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M Vorobyev

Publications and source records attributed to M Vorobyev.

13 recordsLinked to original sources

Photoreceptor spectral sensitivities in terrestrial animals: adaptations for luminance and colour vision.

This review outlines how eyes of terrestrial vertebrates and insects meet the competing requirements of coding both spatial and spectral information. There is no unique solution to this problem. Thus, mammals and honeybees use their long-wavelength receptors for both achromatic (luminance) and colour vision, whereas flies and birds probably use separate sets of photoreceptors for the two purposes. In particular, we look at spectral tuning and diversification among 'long-wavelength' receptors (sensitivity maxima at greater than 500 nm), which play a primary role in luminance vision. Data on spectral sensitivities and phylogeny of visual photopigments can be incorporated into theoretical models to suggest how eyes are adapted to coding natural stimuli. Models indicate, for example, that animal colour vision--involving five or fewer broadly tuned receptors--is well matched to most natural spectra. We can also predict that the particular objects of interest and signal-to-noise ratios will affect the optimal eye design. Nonetheless, it remains difficult to account for the adaptive significance of features such as co-expression of photopigments in single receptors, variation in spectral sensitivities of mammalian L-cone pigments and the diversification of long-wavelength receptors that has occurred in several terrestrial lineages.

Adaptation, Biological↗

Discrimination of coloured patterns by honeybees through chromatic and achromatic cues.

We investigated pattern discrimination by worker honeybees, Apis mellifera, focusing on the roles of spectral cues and the angular size of patterns. Free-flying bees were trained to discriminate concentric patterns in a Y-maze. The rewarded pattern could be composed of either a cyan and a yellow colour, which presented both different chromatic and achromatic L-receptor contrast, or an orange and a blue colour, which presented different chromatic cues, but the same L-receptor contrast. The non-rewarded alternative was either a single-coloured disc with the colour of the central disc or the surrounding ring of the pattern, a checkerboard pattern with non-resolvable squares, the reversed pattern, or the elements of the training pattern (disc or ring alone). Bees resolved and learned both colour elements in the rewarded patterns and their spatial properties. When the patterns subtended large visual angles, this discrimination used chromatic cues only. Patterns with yellow or orange central discs were generalised toward the yellow and orange colours, respectively. When the patterns subtended a visual angle close to the detection limit and L-receptor contrast was mediating discrimination, pattern perception was reduced: bees perceived only the pattern element with higher contrast.

Animals↗

Detection of coloured patterns by honeybees through chromatic and achromatic cues.

We asked whether the detection range of two-coloured centre-surround patterns differs from that of single-coloured targets. Honeybees Apis mellifera were trained to distinguish between the presence and absence of a single-coloured disc or a coloured pattern at different visual angles. The patterns presented colours which were either different in chromatic and L-receptor contrasts to the background, equal in chromatic but different in L-receptor contrasts, or vice-versa. Patterns with colours presenting only chromatic contrast were also tested. Patterns with higher L-receptor contrast in its outer than in its inner element were better detected than patterns with a reversed L-contrast distribution. However, both were detected worse than single-coloured discs of the respective colours. When the L-receptor contrast was the same for both elements, the detection range of the two-coloured and single-coloured targets was the same. Patterns whose colours lacked L-receptor contrast were detected just as single-coloured targets of the same colours. These results demonstrate that both chromatic and L-receptor contrasts mediate the detection of coloured patterns and that particular distributions of L-receptor contrast within a target are better detected than others. This finding is consistent with the intervention of neurons with centre-surround receptive fields in the detection of coloured patterns.

Animals↗

Colour thresholds and receptor noise: behaviour and physiology compared.

