The influence of olfaction on the perception of high-fidelity computer graphics

The computer graphics industry is constantly demanding more realistic images and animations. However, producing such high quality scenes can take a long time, even days, if rendering on a single PC. One of the approaches that can be used to speed up rendering times is Visual Perception, which exploits the limitations of the Human Visual System, since the viewers of the results will be humans. Although there is an increasing body of research into how haptics and sound may affect a viewer's perception in a virtual environment, the in uence of smell has been largely ignored. The aim of this thesis is to address this gap and make smell an integral part of multi-modal virtual environments. In this work, we have performed four major experiments, with a total of 840 participants. In the experiments we used still images and animations, related and unrelated smells and finally, a multi-modal environment was considered with smell, sound and temperature. Beside this, we also investigated how long it takes for an average person to adapt to smell and what affect there may be when performing a task in the presence of a smell. The results of this thesis clearly show that a smell present in the environment firstly affects the perception of object quality within a rendered image, and secondly, enables parts of the scene or the whole animation to be selectively rendered in high quality while the rest can be rendered in a lower quality without the viewer noticing the drop in quality. Such selective rendering in the presence of smell results in significant computational performance gains without any loss in the quality of the image or animations perceived by a viewer

Audio-visual-olfactory resource allocation for tri-modal virtual environments

© 2019 IEEE. Virtual Environments (VEs) provide the opportunity to simulate a wide range of applications, from training to entertainment, in a safe and controlled manner. For applications which require realistic representations of real world environments, the VEs need to provide multiple, physically accurate sensory stimuli. However, simulating all the senses that comprise the human sensory system (HSS) is a task that requires significant computational resources. Since it is intractable to deliver all senses at the highest quality, we propose a resource distribution scheme in order to achieve an optimal perceptual experience within the given computational budgets. This paper investigates resource balancing for multi-modal scenarios composed of aural, visual and olfactory stimuli. Three experimental studies were conducted. The first experiment identified perceptual boundaries for olfactory computation. In the second experiment, participants (N=25) were asked, across a fixed number of budgets (M=5), to identify what they perceived to be the best visual, acoustic and olfactory stimulus quality for a given computational budget. Results demonstrate that participants tend to prioritize visual quality compared to other sensory stimuli. However, as the budget size is increased, users prefer a balanced distribution of resources with an increased preference for having smell impulses in the VE. Based on the collected data, a quality prediction model is proposed and its accuracy is validated against previously unused budgets and an untested scenario in a third and final experiment

Halcyornis toliapicus (aves: Lower Eocene, England) indicates advanced neuromorphology in Mesozoic Neornithes

Our recent X-ray micro computer-tomographic (μCT) investigations of Prophaethon shrubsolei and Odontopteryx toliapica from the Lower Eocene London Clay Formation of England revealed the avian brain to have been essentially modern in form by 55 Ma, but that an important vision-related synapomorphy of living birds, the eminentia sagittalis of the telencephalon, was poorly developed. This evidence suggested that the feature probably appeared close to the end of the Mesozoic. Here we use μCT analysis to describe the endocranium of Halcyornis toliapicus, also from the London Clay Formation. The affinities of Halcyornis have been hotly debated, with the taxon referred to the Charadriiformes (Laridae), Coraciiformes (Alcedinidae, and its own family Halcyornithidae) and most recently that Halcyornithidae may be a possible senior synonym of Pseudasturidae (Pan-Psittaciformes). Unlike Prophaethon and Odontopteryx, the eminentia sagittalis of Halcyornis is strongly developed and comparable to that of living species. Like those London Clay taxa, the eminentia sagittalis occupies a rostral position on the telencephalon. The senses of Halcyornis appear to have been well developed. The length of the cochlear duct of the inner ear indicates a hearing sensitivity within the upper range of living species, and enlarged olfactory lobes suggest a reasonable reliance on sense of smell. The optic nerves were especially well developed which, together with the strong development of the eminentia sagittalis, indicates a high degree of visual specialization in Halcyornis. The advanced development of the eminentia sagittalis further supports a Mesozoic age for the appearance of this structure and associated neural architectural complexity found in extant Aves. The eminentia sagittalis of living Psittaciformes is situated caudally on the telencephalon, making a Pan-Psittaciformes relationship unlikely for Halcyorni

Was There a Sensory Trade-off in Primate Evolution? The Vomeronasal Groove as a Means of Understanding the Vomeronasal System in the Fossil Record.

