Modelling colour constancy in fish: implications for vision and signalling in water.

Ratios of excitations in each cone-photoreceptor class produced by light reflected from pairs of surfaces in a scene are almost invariant under natural illuminant changes. The stability of these spatially defined ratios may explain the remarkable ability of human observers to efficiently discriminate illuminant changes from changes in surface reflectances. Spatial cone-excitation ratios are not, however, exactly invariant. This study is concerned with observers' sensitivity to these invariance violations. Simulations of Mondrian paintings with either 49 or two natural surfaces under Planckian illuminants were presented as images on a computer-controlled display in a two-interval experimental design: in one interval, the surfaces underwent an illuminant change; in the other interval, the surfaces underwent the same change but the images were then corrected so that, for each cone class, ratios of excitations were preserved exactly. Although the intervals with corrected images corresponded individually to highly improbable natural events, observers systematically misidentified them as containing the illuminant changes, the probability of error increasing as the violation of invariance in the other interval increased. For the range of illuminants and surfaces tested, sensitivity to violations of invariance was found to depend on cone class: it was greatest for long-wavelength-sensitive cones and least for short-wavelength-sensitive cones. Spatial cone-excitation ratios, or some closely related quantities, seem to be the cues preferred by observers for making inferences about surface illuminant changes.

The red-green axis of colour vision evolved recently in primate evolutionary history. Signals serving red-green colour vision travel together with signals serving spatial vision, in the parvocellular (PC) division of the subcortical visual pathway. However, the question of whether receptive fields of PC pathway cells are specialized to transmit red-green colour signals remains unresolved. We addressed this question in single-cell recordings from the lateral geniculate nucleus of anaesthetized marmosets. Marmosets show a high proportion of dichromatic (red-green colour-blind) individuals, allowing spatial and colour tuning properties of PC cells to be directly compared in dichromatic and trichromatic visual systems. We measured spatial frequency tuning for sine gratings that provided selective stimulation of individual photoreceptor types. We found that in trichromatic marmosets, the foveal visual field representation is dominated by red-green colour-selective PC cells. Colour selectivity of PC cells is reduced at greater eccentricities, but cone inputs to centre and surround are biased to create more selectivity than predicted by a purely 'random wiring' model. Thus, one-to-one connections in the fovea are sufficient, but not necessary, to create colour-selective responses. The distribution of spatial tuning properties for achromatic stimuli shows almost complete overlap between PC cells recorded in dichromatic and trichromatic marmosets. These data indicate that transmission of red-green colour signals has been enabled by centre-surround receptive fields of PC cells, and has not altered the capacity of PC cells to serve high-acuity vision at high stimulus contrast.

Many species of stomatopod crustaceans have multiple spectral classes of photoreceptors in their retinas. Behavioral evidence also indicates that stomatopods are capable of discriminating objects by their spectral differences alone. Most animals use only two to four different types of photoreceptors in their color vision systems, typically with broad sensitivity functions, but the stomatopods apparently include eight or more narrowband photoreceptor classes for color recognition. It is also known that stomatopods use several colored body regions in social interactions. To examine why stomatopods may be so ‘concerned’ with color, we measured the absorption spectra of visual pigments and intrarhabdomal filters, and the reflectance spectra from different parts of the bodies of several individuals of the gonodactyloid stomatopod species, Gonodactylus smithii. We then applied a model of multiple dichromatic channels for color encoding to examine whether the finely tuned color vision was specifically co-evolved with their complex color signals. Although the eye design of stomatopods seems suitable for detecting color signals of their own, the detection of color signals from other animals, such as reef fishes, can be enhanced as well. Color vision in G. smithii is therefore not exclusively adapted to detect its own color signals, but the spectral tuning of some photoreceptors (e.g. midband Rows 2 and 3) enhances the contrast of certain color signals to a large enough degree to make co-evolution between color vision and these rather specific color signals likely.

