Dichromatic Colour Vision in Wallabies as Characterised by Three Behavioural Paradigms

Despite lacking genetic evidence of a third cone opsin in the retina of any Australian marsupial, most species tested so far appear to be trichromatic. In the light of this, we have re-examined colour vision of the tammar wallaby which had previously been identified as a dichromat. Three different psychophysical tests, based on an operant conditioning paradigm, were used to confirm that colour perception in the wallaby can be predicted and conclusively explained by the existence of only two cone types. Firstly, colour-mixing experiments revealed a Confusion Point between the three primary colours of a LCD monitor that can be predicted by the cone excitation ratio of the short- and middle-wavelength sensitive cones. Secondly, the wavelength discrimination ability in the wallaby, when tested with monochromatic stimuli, was found to be limited to a narrow range between 440 nm and 500 nm. Lastly, an experiment designed to test the wallaby’s ability to discriminate monochromatic lights from a white light provided clear evidence for a Neutral Point around 485 nm where discrimination consistently failed. Relative colour discrimination seemed clearly preferred but it was possible to train a wallaby to perform absolute colour discriminations. The results confirm the tammar wallaby as a dichromat, and so far the only behaviourally confirmed dichromat among the Australian marsupials.     

On the neuronal basis of figure-ground discrimination by relative motion in the visual system of the fly

A fly can discriminate an object (figure) from its background on the basis of motion information alone. This information processing task has been analysed, so far, mainly in behavioural studies but also in electrophysiological experiments (Reichardt et al., 1983). The present study represents a further attempt to bridge the gap between the behavioural and the neuronal level. It is based on behavioural and electrophysiological experiments as well as on computer simulations. The characteristic properties of figureground discrimination behaviour impose specific constraints on the spatial integration properties of the output cells of the underlying neuronal network, the heterolateral interactions in their input circuitry, as well as on the range of variability of their response. These constraints are derived partly from previous behavioural studies (Reichardt et al., 1983), partly, however, from behavioural response characteristics which have not been addressed explicitly so far. They are interpreted in terms of one of the alternative model circuits shown by Reichardt et al. (1983) to be sufficient to account for figure-ground discrimination. It will be demonstrated, however, that this can be done equally well by means of a further alternative model circuit. These constraints are used in the electrophysiological analysis for establishing visual interneurones as output elements of the neuronal network underlying figure-ground discrimination.In the behavioural experiments on figure-ground discrimination as well as on the optomotor course control the yaw torque generated by the tethered flying fly under visual stimulation was used as a measure for the strength and time course of the reaction. Therefore, it has initially been proposed that the three Horizontal Cells, which are regarded as the output elements of the neuronal network underlying the optomotor reaction (e.g. Hausen, 1981), might also control yaw torque generation in figure-ground discrimination (Reichardt et al., 1983). New behavioural data show, however, that the Horizontal Cells do not meet all the constraints imposed on the presumed output cells of the figure-ground discrimination network: (1) The Horizontal Cells are not sensitive enough to motion of small objects. (2) The heterolateral interactions within their input circuitry are not in accordance with the behavioural data (see also Reichardt et al., 1983). (3) The variability found in the time course of certain components of the yaw torque response to relative motion of figure and ground cannot be explained by their response characteristics. Hence, the Horizontal Cells cannot account for figure-ground discrimination on their own and additional output cells of the optic lobes with different functional properties are required to accomplish this task.     

Virtual retinal display system with nodal point image of a laser beam: construction example and evaluation of subjective brightness perception]

We investigated the perception of brightness for red monochromatic laser light. For this purpose, a modified virtual retinal display (VRD) was constructed. The modification involved projecting the laser beam into the eye. In our VRD, the laser beam pivots in the nodal point of the eye (badal system). The displayed image therefore does not depend on the refractive state of the eye. Brightness perception was assessed by means of psychophysical experiments. The results of these experiments indicate that perception of brightness at 652 nm increases more rapidly with increasing physical stimulus than does the perception of white light (colour temperature 2'935 K). At a wavelength of 652 nm, an optical power of 79 nW is required to produce an image subtending 2 degrees of equal brightness, as is perceived with an image of about 2'900 cd/m2 subtending 2 degrees.     

