given a choice, birds consistently selected laying substrates that made visual detection of their egg outline most challenging. However, the strategy for maximizing camouflage varied with the degree of egg maculation. Females laying heavily maculated eggs selected the substrate that more closely matched egg maculation color properties, leading to camouflage through disruptive coloration. For lightly maculated eggs, females chose a substrate that best matched their egg background coloration, suggesting background matching. Our results show that quail "know" their individual egg patterning and seek out a nest position that provides most effective camouflage for their individual phenotype.

Perception of suprathreshold naturalistic changes in colored natural images.

Simple everyday tasks, such as visual search, require a visual system that is sensitive to differences. Here we report how observers perceive changes in natural image stimuli, and what happens if objects change color, position, or identity-i.e., when the external scene changes in a naturalistic manner. We investigated whether a V1-based difference-prediction model can predict the magnitude ratings given by observers to suprathreshold differences in numerous pairs of natural images. The model incorporated contrast normalization and surround suppression, and elongated receptive-fields. Observers' ratings were better predicted when the model included phase invariance, and even more so when the stimuli were inverted and negated to lessen their semantic impact. Some feature changes were better predicted than others: the model systematically underpredicted observers' perception of the magnitude of blur, but over-predicted their ability to report changes in textures.

Magnitude of perceived change in natural images may be linearly proportional to differences in neuronal firing rates.

We are studying how people perceive naturalistic suprathreshold changes in the colour, size, shape or location of items in images of natural scenes, using magnitude estimation ratings to characterise the sizes of the perceived changes in coloured photographs. We have implemented a computational model that tries to explain observers' ratings of these naturalistic differences between image pairs. We model the action-potential firing rates of millions of neurons, having linear and non-linear summation behaviour closely modelled on real VI neurons. The numerical parameters of the model's sigmoidal transducer function are set by optimising the same model to experiments on contrast discrimination (contrast 'dippers') on monochrome photographs of natural scenes. The model, optimised on a stimulus-intensity domain in an experiment reminiscent of the Weber-Fechner relation, then produces tolerable predictions of the ratings for most kinds of naturalistic image change. Importantly, rating rises roughly linearly with the model's numerical output, which represents differences in neuronal firing rate in response to the two images under comparison; this implies that rating is proportional to the neuronal response.

Search for gross illumination discrepancies in images of natural objects.

Shadows may be "discounted" in human visual perception because they do not provide stable, lighting-invariant, information about the properties of objects in the environment. Using visual search, R. A. Rensink and P. Cavanagh (2004) found that search for an upright discrepant shadow was less efficient than for an inverted one. Here we replicate and extend this work using photographs of real objects (pebbles) and their shadows. The orientation of the target shadows was varied between 30 and 180 degrees. Stimuli were presented upright (light from above, the usual situation in the world) or inverted (light from below, unnatural lighting). RTs for upright images were slower for shadows angled at 30 degrees, exactly as found by Rensink and Cavanagh. However, for all other shadow angles tested, the RTs were faster for upright images. This suggests, for small discrepancies in shadow orientation, a switch of processing from a relatively coarse-scaled shadow system to other general-purpose visual routines. Manipulations of the visual heterogeneity of the pebbles that cast the shadows differentially influenced performance. For inverted images, heterogeneity had the expected influence: reducing search efficiency and increasing overall search time. This effect was greatly reduced when images were presented upright, presumably when the distractors were processed as shadows. We suggest that shadows may be processed in a functionally separate, spatially coarse, mechanism. The pattern of results suggests that human vision does not use a shadow-suppressing system in search tasks.

Camouflage and visual perception.

How does an animal conceal itself from visual detection by other animals? This review paper seeks to identify general principles that may apply in this broad area. It considers mechanisms of visual encoding, of grouping and object encoding, and of search. In most cases, the evidence base comes from studies of humans or species whose vision approximates to that of humans. The effort is hampered by a relatively sparse literature on visual function in natural environments and with complex foraging tasks. However, some general constraints emerge as being potentially powerful principles in understanding concealment--a 'constraint' here means a set of simplifying assumptions. Strategies that disrupt the unambiguous encoding of discontinuities of intensity (edges), and of other key visual attributes, such as motion, are key here. Similar strategies may also defeat grouping and object-encoding mechanisms. Finally, the paper considers how we may understand the processes of search for complex targets in complex scenes. The aim is to provide a number of pointers towards issues, which may be of assistance in understanding camouflage and concealment, particularly with reference to how visual systems can detect the shape of complex, concealed objects.