Avian Retinal Carotenoid Accumulation: Ecophysiological Constraints and Behavioral Consequences

abstract: The elaborate signals of animals are often costly to produce and maintain, thus communicating reliable information about the quality of an individual to potential mates or competitors. The properties of the sensory systems that receive signals can drive the evolution of these signals and shape their form and function. However, relatively little is known about the ecological and physiological constraints that may influence the development and maintenance of sensory systems. In the house finch (Carpodacus mexicanus) and many other bird species, carotenoid pigments are used to create colorful sexually selected displays, and their expression is limited by health and dietary access to carotenoids. Carotenoids also accumulate in the avian retina, protecting it from photodamage and tuning color vision. Analogous to plumage carotenoid accumulation, I hypothesized that avian vision is subject to environmental and physiological constraints imposed by the acquisition and allocation of carotenoids. To test this hypothesis, I carried out a series of field and captive studies of the house finch to assess natural variation in and correlates of retinal carotenoid accumulation and to experimentally investigate the effects of dietary carotenoid availability, immune activation, and light exposure on retinal carotenoid accumulation. Moreover, through dietary manipulations of retinal carotenoid accumulation, I tested the impacts of carotenoid accumulation on visually mediated foraging and mate choice behaviors. My results indicate that avian retinal carotenoid accumulation is variable and significantly influenced by dietary carotenoid availability and immune system activity. Behavioral studies suggest that retinal carotenoid accumulation influences visual foraging performance and mediates a trade-off between color discrimination and photoreceptor sensitivity under dim-light conditions. Retinal accumulation did not influence female choice for male carotenoid-based coloration, indicating that a direct link between retinal accumulation and sexual selection for coloration is unlikely. However, retinal carotenoid accumulation in males was positively correlated with their plumage coloration. Thus, carotenoid-mediated visual health and performance or may be part of the information encoded in sexually selected coloration.Dissertation/ThesisPh.D. Biology 201

Perception-Aware Optimisation Methodologies for Quantum Dot Based Displays and Lighting

Human colour vision acuity is limited. This presents opportunities to leverage these perceptual limits to achieve engineering optimisations for devices and systems that interact with the human vision system. This dissertation presents the results of few investigations we carried out into quantifying these limits and several optimisation methodologies that we devised. The first step in this process is to quantify the acuity of human colour vision. We obtained a large corpus of colour matching data from a mobile video game called Specimen. We examine what questions about human vision this dataset allows us to answer and explore global statistics about colour vision based on this data on 41,000 players from 175 countries. We show that we can use the information in this dataset to infer potential candidate functions for the spectral sensitivities of each person in the dataset. The human eye acts like a many to one function; quantifiably different spectra can look like the same colour. This is referred to as metamerism. From a device perspective, different spectra consume different amounts of energy to generate. We show that we can use these two properties to elicit the same colour sensation using less energy. In the colour samples we evaluated, we show that we can achieve up to 10 times less power consumption while achieving a colour match. Given that one cannot change the emission spectrum of a display after fabrication, we propose the use of a multi-primary colour display to achieve this. We present two indices for quantifying the metameric capacity of such a display and its ability to save energy. The emission spectrum of a quantum dot (QD) based device is very narrow. Previous work in the literature suggested that narrow bandwidth spectra can lead to observer metameric breakdown; different observers disagreeing on the perceived ‘colour’ of a spectrum. We show that this might not be the case, using modern colour science tools, and show how metameric breakdown in a display could be minimised by carefully choosing the primary emission wavelengths. The limited colour acuity of human vision implies that people cannot notice small differences in colour. This fact has been used to create approximate colour transformation algorithms that subtly change colours in images such that they consume less energy when displayed on an emissive pixel display without causing unacceptable visual artefacts. We conducted a user study to gather information about the effect of one such colour transform called Crayon. We present a method for effectively picking the optimal transform parameters for Crayon, based on the user study results. The method presented calculates these parameters based on the properties of the image being transformed such that the power saving can be maximised while minimising the loss of image quality. The user study results show that we can achieve up to 50% power saving with a majority of the study participants reporting a negligible degradation in image quality in the transformed images. We additionally investigate a hypothesis that was presented stating that images with large amounts of highly luminous pixels cause increased power consumption in OLED displays due to localised display heating. We show that this hypothesis is wrong. We also investigate if sub-pixel rendering in Pentile displays can be used to reduce display power consumption by intentionally turning off random sub-pixels. However, we present a negative result showing that even single-pixel artefacts are observable on the test platform and thus, this cannot be used to improve display power efficiency. The narrow-band optical emissions of QD based devices mixed with their ability to be fabricated through solution processing can be used to mix multiple QDs together to build devices that generate arbitrary spectral shapes. We show how to use this property in an numerical optimisation based design framework to create lighting devices with a high colour rendering index (CRI). We evaluate the effects of different cost functions and initialisation strategies, and show that, we are able to design devices with a CRI > 96 using only four different QD primaries. We use a charge-transport based simulator to asses the electric properties of the designed devices. We also showcase initial work done on a modular software interface and a material library we developed for this simulator.EPSRC DTP studentship award RG84040:EP/N509620/