Separate colour-opponent mechanisms underlie the detection and discrimination of moving chromatic targets.

Current opinion holds that human colour vision is mediated primarily via a colour-opponent pathway that carries information about both wavelength and luminance contrast (type I). However, some authors argue that chromatic sensitivity may be limited by a different geniculostriate pathway, which carries information about wavelength alone (type II). We provide psychophysical evidence that both pathways may contribute to the perception of moving, chromatic targets in humans, depending on the nature of the visual discrimination. In experiment 1, we show that adaptation to drifting, red-green stimuli causes reductions in contrast sensitivity for both the detection and direction discrimination of moving chromatic targets. Importantly, the effects of adaptation are not directionally specific. In experiment 2, we show that adaptation to luminance gratings results in reduced sensitivity for the direction discrimination, but not the detection of moving chromatic targets. We suggest that sensitivity for the direction discrimination of chromatic targets is limited by a colour-opponent pathway that also conveys luminance-contrast information, whereas the detection of such targets is limited by a pathway with access to colour information alone. The properties of these pathways are consistent with the known properties of type-I and type-II neurons of the primate parvocellular lateral geniculate nucleus and their cortical projections. These findings may explain the known differences between detection and direction discrimination thresholds for chromatic targets moving at low to moderate velocities.     

The asynchrony of consciousness.

We present below a simple hypothesis on what we believe is a characteristic of visual consciousness. It is derived from facts about the visual brain revealed in the past quarter of a century, but it relies most especially on psychophysical evidence which shows that different attributes of the visual scene are consciously perceived at different times. This temporal asynchrony in visual perception reveals, we believe, a plurality of visual consciousnesses that are asynchronous with respect to each other, reflecting the modular organization of the visual brain. We further hypothesize that when two attributes (e.g. colour and motion) are presented simultaneously, the activity of cells in a given processing system is sufficient to create a conscious experience of the corresponding attribute (e.g. colour), without the necessity for interaction with the activities of cells in other processing systems (e.g. motion). Thus, any binding of the activity of cells in different systems should be more properly thought of as a binding of the conscious experiences generated in each system.     

Exposure to Organic Solvents Used in Dry Cleaning Reduces Low and High Level Visual Function

PurposeTo investigate whether exposure to occupational levels of organic solvents in the dry cleaning industry is associated with neurotoxic symptoms and visual deficits in the perception of basic visual features such as luminance contrast and colour, higher level processing of global motion and form (Experiment 1), and cognitive function as measured in a visual search task (Experiment 2).MethodsThe Q16 neurotoxic questionnaire, a commonly used measure of neurotoxicity (by the World Health Organization), was administered to assess the neurotoxic status of a group of 33 dry cleaners exposed to occupational levels of organic solvents (OS) and 35 age-matched non dry-cleaners who had never worked in the dry cleaning industry. In Experiment 1, to assess visual function, contrast sensitivity, colour/hue discrimination (Munsell Hue 100 test), global motion and form thresholds were assessed using computerised psychophysical tests. Sensitivity to global motion or form structure was quantified by varying the pattern coherence of global dot motion (GDM) and Glass pattern (oriented dot pairs) respectively (i.e., the percentage of dots/dot pairs that contribute to the perception of global structure). In Experiment 2, a letter visual-search task was used to measure reaction times (as a function of the number of elements: 4, 8, 16, 32, 64 and 100) in both parallel and serial search conditions.ResultsDry cleaners exposed to organic solvents had significantly higher scores on the Q16 compared to non dry-cleaners indicating that dry cleaners experienced more neurotoxic symptoms on average. The contrast sensitivity function for dry cleaners was significantly lower at all spatial frequencies relative to non dry-cleaners, which is consistent with previous studies. Poorer colour discrimination performance was also noted in dry cleaners than non dry-cleaners, particularly along the blue/yellow axis. In a new finding, we report that global form and motion thresholds for dry cleaners were also significantly higher and almost double than that obtained from non dry-cleaners. However, reaction time performance on both parallel and serial visual search was not different between dry cleaners and non dry-cleaners.ConclusionsExposure to occupational levels of organic solvents is associated with neurotoxicity which is in turn associated with both low level deficits (such as the perception of contrast and discrimination of colour) and high level visual deficits such as the perception of global form and motion, but not visual search performance. The latter finding indicates that the deficits in visual function are unlikely to be due to changes in general cognitive performance.     