Primates have remarkable visual adaptations compared to most other mammals, long explained as associated with a trade-off with olfaction (smell). However, as more information comes to light on the role of olfaction in primate behavior it becomes apparent that olfaction is not a trivial sense. Even humans use smell to communicate, albeit in subtle ways, and the olfactory systems of the lemurs and lorises are very well-developed. Olfaction, however, is actually comprised of two distinct systems - the main olfactory and vomeronasal systems. These two systems overlap in many functions, but the main olfactory system is considered fairly generalized while the vomeronasal system is responsible for detecting odors specifically related to reproduction and predator avoidance. The vomeronasal system is incredibly variable in primates, being well-developed in the lemurs and lorises (strepsirhines) and absent in Old World monkeys and apes (catarrhines). Such variation does imply relaxed selection pressure to maintain a functional vomeronasal system in catarrhines, perhaps in response to gains in visual specialization. The goal of this dissertation is to investigate that evolutionary scenario using a multifaceted approach. A combined approach of comparing histology of the vomeronasal organ (the peripheral organ of the vomeronasal system) and computed tomography of the cranium is used to reveal variation in the vomeronasal organ across primates and to relate the soft-tissue organ to hard-tissue correlates. Indeed, the cartilage that surrounds the soft-tissue vomeronasal organ leaves a distinct impression on the nasal floor, which is here termed the vomeronasal groove . To assess the utility of inferring biological function from gross dimensions of the vomeronasal organ and its groove, vomeronasal organ length is compared to the number of genes underlying vomeronasal olfaction. To test whether or not the main olfactory system is evolving in tandem with the vomeronasal system, a hard-tissue correlate of the main olfaction (area of the cribriform plate) is compared to the number of genes encoding main olfaction. Results indicate that main olfaction and vomeronasal olfaction are affected by evolution differently and that vomeronasal organ length when adjusted for body size has a strong statistical relationship with the proportion of functional vomeronasal receptor genes across mammals. To test whether or not phylogenetic history, ecology, and reproduction strategies affect the evolution of the vomeronasal organ in primates, size-adjusted vomeronasal groove length is compared across related categories. Mating categories, probably reflecting sexual selection, appear to drive variation in vomeronasal groove length in lemurs and lorises, while color vision phenotypes appear to drive variation in the tarsiers, monkeys, and apes. The acquisition of trichromatic color vision in Old World monkeys and apes is associated with vomeronasal organ loss, but trichromatic color vision does not appear to be a primary driving force of vomeronasal organ reduction in other primates. The acquisition of high visual acuity, rather, appears to affect initial reduction in length of the vomeronasal groove in crown haplorhines. Fossils representing various stages of primate evolution show presence of the vomeronasal groove, and the presence of this groove in the recent ancestors of Old World monkeys and apes suggest that the vomeronasal organ was not lost until crown catarrhines (the group containing Old World monkeys and apes) diverged from all other primate lineages. High visual acuity, routine trichromatic color vision, environments with increased visibility, and changes in social dynamics could have shifted the way in which socio-sexual information was perceived in some primates, increasing the priority of visual and main olfactory signals over vomeronasal signals. Thus, a strict trade-off may not have occurred as much as a reallocation of sensory information from the vomeronasal system to vision and main olfaction

Olfaction in mosquitoes

Female mosquitoes are vectors of diseases, affecting both livestock and humans. The host-seeking and identification behaviors of mosquitoes are mediated mainly by olfactory cues. The peripheral olfactory organs of mosquitoes which perceive olfactory cues are the antennae and maxillary palps. These appendages bear numerous hair shaped structures, sensilla, in which olfactory receptor neurons (ORNs) are housed. The ORNs detect and discriminate various odorant molecules and send information regarding odor quality, quantity and spatio-temporal patterns to the central olfactory system in the brain for further analysis. The first goal of this study was to investigate the neuroanatomy of the mosquito central olfactory system. Using different staining techniques, the neuronal architecture of the deutocerebrum as well as 3D reconstructions of antennal lobe (AL) glomeruli were depicted for both sexes of the Afrcian malaria mosquito, Anopheles gambiae and the yellow fever mosquito, Aedes aegypti. To study how mosquitoes detect olfactory cues, single sensillum recordings (SSRs) were performed, which allowed me to investigate electrophysiological properties of individual ORNs housed in four morphological types of the most abundant olfactory sensilla, s. trichodea. I was able to identify 11 functional types which their ORNs displayed distinct responses to a set of compounds. As part of this study, axons of functionally defined ORNs were traced by neurobiotin to indicate which glomeruli they targeted. This resulted in a functional map of AL glomeruli. The map indicated that different functional types of ORNs converged onto different spatially fixed glomeruli. My next step was to identify novel biologically active compounds for the ORNs using gas chromatography coupled SSRs (GC-SSRs). Headspace odors from different human body parts, i.e. armpit, feet and trunk regions as well as from a plant used as a mosquito repellent (Nepeta faassenii) were collected, extracted and eventually injected onto the GC-column. I found that some of the extract components elicited responses in previously defined ORNs as well as in ORNs of the intermediate sensilla. Some of the compounds, which were subsequently identified by using GC-mass spectrometry (GC-MS) were heptanal, octanal, nonanal and decanal