TOWARDS A COMPUTATIONAL MODEL OF RETINAL STRUCTURE AND BEHAVIOR

Human vision is our most important sensory system, allowing us to perceive our surroundings. It is an extremely complex process that starts with light entering the eye and ends inside of the brain, with most of its mechanisms still to be explained. When we observe a scene, the optics of the eye focus an image on the retina, where light signals are processed and sent all the way to the visual cortex of the brain, enabling our visual sensation. The progress of retinal research, especially on the topography of photoreceptors, is often tied to the progress of retinal imaging systems. The latest adaptive optics techniques have been essential for the study of the photoreceptors and their spatial characteristics, leading to discoveries that challenge the existing theories on color sensation. The organization of the retina is associated with various perceptive phenomena, some of them are straightforward and strictly related to visual performance like visual acuity or contrast sensitivity, but some of them are more difficult to analyze and test and can be related to the submosaics of the three classes of cone photoreceptors, like how the huge interpersonal differences between the ratio of different cone classes result in negligible differences in color sensation, suggesting the presence of compensation mechanisms in some stage of the visual system. In this dissertation will be discussed and addressed issues regarding the spatial organization of the photoreceptors in the human retina. A computational model has been developed, organized into a modular pipeline of extensible methods each simulating a different stage of visual processing. It does so by creating a model of spatial distribution of cones inside of a retina, then applying descriptive statistics for each photoreceptor to contribute to the creation of a graphical representation, based on a behavioral model that determines the absorption of photoreceptors. These apparent color stimuli are reconstructed in a representation of the observed scene. The model allows the testing of different parameters regulating the photoreceptor's topography, in order to formulate hypothesis on the perceptual differences arising from variations in spatial organization

Optimising Light Source Spectrum to Reduce the Energy Absorbed by Objects

Light is used to illuminate objects in the built environment. Humans can only observe light reflected from an object. Light absorbed by an object turns into heat and does not contribute to visibility. Since the spectral output of the new lighting technologies can be tuned, it is possible to imagine a lighting system that detects the colours of objects and emits customised light to minimise the absorbed energy. Previous optimisation studies investigated the use of narrowband LEDs to maximise the efficiency and colour quality of a light source. While these studies aimed to tune a white light source for general use, the lighting system proposed here minimises the energy consumed by lighting by detecting colours of objects and emitting customised light onto each coloured part of the object. This thesis investigates the feasibility of absorption-minimising light source spectra and their impact on the colour appearance of objects and energy consumption. Two computational studies were undertaken to form the theoretical basis of the absorption-minimising light source spectra. Computational simulations show that the theoretical single-peak spectra can lower the energy consumption up to around 38 % to 62 %, and double-peak test spectra can result in energy savings up to 71 %, without causing colour shifts. In these studies, standard reference illuminants, theoretical test spectra and coloured test samples were used. These studies are followed by the empirical evidence collected from two psychophysical experiments. Data from the experiments show that observers find the colour appearance of objects equally natural and attractive under spectrally optimised spectra and reference white light sources. An increased colour difference, to a certain extent, is found acceptable, which allows even higher energy savings. However, the translucent nature of some objects may negatively affect the results