Simultaneous and successive colour discrimination in the honeybee (Apis mellifera)

The colour discrimination of individual free-flying honeybees (Apis mellifera) was tested with simultaneous and successive viewing conditions for a variety of broadband reflectance stimuli. For simultaneous viewing bees used form vision to discriminate patterned target stimuli from homogeneous coloured distractor stimuli, and for successive discrimination bees were required to discriminate between homogeneously coloured stimuli. Bees were significantly better at a simultaneous discrimination task, and we suggest this is explained by the inefficiency with which the bees brain can code and retrieve colour information from memory when viewing stimuli successively. Using simultaneous viewing conditions bees discriminated between the test stimuli at a level equivalent to 1 just-noticeable-difference for human colour vision. Discrimination of colours by bees with simultaneous viewing conditions exceeded previous estimates of what is possible considering models of photoreceptor noise measured in bees, which suggests spatial and/or temporal summation of colour signals for fine discrimination tasks. The results show that when behavioural experiments are used to collect data about the mechanisms facilitating colour discrimination in animals, it is important to consider the effects of the stimulus viewing conditions on results.     

A neural model of feature attention in motion perception

We utilize a model of motion perception to link a physiological study of feature attention in cortical motion processing to a psychophysical experiment of motion perception. We explain effects of feature attention by modulatory excitation of neural activity patterns in a framework of biased competition. Our model allows us to qualitatively replicate physiological data concerning attentional modulation and to generate model behavior in a decision experiment that is consistent with psychophysical observations. Furthermore, our investigation makes predictions for future psychophysical experiments.     

Flutter discrimination: neural codes,perception, memory and decision making

Recent studies combining psychophysical and neurophysiological experiments in behaving monkeys have provided new insights into how several cortical areas integrate efforts to solve a vibrotactile discrimination task. In particular, these studies have addressed how neural codes are related to perception, working memory and decision making in this model. The primary somatosensory cortex drives higher cortical areas where past and current sensory information are combined, such that a comparison of the two evolves into a behavioural decision. These and other observations in visual tasks indicate that decisions emerge from highly-distributed processes in which the details of a scheduled motor plan are gradually specified by sensory information.     

A theory for the use of visual orientation information which exploits the columnar structure of striate cortex

A neural model is constructed based on the structure of a visual orientation hypercolumn in mammalian striate cortex. It is then assumed that the perceived orientation of visual contours is determined by the pattern of neuronal activity across orientation columns. Using statistical estimation theory, limits on the precision of orientation estimation and discrimination are calculated. These limits are functions of single unit response properties such as orientation tuning width, response amplitude and response variability, as well as the degree of organization in the neural network. It is shown that a network of modest size, consisting of broadly orientation selective units, can reliably discriminate orientation with a precision equivalent to human performance. Of the various network parameters, the discrimination threshold depends most critically on the number of cells in the hypercolumn. The form of the dependence on cell number correctly predicts the results of psychophysical studies of orientation discrimination. The model system's performance is also consistent with psychophysical data in two situations in which human performance is not optimal. First, interference with orientation discrimination occurs when multiple stimuli activate cells in the same hypercolumn. Second, systematic errors in the estimation of orientation can occur when a stimulus is composed of intersecting lines. The results demonstrate that it is possible to relate neural activity to visual performance by an examination of the pattern of activity across orientation columns. This provides support for the hypothesis that perceived orientation is determined by the distributed pattern of neural activity. The results also encourage the view of neural activity. The results also are determined by the responses of many neurons rather than the sensitivity of individual cells.     