Introduction to research topic: attention and consciousness in different senses

The question of the origin of consciousness has engaged scientists and philosophers for centuries. Early scholars relied on introspection, leading some to conclude that attention is necessary for consciousness, and in some cases equating attention and consciousness. Such a tight relationship between attention and consciousness has also been proposed by many modern theorists (Posner, 1994; Merikle and Joordens, 1997; Mack and Rock, 1998; Chun and Wolfe, 2000; O'Regan and Noe, 2001; Mole, 2008; De Brigard and Prinz, 2010; Prinz, 2011; Cohen et al., 2012). The relationship between attention and consciousness has come under increasing scrutiny with the development of neuroscientific methods. In modern neuroscience, the effects of attention are often objectively defined and measured as reduced reaction time and improved performance. Similarly, conscious awareness of an object is established by a subjective report in combination with objective forced-choice performance (Seth et al., 2008; Sandberg et al., 2011). With these measures in place, a variety of methods has been used to manipulate attention (e.g., cueing, divided attention, etc.) and consciousness [e.g., masking, crowding, and binocular rivalry (Kim and Blake, 2005)]. These empirical studies have culminated in recent proposals that attention and consciousness are supported by different neuronal processes and they are not necessarily correlated all the time (Iwasaki, 1993; Baars, 1997; Hardcastle, 1997; Kentridge et al., 1999; Naccache et al., 2002; Lamme, 2003; Woodman and Luck, 2003; Bachmann, 2006; Koch and Tsuchiya, 2007; van Boxtel et al., 2010). Our original motivation to edit this Research Topic was threefold: (1) to gather and collect current, diverse views on the relationship between consciousness and attention, (2) to invite reviews on consciousness and attention in non-vision modalities, (3) and to invite empirical studies of consciousness and attention. As summarized below, our goals are largely achieved thanks to 17 contributions to this issue

The Murine Accessory Olfactory Bulb as a Model Chemosensory System: Experimental and Computational Analysis of Chemosensory Representations

A common challenge across sensory processing modalities is forming meaningful associations between the neural responses and the outside world. These neural representations of the world must then be integrated across different sensory systems contributing to each individuals perceptual experience. While there has been considerable study of sensory representations in the visual system of humans and multiple model organisms, other sensory domains, including olfaction, are less well understood. In this thesis, I set out to better understand the sensory representations of the mouse accessory olfactory system (AOS), a part of the olfactory system. The mouse AOS, our model chemosensory system, comprises peripheral vomeronasal sensory neurons (VSNs), the accessory olfactory bulb (AOB), and downstream effectors. Our work describes the neural representations of multiple sensory inputs in the AOS, specifically the representations of odorants in high dimensional chemical sensory space in the AOB, and how these representations are shaped by interactions within the circuit. Given the complex nature of olfactory chemosensory representations, the features of our model system may give new perspectives on the neural representation of the outside world. In a neural representation of olfactory information, both the interactions between each receptor and odor compounds as well as the circuit mediated interactions could potentially affect the neural representations of the outside world. The initial neural response comprises component interactions between each receptor and the odor; chemical signals must interact with physical receptors. However, chemosensory processing, such as olfaction, requires interpreting a large variety of potentially overlapping chemical cues from the environment with only a finite number of receptor types. This means that each chemical cue does not necessarily activate only one receptor type or region of the circuit, but rather the cue is likely to be represented by multiple receptor and odor component interactions. Also, the component parts of odors may be processed differently when presented in isolation versus in a more complex mixture, thus allowing the response to a particular odor to vary with chemical context. Moreover, once these component representations exist, interactions within the neural circuit may further shape these responses. For example, one might expect component parts of a complex odor to specifically inhibit other component parts. In the case of the accessory olfactory system this inhibition could be at the receptor level or at the level of the sensory representation in the accessory olfactory bulb (AOB). In Chapter 3, I describe the overall organization of chemosensory representations in the accessory olfactory bulb (AOB), which is found to be a modular map in which the primary associations of functional sensory responses are spatially organized relative to one another. I find these primary associations are condensations of the first order sensory neuron axon terminals, which form population response pooling structures called glomeruli. In these glomeruli, similar response types from those sensory neurons expressing one of the approximately 300 receptor types in the vomeronasal organ (VNO) co-converge. One purpose of converging inputs of neurons expressing the same receptor is likely to minimize noise, and I demonstrate that pooling of like receptor responses into glomeruli does increase neural signal relative to noise. However, I also observed a modular organization among and between glomeruli in which certain types or patterns of chemosensory responses are always spatially adjacent to one another, while others are much farther apart than would be expected by chance. I found this spatial modularity for both ethological stimuli (urine collected from conspecifics with widely divergent physiological endocrine status) and individual sulfated steroids. In Chapter 4, I explore the consequences of changing sensory context, specifically the presentation of multiple compounds, and the role that inhibition plays in the neural representation of the sensory stimuli. First, I tested whether the circuit responds differently to demands to represent a single odor than to demands to represent multiple odors by using odors that activate glomeruli both inside and outside of modules. I found that responses to mixtures rapidly diverge from the responses of individual component parts. Moreover, there was an effect of inhibition in modulating the response to preferred stimuli in all glomeruli. However, initial analysis of one type of pregnanolone responsive glomeruli demonstrated that the divergent response to mixtures in this type of glomerulus was not mediated by inhibition at the glomerular level, but was rather attributable to bottom-up effects from the interactions of multiple ligands with chemosensory receptors in the VNO. Nonetheless, I also demonstrated that in the AOB, the axon terminals of the same sensory neurons (glomeruli) are organized into modules that allow for feedback inhibition. Significant ionotropic glutamate receptor signal modulation was observed within modules, demonstrating that there are inhibition mediated effects in the representation of complex mixtures when glomeruli are co-locally arranged. Specifically, at both the level of the VSNs and also in AOB glomeruli, the response to allopregnanolone sulfate is inhibited by co-presentation with estradiol sulfate. This both significantly increases the relative representation of estradiol sulfate and shifts representation of allopregnanolone primarily within modules. These types of context dependent interactions depend on the spatial organization described in Chapter 3 as well as mixture context, and have the potential to optimize the representation of some chemical cues in a context specific manner

