Colour Vision in Birds : Comparing behavioural thresholds and model predictions

Birds use colour vision for many biologically relevant behaviours such as foraging and mate choice. Bird colour vision is mediated by four types of single cones, giving them an extra dimension of colour information compared to trichromatic humans. The cone photoreceptors of birds have coloured oil droplets that are assumed to increase the discriminability of colours in bright light at the cost of dim light sensitivity. In this thesis I present four studies where we have trained chickens to perform colour discrimination and tested the limits of their behavioural performance. In paper I we tested how small colour differences chickens can discriminate. This allowed us to test the predictions of the most well established model for bird colour vision, the receptor noise limited model. There was a reasonably good fit between model and behaviour. Furthermore, we tested in how dim light chickens could discriminate colours and found that the intensity threshold was affected by the colour difference between the stimuli and their intensity. In Paper II we continued testing colour discrimination in dim light and tested the hypothesis that chickens sum the signals from many photoreceptors to increase contrast sensitivity at the cost of spatial resolution in dim light, so called spatial pooling. We used food containers covered with larger, smaller, more or fewer colour patches. Supporting the hypothesis, the containers covered by more colour could be discriminated in dimmer light. In Paper III we tested colour constancy, the ability to maintain colour perception in different spectral illuminations that would otherwise confuse colour perception. Our aim was to find the largest illumination change that chicken colour constancy could tolerate. We found that chicken colour constancy could tolerate larger illumination changes when discriminating stimuli that were more different from each other. In paper IV we continued the work on colour constancy but allowed the chickens to use relative colour learning, which was specifically excluded in paper III. In Paper IV we found that their colour constancy could tolerate larger illumination changes. In nature relative colour cues are available and may be an important aspect of colour learning and perception. These results suggest that such cues can make colour constancy more robust to larger illumination changes. In both experiments chicken colour constancy was improved if they were adapted for 5 minutes in the tested illumination before performing the discrimination task. We compared the illuminations for which chickens retained colour constancy, to the difference between natural illuminations and we can conclude that chickens are well equipped to maintain accurate colour perception when changing between habitats in the wild. Objects are detected both by their chromatic and achromatic contrasts. The receptor noise limited model can be used to predict discriminability through both chromatic and achromatic vision. To use the model reliably its assumptions and predictions must be compared to behavioural results. This has been done for the chromatic version of the model but not the achromatic. In Paper V we compiled all known chromatic and achromatic contrast detection thresholds, and used them to derive the limiting noise level to be used when predicting visual discrimination in a range of animals. We discuss the limitations of using modelling in the wild such as the need to consider the spatial pattern of the stimuli and the light intensities in which the modelling occurs

Factors affecting brightness and colour vision under water

Both theoretical and practical importance can be attached to attempts to model human threshold and supra-threshold visual performance under water. Previously, emphasis has been given to the integration of visual data from experiments conducted in air with data of the physical specification of the underwater light field. However, too few underwater studies have been undertaken for the validity of this approach to be assessed. The present research therefore was concerned with the acquisition of such data. Four experiments were carried out: (a) to compare the predicted and obtained detection thresholds of achromatic targets, (b) to measure the relative recognition thresholds of coloured targets, (c) to compare the predicted and obtained supra-threshold appearance of coloured targets at various viewing distances and under different experimental instructions, (d) to compare the predicted and obtained detection thresholds for achromatic targets under realistic search conditions. Within each experiment, observers were tested on visual tasks in the field and in laboratory simulations. Physical specifications of targets and backgrounds were determined by photometry and spectroradiometry. The data confirmed that: (a) erroneous predictions of the detection threshold could occur when the contributions of absorption and scattering to the attenuation of light were not differentiated, (b) the successful replication of previous findings for the relative recognition thresholds of colours depended on the brightness of the targets, (c) the perceived change in target colour with increasing viewing distance was less than that measured physically, implying the presence of a colour constancy mechanism other than chromatic adaptation and simultaneous colour contrast; the degree of colour constancy also varied with the type of target and experimental instructions, (d) the successful prediction of the effects of target-observer motion and target location uncertainty required more than simple numerical corrections to the basic detection threshold model. It was concluded that further progress in underwater visibility modelling is possible provided that the tendency to oversimplify human visual performance is suppressed