Fundus-controlled perimetry (microperimetry): Application as outcome measure in clinical trials

YesFundus-controlled perimetry (FCP, also called 'microperimetry') allows for spatially-resolved mapping of visual sensitivity and measurement of fixation stability, both in clinical practice as well as research. The accurate spatial characterization of visual function enabled by FCP can provide insightful information about disease severity and progression not reflected by best-corrected visual acuity in a large range of disorders. This is especially important for monitoring of retinal diseases that initially spare the central retina in earlier disease stages. Improved intra- and inter-session retest-variability through fundus-tracking and precise point-wise follow-up examinations even in patients with unstable fixation represent key advantages of these technique. The design of disease-specific test patterns and protocols reduces the burden of extensive and time-consuming FCP testing, permitting a more meaningful and focused application. Recent developments also allow for photoreceptor-specific testing through implementation of dark-adapted chromatic and photopic testing. A detailed understanding of the variety of available devices and test settings is a key prerequisite for the design and optimization of FCP protocols in future natural history studies and clinical trials. Accordingly, this review describes the theoretical and technical background of FCP, its prior application in clinical and research settings, data that qualify the application of FCP as an outcome measure in clinical trials as well as ongoing and future developments

Doctor of Philosophy

dissertationIt is imperative to obtain a complete network graph of at least one representative retina if we are to fully understand vertebrate vision. Synaptic connectomics endeavors to construct such graphs. Though previously prevented by hardware and software limitations, the creation of customized viewing and analysis software, affordable data storage, and advances in electron imaging platform control now permit connectome assembly and analysis. The optimal strategy for building complete connectomes utilizes automated transmission electron imaging with 2 nm or better resolution, molecular tags for cell identification, open access data volumes for navigation, and annotation with open source tools to build three-dimensional cell libraries, complete network diagrams, and connectivity databases. In a few years, the first retinal connectome analyses reveal that many well-studied cells participate in much richer networks than expected. Collectively, these results impel a refactoring of the inner plexiform layer, while providing proof of concept for connectomics as a game-changing approach for a new era of scientific discovery

The Land Experiments in Colour Vision - Colour as a Physical, Phenomenological and Synthetic Object

This thesis analyses the historical and intellectual context of Edwin Land’s experiments in colour vision. I argue that the colour vision research program and retinex theory developed by Land and his colleagues provided a satisfying synthesis of two divergent schools in the history of colour science. The first chapter of this thesis establishes the existence of the “physical” school of colour science. The defining feature of this school was the belief in the colour atomism hypothesis. This is the idea that the colour perceived at a point in the visual field is completely determined by the physical properties of the light rays entering the retina at that point. In other words, there is a one-to-one correspondence between the physical properties of light rays and colour sensation at a point in the visual field. The second chapter establishes the existence of the “phenomenological” school of colour science. The defining feature of this school was the discovery of colour phenomena which could not be accounted for by the colour atomism hypothesis. Among these phenomena were “coloured shadows”, “simultaneous colour contrast”, and “colour constancy”. The third chapter shows how Land’s colour vision research program and retinex theory reconciled these two schools. Land and his colleagues demonstrated that the colour atomism hypothesis is a special case, valid only for points of light. The colour phenomena studied by the “phenomenological” school could be predicted by a computational model – retinex theory – which accounted for colour as it is perceived over a wide visual field, rather than simply at single points. In this process, Land and colleagues built up a new understanding of colour vision as a practical utility evolved for the organism, designed to achieve colour constancy