How animals visually perceive the environment is key to understanding important ecological behaviors, such as predation, foraging, and mating. This study focuses on the visual system properties and visual perception of color in the largemouth bass Micropterus salmoides. This study (1) documents the number and spectral sensitivity of photoreceptors, (2) uses these parameters to model visual perception, and (3) tests the model of color perception using a behavioral assay. Bass possess single cone cells maximally sensitive at 535 nm, twin cone cells maximally sensitive at 614 nm, and rod cells maximally sensitive at 528 nm. A simple model of visual perception predicted that bass should not be able to discern between chartreuse yellow and white nor between green and blue. In contrast, bass should be able to discern red from all achromatic (i.e., gray scale) stimuli. These predictions were partially upheld in behavioral trials. In behavioral trials, bass were first trained to recognize a target color to receive a food reward, and then tested on their ability to differentiate between their target color and a color similar in brightness. Bass trained to red and green could easily discern their training color from all other colors for target colors that were similar in brightness (white and black, respectively). This study shows that bass possess dichromatic vision and do use chromatic (i.e., color) cues in making visual-based decisions.

Animal communication relies on conspicuous signals and compatible signal perception abilities. Good signal perception abilities are particularly important for polymorphic animals where mate choice can be a challenge. Behavioral studies suggest that polymorphic damselflies use their varying body colorations and/or color patterns as communication signal for mate choice and to control mating frequencies. However, solid evidence for this hypothesis combining physiological with spectral and behavioral data is scarce. We investigated this question in the Australian common blue tail damselfly, Ischnura heterosticta, which has pronounced female-limited polymorphism: andromorphs have a male-like blue coloration and gynomorphs display green/grey colors. We measured body color reflectance and investigated the visual capacities of each morph, showing that I. heterosticta have at least three types of photoreceptors sensitive to UV, blue, and green wavelength, and that this visual perception ability enables them to detect the spectral properties of the color signals emitted from the various color morphs in both males and females. We further demonstrate that different color morphs can be discriminated against each other and the vegetation based on color contrast. Finally, these findings were supported by field observations of natural mating pairs showing that mating partners are indeed chosen based on their body coloration. Our study provides the first comprehensive evidence for the function of body coloration on mate choice in polymorphic damselflies.

1 - Colour vision: theories and principles

This paper gives an overview of behavioral studies on the color and polarization vision of the Japanese yellow swallowtail butterfly, Papilio xuthus. We focus on indoor experiments on foraging individuals. Butterflies trained to visit a disk of certain color correctly select that color among various other colors and/or shades of gray. Correct selection persists under colored illumination, but is systematically shifted by background colors, indicating color constancy and simultaneous color contrast. While their eyes contain six classes of spectral receptors, their wavelength discrimination performance indicates that their color vision is tetrachromatic. P. xuthus innately prefers brighter targets, but can be trained to select dimmer ones under certain conditions. Butterflies trained to a dark red stimulus select an orange disk presented on a bright gray background over one on dark gray. The former probably appears darker to them, indicating brightness contrast. P. xuthus has a strong innate preference for vertically polarized light, but the selection of polarized light changes depending on the intensity of simultaneously presented unpolarized light. Discrimination of polarization also depends on background intensity. Similarities between brightness and polarization vision suggest that P. xuthus perceive polarization angle as brightness, such that vertical polarization appears brighter than horizontal polarization.

A color constancy mechanism is proposed to perceive the image color from the strengths of three RGB responses of a color camera or the like, independent of the color of the light illuminating the object. The first step of the method is to use a finite-dimensional linear model to estimate the color signals. It is shown that any color signal can be characterized as a linear combination of four principal component basis functions. Once the color signals have been estimated the unknown illuminant can be determined by S. Tominaga and B.A. Wandell's (1989) estimation method, which is based on a dichromatic model. In color constancy, the knowledge of surface reflectance helps in determining the canonical color descriptors despite the variations in the spectral power distribution of the ambient light. It is shown that the estimate of surface reflectance is derived from both the estimated color signal and illuminant straightforwardly. The model of a combination of the estimations of the surface spectral reflectance and the color signals can be employed in designing a color constancy electronic video camera which indeed improves the shortcomings of the traditional video camcorders.< <ETX xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">&gt;</ETX>