Illuminant Estimation by Voting

Obtaining an estimate of the illuminant color is an important component in many image analysis applications. Due to the complexity of the problem many restrictive assumptions are commonly applied, making the existing illuminant estimation methodologies not widely applicable on natural images. We propose a methodology which analyzes a large number of regions in an image. An illuminant estimate is obtained independently from each region and a global illumination color is computed by consensus. Each region itself is mainly composed by pixels which simultaneously exhibit both diffuse and specular reflection. This allows for a larger inclusion of pixels than purely specularitybased methods, while avoiding, at the same time, some of the restrictive assumptions of purely diffuse-based approaches. As such, our technique is particularly well-suited for analyzing real-world images. Experiments with laboratory data show that our methodology outperforms 75 % of other illuminant estimation methods. On natural images, the algorithm is very stable and provides qualitatively correct estimates. 1

Estimation of illuminants from color signals of illuminated objects

Color constancy is the ability of the human visual systems to discount the effect of the illumination and to assign approximate constant color descriptions to objects. This ability has long been studied and widely applied to many areas such as color reproduction and machine vision, especially with the development of digital color processing. This thesis work makes some improvements in illuminant estimation and computational color constancy based on the study and testing of existing algorithms. During recent years, it has been noticed that illuminant estimation based on gamut comparison is efficient and simple to implement. Although numerous investigations have been done in this field, there are still some deficiencies. A large part of this thesis has been work in the area of illuminant estimation through gamut comparison. Noting the importance of color lightness in gamut comparison, and also in order to simplify three-dimensional gamut calculation, a new illuminant estimation method is proposed through gamut comparison at separated lightness levels. Maximum color separation is a color constancy method which is based on the assumption that colors in a scene will obtain the largest gamut area under white illumination. The method was further derived and improved in this thesis to make it applicable and efficient. In addition, some intrinsic questions in gamut comparison methods, for example the relationship between the color space and the application of gamut or probability distribution, were investigated. Color constancy methods through spectral recovery have the limitation that there is no effective way to confine the range of object spectral reflectance. In this thesis, a new constraint on spectral reflectance based on the relative ratios of the parameters from principal component analysis (PCA) decomposition is proposed. The proposed constraint was applied to illuminant detection methods as a metric on the recovered spectral reflectance. Because of the importance of the sensor sensitivities and their wide variation, the influence from the sensor sensitivities on different kinds of illuminant estimation methods was also studied. Estimation method stability to wrong sensor information was tested, suggesting the possible solution to illuminant estimation on images with unknown sources. In addition, with the development of multi-channel imaging, some research on illuminant estimation for multi-channel images both on the correlated color temperature (CCT) estimation and the illuminant spectral recovery was performed in this thesis. All the improvement and new proposed methods in this thesis are tested and compared with those existing methods with best performance, both on synthetic data and real images. The comparison verified the high efficiency and implementation simplicity of the proposed methods

The time-course of colour vision

Four experiments are presented, each investigating temporal properties of colour vision processing in human observers. The first experiment replicates and extends an experiment by Stromeyer et al. (1991). We look for a phase difference between combined temporal modulations in orthogonal directions in colour space, which might null the often-claimed latency of signals originating from the short-wavelength sensitive cones (S-cones). We provide another estimate of the magnitude of this latency, and give evidence to suggest that it originates early in the chromatic pathway, before signals from S-cones are combined with those that receive opposed L- and M-cone input. In the second experiment we adapt observers to two stimuli that are matched in the mean and amplitude of modulation they offer to the cone classes and to the cardinal opponent mechanisms, but that differ in chromatic appearance, and hence their modulation of later colour mechanisms. Chromatic discrimination thresholds after adaptation to these two stimuli differ along intermediate directions in colour space, and we argue that these differences reveal the adaptation response of central colour mechanisms. In the third experiment we demonstrate similar adaptation using the same stimuli, measured with reaction times rather than thresholds. In the final experiment, we measure the degree to which colour constancy is achieved as a function of time in a simulated stimulus environment in which the illuminant changes periodically. We find that perfect constancy is not achieved instantaneously after an illuminant chromaticity shift and that constancy of colour appearance judgements increases over several seconds