Effect of Olfaction on the Perception of Movie Clips

Odours are important to many species but their effect on human perception in the context of concurrent auditory and visual stimulation has received little investigation. Here we examined how the experience of viewing audio-visual movie clips changes when accompanied by congruent or incongruent odours. Using an olfactometer to control odourant delivery, thirty-five undergraduate students from Western University were randomly presented 36 different odour-video pairs twice. Following each presentation, participants completed three Likert scales to assess multisensory interaction in terms of engagement, pleasantness, and emotional arousal. Comparison of congruent and incongruent odours to the no odour control condition revealed that incongruent odours had a greater effect than congruent odours on participant ratings, and that this effect acted to negatively influence experience, reducing engagement, pleasantness, and emotional arousal. There was little difference between congruent odours and no odour on ratings of engagement and emotional arousal; however, even congruent odours reduced pleasantness ratings, suggesting all odours used were, to an extent, unpleasant. An interaction suggested that certain movies were more strongly modulated by odour than others. We interpret our results as evidence of crossmodal competition, in which the presence of an odour leads to suppression of the auditory and visual modalities. This was confirmed using functional magnetic resonance imaging in a single participant. Future research should continue to investigate the surprising role odour plays in multisensory interaction

Cortical network and connectivity underlying hedonic olfactory perception

Objective. The emotional response to olfactory stimuli implies the activation of a complex cascade of events triggered by structures lying in the limbic system. However, little is known about how this activation is projected up to cerebral cortex and how different cortical areas dynamically interact each other. Approach. In this study, we acquired EEG from human participants performing a passive odor-perception task with odorants conveying positive, neutral and negative valence. A novel methodological pipeline integrating global field power (GFP), independent component analysis (ICA), dipole source localization was applied to estimate effective connectivity in the challenging scenario of single-trial low-synchronized stimulation. Main results. We identified the brain network and the neural paths, elicited at different frequency bands, i.e. θ (4-7Hz), α (8-12Hz) and β (13-30Hz), involved in odor valence processing. This brain network includes the orbitofrontal cortex (OFC), the cingulate gyrus (CgG), the superior temporal gyrus (STG), the posterior cingulate cortex/precuneus (PCC/PCu) and the parahippocampal gyrus (PHG). It was analyzed using a time-varying multivariate autoregressive model to resolve time-frequency causal interactions. Specifically, the OFC acts as the main node for odor perception and evaluation of pleasant and unpleasant stimuli, whereas no specific path was observed for a neutral stimulus. Significance. The results introduce new evidences on the role of the OFC during hedonic perception and underpin its specificity during the odor valence assessment. Our findings suggest that, after the odor onset different, bidirectional interactions occur between the OFC and other brain regions associated with emotion recognition/categorization and memory according to the stimulus valence. This outcome unveils how the hedonic olfactory network dynamically changes based on odor valence