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1.
Sci Rep ; 10(1): 9246, 2020 06 08.
Article in English | MEDLINE | ID: mdl-32514003

ABSTRACT

An apparent and common feature of aposematic patterns is that they contain a high level of achromatic (luminance) contrast, for example, many warning signals combine black spots and stripes with a lighter colour such as yellow. However, the potential importance of achromatic contrast, as distinct from colour contrast, in reducing predation has been largely overlooked. Here, using domestic chicks as a model predator, we manipulated the degree of achromatic contrast in warning patterns to test if high luminance contrast in aposematic signals is important for deterring naïve predators. We found that the chicks were less likely to approach and eat prey with high contrast compared to low contrast patterns. These findings suggest that aposematic prey patterns with a high luminance contrast can benefit from increased survival through eliciting unlearned biases in naïve avian predators. Our work also highlights the importance of considering luminance contrast in future work investigating why aposematic patterns take the particular forms that they do.

2.
Iperception ; 10(5): 2041669519877435, 2019.
Article in English | MEDLINE | ID: mdl-31555431

ABSTRACT

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.

3.
Sci Rep ; 8(1): 6599, 2018 04 26.
Article in English | MEDLINE | ID: mdl-29700366

ABSTRACT

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.


Subject(s)
Biological Mimicry , Color , Physical Appearance, Body , Pigmentation , Animals , Male
4.
R Soc Open Sci ; 5(2): 170801, 2018 Feb.
Article in English | MEDLINE | ID: mdl-29515822

ABSTRACT

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.

5.
Iperception ; 9(1): 2041669517752372, 2018.
Article in English | MEDLINE | ID: mdl-29375752

ABSTRACT

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

6.
Gen Comp Endocrinol ; 256: 123-129, 2018 01 15.
Article in English | MEDLINE | ID: mdl-28596079

ABSTRACT

The genomics era has brought along the completed sequencing of a large number of bird genomes that cover a broad range of the avian phylogenetic tree (>30 orders), leading to major novel insights into avian biology and evolution. Among recent findings, the discovery that birds lack a large number of protein coding genes that are organized in highly conserved syntenic clusters in other vertebrates is very intriguing, given the physiological importance of many of these genes. A considerable number of them play prominent endocrine roles, suggesting that birds evolved compensatory genetic or physiological mechanisms that allowed them to survive and thrive in spite of these losses. While further studies are needed to establish the exact extent of avian gene losses, these findings point to birds as potentially highly relevant model organisms for exploring the genetic basis and possible therapeutic approaches for a wide range of endocrine functions and disorders.


Subject(s)
Birds/genetics , Endocrine System/physiology , Genomics , Animals , Genes
7.
Sci Rep ; 7(1): 13672, 2017 10 20.
Article in English | MEDLINE | ID: mdl-29057907

ABSTRACT

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.


Subject(s)
Pigmentation , Visual Perception , Animals , Biological Evolution , Color , Computer Simulation , Environment , Female , Humans , Male , Orientation , Photic Stimulation , Plant Leaves , Predatory Behavior , Psychophysics
8.
Am Nat ; 186(4): 553-63, 2015 Oct.
Article in English | MEDLINE | ID: mdl-26655578

ABSTRACT

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.


Subject(s)
Environment , Light , Pigmentation , Visual Perception , Animals , Color , Computer Simulation , Cues
9.
Sci Rep ; 5: 14093, 2015 Sep 14.
Article in English | MEDLINE | ID: mdl-26364979

ABSTRACT

The dopamine transporter (DAT) is a major regulator of synaptic dopamine (DA) availability. It plays key roles in motor control and motor learning, memory formation, and reward-seeking behavior, is a major target of cocaine and methamphetamines, and has been assumed to be conserved among vertebrates. We have found, however, that birds, crocodiles, and lizards lack the DAT gene. We also found that the unprecedented loss of this important gene is compensated for by the expression of the noradrenaline transporter (NAT) gene, and not the serotonin transporter genes, in dopaminergic cells, which explains the peculiar pharmacology of the DA reuptake activity previously noted in bird striatum. This unexpected pattern contrasts with that of ancestral vertebrates (e.g. fish) and mammals, where the NAT gene is selectively expressed in noradrenergic cells. DA circuits in birds/reptiles and mammals thus operate with an analogous reuptake mechanism exerted by different genes, bringing new insights into gene expression regulation in dopaminergic cells and the evolution of a key molecular player in reward and addiction pathways.