Photoreceptor noise sets an absolute limit for the accuracy of colour discrimination. We compared colour thresholds in the honeybee (Apis mellifera) with this limit. Bees were trained to discriminate an achromatic stimulus from monochromatic lights of various wavelengths as a function of their intensity. Signal-to-noise ratios were measured by intracellular recordings in the three spectral types of photoreceptor cells. To model thresholds we assumed that discrimination was mediated by opponent mechanisms whose performance was limited by receptor noise. Most of the behavioural thresholds were close to those predicted from receptor signal-to-noise ratios, suggesting that colour discrimination in honeybees is affected by photoreceptor noise. Some of the thresholds were lower than this theoretical limit, which indicates summation of photoreceptor cell signals.

Animals↗

Spectral tuning of dichromats to natural scenes.

Multispectral images of natural scenes were collected from both forests and coral reefs. We varied the wavelength position of receptors in hypothetical dichromatic visual systems and, for each receptor pair estimated the percentage of discriminable points in natural scenes. The optimal spectral tuning predicted by this model results in photoreceptor pairs very like those of forest dwelling, dichromatic mammals and of coral reef fishes. Variations of the natural illuminants in forests have little or no effect on optimal spectral tuning, but variations of depth in coral reefs have moderate effects on the spectral placement of S and L cones. The ratio of S and L cones typically found in dichromatic mammals reduces the discriminability of forest scenes; in contrast, the typical ratio of S and L cones in coral reef fishes achieves nearly the optimal discrimination in coral reef scenes.

Animals↗

Detection of bright and dim colours by honeybees.

Honeybees, Apis mellifera, were trained to detect coloured disks with either a strong or a weak intensity difference against the background. Green, blue, ultraviolet-reflecting white and grey papers were reciprocally combined as targets or backgrounds, providing strong chromatic and/or achromatic cues. The behavioural performance of the honeybees was always symmetrical for both reciprocal target/background combinations of a colour pair, thus showing that target detection is independent of whether the colour is presented as a background or as a target in combination with the other colour. Bright targets against dim backgrounds and vice versa were detected more reliably than dim target/background combinations. This result favours the general assumption that the detectability of a coloured stimulus increases with increasing intensity.

Animals↗

Characterization of natural illuminants in forests and the use of digital video data to reconstruct illuminant spectra.

We describe illumination spectra in forests and show that they can be accurately recovered from recorded digital video images. Natural illuminant spectra of 238 samples measured in temperate forests were characterized by principal-component analysis. The spectra can be accurately approximated by the mean and the first two principal components. Compared with illumination under open skies, the loci of forest illuminants are displaced toward the green region in the chromaticity plots, and unlike open sky illumination they cannot be characterized by correlated color temperature. We show that it is possible to recover illuminant spectra accurately from digital video images by a linear least-squares-fit estimation technique. The use of digital video data in spectral analysis provides a promising new approach to the studies of the spatial and temporal variation of illumination in natural scenes and the understanding of color vision in natural environments.

Color↗

Accurate memory for colour but not pattern contrast in chicks.

The visual displays of animals and plants often look dramatic and colourful to us, but what information do they convey to their intended, non-human, audience [1] [2]? One possibility is that stimulus values are judged accurately - so, for example, a female might choose a suitor if he displays a specific colour [3]. Alternatively, as for human advertising, displays may attract attention without giving information, perhaps by exploiting innate preferences for bright colours or symmetry [2] [4] [5]. To address this issue experimentally, we investigated chicks' memories of visual patterns. Food was placed in patterned paper containers which, like seed pods or insect prey, must be manipulated to extract food and their patterns learnt. To establish what was learnt, birds were tested on familiar stimuli and on alternative stimuli of differing colour or contrast. For colour, birds selected the trained stimulus; for contrast, they preferred high contrast patterns over the familiar. These differing responses to colour and contrast show how separate components of display patterns could serve different roles, with colour being judged accurately whereas pattern contrast attracts attention.

Animals↗

Colour vision of domestic chicks.