We have examined visual functions, including colour vision, in a patient with bilateral cortical lesions involving mainly the fusiform and lingual gyri, areas known to be involved in the central processing of chromatic stimuli. The patient has near normal (6/9) acuity, and his responses to tests of binocular function and spatial vision are normal, as are his discrimination of changes in target speed and surface lightness. He does, however, exhibit minor losses in the upper visual field, mild prosopagnosia and topographical agnosia, all conditions commonly associated with cerebral achromatopsia. Colour matches and spectral response data establish that his cone photoreceptors have normal spectral characteristics and his spectral sensitivity measured against a white background reveals normal postreceptoral chromatic function. The patient's colour discrimination for differences in wavelength, hue or saturation is, however, impaired and his colour naming is significantly disturbed, particularly for blues and greens. We have determined the areas of the chromaticity chart that correspond to his naming categories for surface colours, and show that changes in illuminant cause him to alter the names of surface colours in a manner consistent with the changes in their chromaticities. Other subjects with normal or congenital red-green deficient colour vision make many fewer name changes under changes in illuminant. We conclude that the patient's colour constancy is impaired as a consequence of abnormal central processing of colour vision.

Perception of colour depends on the spectral composition of light that reaches retina and depends as well on various mechanisms of visual system that processes information flow. The few important mechanisms can be distinguished in colour perception: colour adaptation, colour constancy and colour contrast. If the visual field has only one coloured object, then colour perception will be determined by spectral composition of the light and colour adaptation. Whereas mechanisms of colour constancy and colour contrast switches on when in the visual field there are at least two colour objects. Von Kries (1905) have attributed colour constancy phenomenon to theory of receptoral adaptation. But this is theory is applicable to local processes happening in relatively small size of visual field. Craven and Foster (1992) shows that receptor excitation ratios remains constant during change of illumination. But again remains unexplained colour change during adaptation. So, the hypothesis is raised that two processes attribute colour perception: local colour contrast calculation and global adaptation to the background. Therefore the experiments have been carried out to establish the colour contrast and background adaptation impact on colour perception.Four subjects with normal colour vision participated in the experiments. 40 Munsell samples (value 7 and chroma 4) illuminated with one reference illuminant (standard C) and 6 test illuminants (standard A, standard S, cardinal red, cardinal green, cardinal yellow and cardinal blue) were simulated on computer monitor. The dark-adapted subjects have been shown colour samples on neutral background illuminated with one of test illuminants. The task was to match the colour stimuli appearance under reference illuminant. The sequential asymmetric matching procedure was carried out. First subject adapts to neutral background under illuminan C then test stimulus and test background appears for limited time followed by readaptation to neutral background. Then subject is asked to adjust the colour mach under illuminant C. Two different experiment paradigms have been used. In the first one the stimulus had 2° on neutral background covering 20° of visual field. In the second paradigm the stimulus size was the same but the background was covering the all visual field. The adaptation and presentation timings were 1, 5, 30 or 60 seconds.Results under the first experiment paradigm show partial colour constancy for all subjects and for all test illuminants under various adaptation times. Despite quite a long adaptation time of 30 seconds subjects was unable to achieve full colour constancy. Colour constancy improves under second experiment paradigm. 60-second adaptation time is enough to achieve full colour constancy. The less adaptation time (1 or 30 second) gives partial colour constancy.The subject sees two colours (stimulus and background) during the experiment. The colour perception of stimulus and background changes during adaptation time. In the first 5 seconds of adaptation 40–60% of full adaptation level is reached (Fairchild and Reniff, 1995; Werner et al., 2000). Wesner and Shevell (1992, 1994) shown that signals from all visual field have influence on colour perception of object. The first part of our experiment shows that neutral background of 20° gives partial adaptation to background colour. Therefore, increase background illumination to full visual field gives full adaptation to background. So, subject perceives different coloured background under various illuminants as the same neutral (grey) background.We can state that two different systems have been distinguished in the process of colour perception. One system evaluates the colour difference between the stimulus and the background. Second system evaluates the colour of the background. Perceived colour difference during adaptation does not change much, but perceived colour of the background changes a lot and drifts towards neutral colour. The hypothesis of two level colour perception is supported by experimental data.&#x0D;