Subject(s)
Dopamine Plasma Membrane Transport Proteins/metabolism , Animals , Birds , Corpus Striatum/metabolism , Dopamine/metabolism , Dopamine Plasma Membrane Transport Proteins/classification , Dopamine Plasma Membrane Transport Proteins/genetics , Dopaminergic Neurons/metabolism , Male , Norepinephrine Plasma Membrane Transport Proteins/genetics , Norepinephrine Plasma Membrane Transport Proteins/metabolism , Phylogeny , Promoter Regions, Genetic , Regulatory Elements, Transcriptional/genetics , Reptiles
10.
PLoS One ; 10(7): e0130093, 2015.
Article in English | MEDLINE | ID: mdl-26134671

ABSTRACT

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.


Subject(s)
Face/anatomy & histology , Fertility/physiology , Luteal Phase/physiology , Ovulation/physiology , Adolescent , Adult , Beauty , Color , Face/blood supply , Female , Humans , Pattern Recognition, Visual/physiology , Photography , Sexual Behavior/physiology , Voice/physiology
11.
Funct Ecol ; 29(9): 1165-1177, 2015 09.
Article in English | MEDLINE | ID: mdl-26937063

ABSTRACT

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.

12.
Curr Biol ; 23(3): 260-4, 2013 Feb 04.
Article in English | MEDLINE | ID: mdl-23333313

ABSTRACT

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.


Subject(s)
Nesting Behavior , Ovum , Pigmentation , Quail , Visual Perception , Animals , Choice Behavior , Female
13.
J Vis ; 10(4): 12.1-22, 2010 Apr 27.
Article in English | MEDLINE | ID: mdl-20465332

ABSTRACT

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.


Subject(s)
Color Perception/physiology , Form Perception/physiology , Models, Neurological , Sensory Thresholds/physiology , Contrast Sensitivity/physiology , Humans , Motion Perception/physiology , Nonlinear Dynamics , Photic Stimulation/methods , Predictive Value of Tests , Psychophysics , Visual Fields/physiology
14.
Seeing Perceiving ; 23(4): 349-72, 2010.
Article in English | MEDLINE | ID: mdl-21466148

ABSTRACT

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.


Subject(s)
Contrast Sensitivity/physiology , Discrimination, Psychological/physiology , Memory/physiology , Models, Theoretical , Neurons/physiology , Humans , Photic Stimulation
15.
J Vis ; 9(1): 37.1-14, 2009 Jan 29.
Article in English | MEDLINE | ID: mdl-19271907

ABSTRACT

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.


Subject(s)
Attention , Lighting , Visual Perception , Humans , Photic Stimulation/methods , Reaction Time , Visual Perception/physiology , Young Adult
16.
Br J Dermatol ; 160(5): 1094-7, 2009 May.
Article in English | MEDLINE | ID: mdl-19120338

ABSTRACT

BACKGROUND: Epidermolysis bullosa (EB) encompasses a heterogeneous group of inherited skin disorders associated with trauma-induced blistering. The junctional forms of EB (JEB), Herlitz JEB, non-Herlitz JEB and JEB associated with pyloric atresia have all been attributed to autosomal recessive inheritance. We describe a 7-year-old girl with defective dental enamel, trauma-induced blistering and subsequent scarring. Her mother, a carrier of the mutation p.G627V in the collagen XVII gene (COL17A1) had evidence of hypoplastic dental enamel without skin blistering. Her grandmother had non-Herlitz JEB as a result of a compound heterozygous mutation in COL17A1 (p.G627V and c.3514ins25). OBJECTIVES: To explore the molecular, ultrastructural and immunofluorescence findings of the first case of dominant JEB. METHODS: Mutational analysis of COL17A1 was performed on the proband's genomic DNA. In addition, transmission electron microscopy and immunofluorescence microscopy were performed on a nonlesional skin biopsy from the proband and an unrelated healthy control. RESULTS: Direct sequencing revealed a heterozygous glycine substitution mutation, p.G627V, in COL17A1. No discernible morphological abnormalities were found on transmission electron microscopy; however, immunofluorescence microscopy revealed findings of an altered distribution pattern for type XVII collagen epitopes close to the dermal-epidermal junction. CONCLUSION: This report describes the first case of dominant JEB. Although some heterozygous mutations in COL17A1 are known to cause dental abnormalities none were associated with skin fragility. The dominant-negative interference between the proband's mutated type XVII collagen and the wild-type allele appears to render the skin prone to trauma-induced blister formation. Alternatively, other undisclosed modifying genetic or epigenetic factors might explain why the patient gets blistering whereas her mother, who has the same COL17A1 mutation, has no skin fragility.


Subject(s)
Autoantigens/genetics , Dental Enamel/abnormalities , Epidermolysis Bullosa, Junctional/genetics , Non-Fibrillar Collagens/genetics , Blister/etiology , Child , DNA Mutational Analysis , Dental Enamel/pathology , Epidermolysis Bullosa, Junctional/pathology , Female , Genetic Variation/genetics , Heterozygote , Humans , Microscopy, Fluorescence , Pedigree , Collagen Type XVII
17.
Philos Trans R Soc Lond B Biol Sci ; 364(1516): 449-61, 2009 Feb 27.
Article in English | MEDLINE | ID: mdl-18990671

ABSTRACT

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.