Objective and subjective indexes of auditory motion processing

The objective and subjective indexes of sound stimulus discrimination have been studied in order to get insight into individual stages of signal processing in the human brain. The experiment employed two methods: electrophysiological (mismatch negativity or MMN recording) and psychophysical (two-alternative forced choice). Two types of spatial sound stimuli simulated gradual and abrupt sound motion from the head midline. The subjective discrimination between the gradual and abrupt motions was estimated as a function of the stimulus trajectory length. MMN as an objective index of spatial discrimination has been obtained in response to the subthreshold and the suprathreshold levels of psychophysical discrimination. An increase in the angular displacement of the moving stimuli resulted in an increase in both the MMN amplitude and the subjective discrimination, although their correlation remained below the significance level. The results obtained are discussed from the point of view of preconscious perception of auditory spatial information.     

Colour Vision Impairment in Young Alcohol Consumers

Alcohol consumption among young adults is widely accepted in modern society and may be the starting point for abusive use of alcohol at later stages of life. Chronic alcohol exposure can lead to visual function impairment. In the present study, we investigated the spatial luminance contrast sensitivity, colour arrangement ability, and colour discrimination thresholds on young adults that weekly consume alcoholic beverages without clinical concerns. Twenty-four young adults were evaluated by an ophthalmologist and performed three psychophysical tests to evaluate their vision functions. We estimated the spatial luminance contrast sensitivity function at 11 spatial frequencies ranging from 0.1 to 30 cycles/degree. No difference in contrast sensitivity was observed comparing alcohol consumers and control subjects. For the evaluation of colour vision, we used the Farnsworth-Munsell 100 hue test (FM 100 test) to test subject’s ability to perform a colour arrangement task and the Mollon-Reffin test (MR test) to measure subject’s colour discrimination thresholds. Alcohol consumers made more mistakes than controls in the FM100 test, and their mistakes were diffusely distributed in the FM colour space without any colour axis preference. Alcohol consumers also performed worse than controls in the MR test and had higher colour discrimination thresholds compared to controls around three different reference points of a perceptually homogeneous colour space, the CIE 1976 chromaticity diagram. There was no colour axis preference in the threshold elevation observed among alcoholic subjects. Young adult weekly alcohol consumers showed subclinical colour vision losses with preservation of spatial luminance contrast sensitivity. Adolescence and young adult age are periods of important neurological development and alcohol exposure during this period of life might be responsible for deficits in visual functions, especially colour vision that is very sensitive to neurotoxicants.     

Comparative psychophysics of bumblebee and honeybee colour discrimination and object detection

Bumblebee (Bombus terrestris) discrimination of targets with broadband reflectance spectra was tested using simultaneous viewing conditions, enabling an accurate determination of the perceptual limit of colour discrimination excluding confounds from memory coding (experiment 1). The level of colour discrimination in bumblebees, and honeybees (Apis mellifera) (based upon previous observations), exceeds predictions of models considering receptor noise in the honeybee. Bumblebee and honeybee photoreceptors are similar in spectral shape and spacing, but bumblebees exhibit significantly poorer colour discrimination in behavioural tests, suggesting possible differences in spatial or temporal signal processing. Detection of stimuli in a Y-maze was evaluated for bumblebees (experiment 2) and honeybees (experiment 3). Honeybees detected stimuli containing both green-receptor-contrast and colour contrast at a visual angle of approximately 5 degrees , whilst stimuli that contained only colour contrast were only detected at a visual angle of 15 degrees . Bumblebees were able to detect these stimuli at a visual angle of 2.3 degrees and 2.7 degrees , respectively. A comparison of the experiments suggests a tradeoff between colour discrimination and colour detection in these two species, limited by the need to pool colour signals to overcome receptor noise. We discuss the colour processing differences and possible adaptations to specific ecological habitats.     