Preferred lightness and chromatic image contrast reproduction

In this study, the image preference as a function of lightness and chromatic contrast of images produced on an ink-jet printer is examined. The purpose is to develop image manipulation rules, useful in the development of printer algorithms to produce images that are preferred by viewers over images that have been printed without application of these rules. Five images are used during the psychophysical experiment, two business graphics and three pictorial, processed in three different ways in RLAB color space, once having only the tightness contrast varied, then only the chromatic contrast, and finally both lightness and chromatic contrast varied. The results showed that for the graphics images seen without a CRT original used for comparison, the mean preference was an increase in lightness contrast, while with an original available for comparison, the mean preference indicated a decrease in both lightness and chromatic contrast. For pictorial images, in the first phase of the experiment the mean preference was an increase in both lightness and chromatic contrast, and after comparison, a decrease in lightness and simultaneous decrease in lightness and chromatic contrast are the most preferred

Metamer Mismatching in Practice versus Theory

Metamer mismatching (the phenomenon that two objects matching in color under one illuminant may not match under a different illuminant) potentially has important consequences for color perception. Logvinenko et al. [PLoS ONE 10, e0135029 (2015)] show that in theory the extent of metamer mismatching can be very significant. This paper examines metamer mismatching in practice by computing the volumes of the empirical metamer mismatch bodies and comparing them to the volumes of the theoretical mismatch bodies. A set of more than 25 million unique reflectance spectra is assembled using datasets from several sources. For a given color signal (e.g., CIE XYZ) recorded under a given first illuminant, its empirical metamer mismatch body for a change to a second illuminant is computed as follows: the reflectances having the same color signal when lit by the first illuminant (i.e., reflect metameric light) are computationally relit by the second illuminant, and the convex hull of the resulting color signals then defines the empirical metamer mismatch body. The volume of these bodies is shown to vary systematically with Munsell value and chroma. The empirical mismatch bodies are compared to the theoretical mismatch bodies computed using the algorithm of Logvinenko et al. [IEEE Trans. Image Process. 23, 34 (2014)]. There are three key findings: (1) the empirical bodies are found to be substantially smaller than the theoretical ones; (2) the sizes of both the empirical and theoretical bodies show a systematic variation with Munsell value and chroma; and (3) applied to the problem of color-signal prediction, the centroid of the empirical metamer mismatch body is shown to be a better predictor of what a given color signal might become under a specified illuminant than state-of-the-art methods

Colour constancy in dichromats and trichromats: dependence on task

An important topic in the field of colour vision is the impact of colour vision deficiencies on daily life tasks. Investigating the extent to which colour constancy (i.e. the ability to recognise surface colour under different illuminants) is preserved in colour vision- deficient observers can provide us with insight into the nature and function of trichromatic colour vision. The first chapter of this thesis provides a summary of the very basics of colour vision, colour vision deficiencies, as well as colour constancy. Studies conducted on the colour constancy abilities of colour-vision-deficient observers versus those with normal colour vision are reviewed. The second chapter presents and reports the aims and methods of the proposed experiment (which could not take place due to the COVID-19 pandemic). This experiment investigated the colour constancy abilities of trichromats versus dichromats using two different colour constancy tasks (2D achromatic adjustment vs. 3D blocks-copying/selection task) and aimed to show how colour constancy depends on observer type as well as task type. The third chapter comprises of a computerised simulation. This simulation aimed to model the colour constancy of “ideal” observers when presented with various surfaces and illuminants. These observers involve simulated normal trichromats, anomalous trichromats and dichromats. A variety of yellow, blue, green and red illuminant shifts (from neutral daylight) were used, and surface chromaticity and observer types were compared. Overall, whilst no three-way interaction between illuminant shift, surface chromaticity and observer type were found in the simulation, strong main effects were found. It is suggested that a combination of simulated and experimental research is needed to understand the colour constancy mechanisms underpinning dichromacy and trichromacy at multiple levels (cone-based, cognitive and computational)