Subject(s)
Adaptation, Biological/physiology , Appetitive Behavior/physiology , Pigmentation , Visual Fields/physiology , Visual Perception/physiology , Animals
18.
ACS Appl Mater Interfaces ; 1(9): 2021-9, 2009 Sep.
Article in English | MEDLINE | ID: mdl-20355828

ABSTRACT

The effect of low levels of cross-linking on the adhesive and mechanical properties of waterborne pressure-sensitive adhesives was investigated. We have taken advantage of a core-shell latex particle morphology obtained by emulsion polymerization to create a heterogeneous structure of cross-links without major modification of the monomer composition. The latex particles comprise a shell containing cross-linkable diacetone acrylamide (DAAM) repeat units localized on the periphery of a slightly softer core copolymer of very similar composition. Adipic acid dihydrazide was added to the latex prior to film formation to react with DAAM repeat units and affect interfacial cross-linking between particles in the adhesive films. The honeycomb-like structure obtained after drying of the latex results in a good balance between the dissipative properties required for adhesion and the resistance to creep. The characterization of the mechanical properties of the films shows that the chosen cross-linking method creates a percolating lightly cross-linked network, swollen with a nearly un-cross-linked component. With this cross-linking method, the linear viscoelastic properties of the soft films are nearly unaffected by the cross-linking while the nonlinear tensile properties are greatly modified. As a result, the long-term shear resistance of the adhesive film improves very significantly while the peel force remains nearly the same. A simple rheological model is used to interpret qualitatively the changes in the material parameters induced by cross-linking.


Subject(s)
Membranes, Artificial , Polymers/chemistry , Surface Properties , Adhesives , Cross-Linking Reagents/chemistry , Elasticity , Materials Testing , Particle Size , Tensile Strength , Viscosity
19.
Proc Natl Acad Sci U S A ; 105(33): 11634-9, 2008 Aug 19.
Article in English | MEDLINE | ID: mdl-18695241

ABSTRACT

Polar regions are particularly sensitive to climate change, with the potential for significant feedbacks between ocean circulation, sea ice, and the ocean carbon cycle. However, the difficulty in obtaining in situ data means that our ability to detect and interpret change is very limited, especially in the Southern Ocean, where the ocean beneath the sea ice remains almost entirely unobserved and the rate of sea-ice formation is poorly known. Here, we show that southern elephant seals (Mirounga leonina) equipped with oceanographic sensors can measure ocean structure and water mass changes in regions and seasons rarely observed with traditional oceanographic platforms. In particular, seals provided a 30-fold increase in hydrographic profiles from the sea-ice zone, allowing the major fronts to be mapped south of 60 degrees S and sea-ice formation rates to be inferred from changes in upper ocean salinity. Sea-ice production rates peaked in early winter (April-May) during the rapid northward expansion of the pack ice and declined by a factor of 2 to 3 between May and August, in agreement with a three-dimensional coupled ocean-sea-ice model. By measuring the high-latitude ocean during winter, elephant seals fill a "blind spot" in our sampling coverage, enabling the establishment of a truly global ocean-observing system.


Subject(s)
Ice , Seals, Earless , Seawater , Animals , Temperature
20.
Proc Biol Sci ; 275(1649): 2299-308, 2008 Oct 22.
Article in English | MEDLINE | ID: mdl-18628119

ABSTRACT

Natural visual scenes are rich in information, and any neural system analysing them must piece together the many messages from large arrays of diverse feature detectors. It is known how threshold detection of compound visual stimuli (sinusoidal gratings) is determined by their components' thresholds. We investigate whether similar combination rules apply to the perception of the complex and suprathreshold visual elements in naturalistic visual images. Observers gave magnitude estimations (ratings) of the perceived differences between pairs of images made from photographs of natural scenes. Images in some pairs differed along one stimulus dimension such as object colour, location, size or blur. But, for other image pairs, there were composite differences along two dimensions (e.g. both colour and object-location might change). We examined whether the ratings for such composite pairs could be predicted from the two ratings for the respective pairs in which only one stimulus dimension had changed. We found a pooling relationship similar to that proposed for simple stimuli: Minkowski summation with exponent 2.84 yielded the best predictive power (r=0.96), an exponent similar to that generally reported for compound grating detection. This suggests that theories based on detecting simple stimuli can encompass visual processing of complex, suprathreshold stimuli.


Subject(s)
Sensory Thresholds/physiology , Visual Perception/physiology , Female , Humans , Male , Models, Biological , Models, Statistical , Photic Stimulation/methods
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