The colour vision of domestic chicks (Gallus gallus) was investigated by training them to small food containers decorated with tilings of grey and coloured rectangles. Chicks learn to recognise the colour quickly and accurately. Chicks have four types of single-cone photoreceptor sensitive to ultraviolet, short-, medium- or long-wavelength light. To establish how these receptors are used for colour vision, stimuli were designed to be distinguished only by specific combinations of receptors. We infer (1) that all four single cones are used, and (2) that their outputs are encoded by at least three opponency mechanisms: one comparing the outputs of ultraviolet- and short-wavelength-sensitive receptors, one comparing the outputs of medium- and long-wavelength receptors and a third comparing of the outputs of short- and long- and/or medium-wavelength receptors. Thus, the chicks have tetrachromatic colour vision. These experiments do not exclude a role for the fifth cone type, double cones, but other evidence suggests that these cones serve luminance-based tasks, such as motion detection, and not colour recognition.

Animals↗

Receptor noise as a determinant of colour thresholds.

Inferences about mechanisms at one particular stage of a visual pathway may be made from psychophysical thresholds only if the noise at the stage in question dominates that in the others. Spectral sensitivities, measured under bright conditions, for di-, tri-, and tetrachromatic eyes from a range of animals can be modelled by assuming that thresholds are set by colour opponency mechanisms whose performance is limited by photoreceptor noise, the achromatic signal being disregarded. Noise in the opponency channels themselves is therefore not statistically independent, and it is not possible to infer anything more about the channels from psychophysical thresholds. As well as giving insight into mechanisms of vision, the model predicts the performance of colour vision in animals where physiological and anatomical data on the eye are available, but there are no direct measurements of perceptual thresholds. The model, therefore, is widely applicable to comparative studies of eye design and visual ecology.

Animals↗

Tetrachromacy, oil droplets and bird plumage colours.

There is a growing body of data on avian eyes, including measurements of visual pigment and oil droplet spectral absorption, and of receptor densities and their distributions across the retina. These data are sufficient to predict psychophysical colour discrimination thresholds for light-adapted eyes, and hence provide a basis for relating eye design to visual needs. We examine the advantages of coloured oil droplets, UV vision and tetrachromacy for discriminating a diverse set of avian plumage spectra under natural illumination. Discriminability is enhanced both by tetrachromacy and coloured oil droplets. Oil droplets may also improve colour constancy. Comparison of the performance of a pigeon's eye, where the shortest wavelength receptor peak is at 410 nm, with that of the passerine Leiothrix, where the ultraviolet-sensitive peak is at 365 nm, generally shows a small advantage to the latter, but this advantage depends critically on the noise level in the sensitivity mechanism and on the set of spectra being viewed.

Adaptation, Ocular↗

Metric analysis of threshold spectral sensitivity in the honeybee.

Behavioral spectral sensitivity curves are frequently used to characterize peripheral stages of visual processing. We test specific hypotheses about the physiology underlying honeybee spectral sensitivity by approximating published sensitivity curves with several metric models. The analysis shows that: (1) models assuming no interactions between different receptor types do not explain the behavioral data. Similarly, neither simple luminance mechanism models (sum of receptor excitations), nor models in which only the most sensitive receptor determines sensitivity fit the data. (2) The minimum number of postreceptoral mechanisms mediating discrimination is two. (3) Both mechanisms are of the chromatic type. Adding an achromatic mechanism decreases the accuracy of approximation.

Animals↗

Colour vision as an adaptation to frugivory in primates.

Most mammals possess two classes of cone, sensitive to short and to long wavelengths of light, but Old World primates (Catarrhini) have distinct medium and long wavelength sensitive classes. The sensitivities of these cones photopigments are alike in all catarrhines with peaks at about 440 nm ('blue'), 533 nm ('green') and 565 nm ('red'). One possible reason for the evolution and conservatism of catarrhine trichromacy is that colour vision is a specialization for finding food. A model of retinal coding of natural spectra, based on discrimination thresholds, is used to examine the usefulness of dichromatic and trichromatic vision for finding fruit, and for identifying fruit and leaves by colour. For identification tasks the dichromat's eye is almost as good as a trichromat's, but the trichromat has an advantage for detecting fruit against a background of leaves.

Adaptation, Physiological↗