Centre-surround stimuli evoke colour appearances (resembling surface colours) which cannot be produced by a single homogeneous spot of light alone (eg brown or grey). Although this seems of great impact to a general theory of colour (including 'colour constancy'), the psychophysics of these 'minimal relational stimuli' is still less well understood than often assumed. On the basis of empirical as well as theoretical observations concerning centre-surround-type stimuli we introduce a relational model of colour coding. At the core of this model is the concept of a three-dimensional linear incremental colour code which behaves differently for increments and decrements. This model takes into account results on 'discounting the background' mechanisms and it is closely related to ratio-based relational concepts and to certain opponent-colour theories. In addition, it provides an analogue to the classical distinction between light and object colours, and covers colour appearances related to object colours as well. Within the conceptual framework offered, problems of complex colour perception (eg 'colour constancy') and judgmental modes are discussed. Conclusions regarding general limitations of three-dimensional modelling in colour perception are derived and corresponding refinements of the relational perspective are briefly outlined.

The color appearance of a stimulus depends on the context in which it is viewed. Well-known examples of context effects include simultaneous color contrast and color constancy. In simultaneous contrast, the color immediately surrounding a test influences its appearance. In color constancy, the visual system adapts to the ambient illumination to keep surface color approximately constant. Color constancy is a context effect because the way that the visual system interprets the light reaching the eye depends on the context defined by the ambient illumination.To predict accurately the color appearance of stimuli rendered on a CRT monitor, we need a theory of color context effects that applies to such stimuli. Simultaneous contrast suggests that the color perceived at a particular monitor location may vary with the immediate context provided by the image at other locations on the monitor (the monitor context). Color constancy suggests that the color appearance of stimuli displayed on a monitor may vary with the ambient illumination of the room in which the monitor is viewed (the illumination context), at least if images from a monitor are processed by the visual system in the same way as reflective surfaces.In this paper, we report initial experiments designed to measure psychophysically how the monitor context and the illumination context influence the appearance of stimuli presented on CRT color monitors. Our emphasis, however, is on the effect of the illumination context.

Most butterfly families expand the number of spectrally distinct photoreceptors in their compound eye by opsin gene duplications together with lateral filter pigments; however, most nymphalid genera have limited diversity, with only three or four spectral types of photoreceptor. Here, we examined the spatial pattern of opsin expression and photoreceptor spectral sensitivities in Heliconius erato, a nymphalid with duplicate ultraviolet opsin genes, UVRh1 and UVRh2 We found that the H. erato compound eye is sexually dimorphic. Females express the two UV opsin proteins in separate photoreceptors, but males do not express UVRh1. Intracellular recordings confirmed that females have three short wavelength-sensitive photoreceptors (λmax=356, ∼390 and 470 nm), while males have two (λmax=390 and ∼470 nm). We also found two long wavelength-sensitive photoreceptors (green, λmax∼555 nm, and red, λmax∼600 nm), which express the same LW opsin. The red cell's shifted sensitivity is probably due to perirhabdomal filtering pigments. Sexual dimorphism of the UV-absorbing rhodopsins may reflect the females' need to discriminate conspecifics from co-mimics. Red-green color vision may be used to detect differences in red coloration on Heliconius wings, or for host-plant identification. Among nymphalids so far investigated, only H. erato is known to possess five spectral classes of photoreceptor; sexual dimorphism of the eye via suppression of one class of opsin (here UVRh1 in males) has not - to our knowledge - been reported in any animal.

Diurnal and nocturnal hawkmoths have been shown to use colour vision for flower discrimination. Here, we present evidence that the nocturnal hawkmoth Deilephila elpenor and the diurnal hawkmoth Macroglossum stellatarum also have colour constancy. Colour constancy was shown in D. elpenor in two multiple-choice experiments with five different bluish colour patches under white and blue illumination and with five yellowish colour patches under white, blue and yellow illumination. The mechanism underlying colour constancy in both species was investigated in two dual-choice experiments. The choice behaviour is consistent with the use of the von Kries coefficient law. Although the moths have colour constancy, they react to the colour of the illumination. They make fewer choices when tested under the changed illumination, where they never receive a reward, compared with the training illumination. Even if colour constancy can be explained by a von Kries adaptation mechanism, the fact that the animals discriminate between different illuminations indicates that some additional process must be involved.

Invariance of surface color representations across illuminant changes in the human cortex