Assessment of novel binocular colour, motion and contrast tests in glaucoma

The effects of glaucoma on binocular visual sensitivity for the detection of various stimulus attributes are investigated at the fovea and in four paracentral retinal regions. The study employed a number of visual stimuli designed to isolate the processing of various stimulus attributes. We measured absolute contrast detection thresholds and functional contrast sensitivity by using Landolt ring stimuli. This psychophysical Landolt C-based contrast test of detection and gap discrimination allowed us to test parafoveally at 6 ° from fixation and foveally by employing interleaved testing locations. First-order motion perception was examined by using moving stimuli embedded in static luminance contrast noise. Red/green (RG) and yellow/blue (YB) colour thresholds were measured with the Colour Assessment and Diagnosis (CAD) test, which utilises random dynamic luminance contrast noise (± 45 %) to ensure that only colour and not luminance signals are available for target detection. Subjects were normal controls (n?=?65) and glaucoma patients with binocular visual field defects (n?=?15) classified based on their Humphrey Field Analyzer mean deviation (MD) scores. The impairment of visual function varied depending on the stimulus attribute and location tested. Progression of loss was noted for all tests as the degree of glaucoma increased. For subjects with mild glaucoma (MD ?0.01 dB to ?6.00 dB) significantly more data points fell outside the normal age-representative range for RG colour thresholds than for any other visual test, followed by motion thresholds. This was particularly the case for the parafoveal data compared with the foveal data. Thus, a multifaceted measure of binocular visual performance, incorporating RG colour and motion test at multiple locations, might provide a better index for comparison with quality of life measures in glaucoma.     

Testing Neuronal Accounts of Anisotropic Motion Perception with Computational Modelling

There is an over-representation of neurons in early visual cortical areas that respond most strongly to cardinal (horizontal and vertical) orientations and directions of visual stimuli, and cardinal- and oblique-preferring neurons are reported to have different tuning curves. Collectively, these neuronal anisotropies can explain two commonly-reported phenomena of motion perception – the oblique effect and reference repulsion – but it remains unclear whether neuronal anisotropies can simultaneously account for both perceptual effects. We show in psychophysical experiments that reference repulsion and the oblique effect do not depend on the duration of a moving stimulus, and that brief adaptation to a single direction simultaneously causes a reference repulsion in the orientation domain, and the inverse of the oblique effect in the direction domain. We attempted to link these results to underlying neuronal anisotropies by implementing a large family of neuronal decoding models with parametrically varied levels of anisotropy in neuronal direction-tuning preferences, tuning bandwidths and spiking rates. Surprisingly, no model instantiation was able to satisfactorily explain our perceptual data. We argue that the oblique effect arises from the anisotropic distribution of preferred directions evident in V1 and MT, but that reference repulsion occurs separately, perhaps reflecting a process of categorisation occurring in higher-order cortical areas.     

A variational approach to behavioral and neuroelectrical laws

Variational methods play a fundamental and unifying role in several fields of physics, chemistry, engineering, economics, and biology, as they allow one to derive the behavior of a system as a consequence of an optimality principle. A possible application of these methods to a model of perception is given by considering a psychophysical law as the solution of an Euler-Lagrange equation. A general class of Lagrangians is identified by requiring the measurability of prothetic continua on interval scales. The associated Hamiltonian (the energy of the process) is tentatively connected with neurophysiological aspects. As an example of the suggested approach a particular choice of the Lagrangian, that is a sufficient condition to obtain classical psychophysical laws, while accounting for psychophysical adaptation and the stationarity of neuronal activity, is used to explore a possible relation between a behavioral law and a neuroelectrical ,response based on the Naka-Rushton model.     

Improved Discrimination of Visual Stimuli Following Repetitive Transcranial Magnetic Stimulation

Background

Repetitive transcranial magnetic stimulation (rTMS) at certain frequencies increases thresholds for motor-evoked potentials and phosphenes following stimulation of cortex. Consequently rTMS is often assumed to introduce a “virtual lesion” in stimulated brain regions, with correspondingly diminished behavioral performance.

Methodology/Principal Findings

Here we investigated the effects of rTMS to visual cortex on subjects'' ability to perform visual psychophysical tasks. Contrary to expectations of a visual deficit, we find that rTMS often improves the discrimination of visual features. For coarse orientation tasks, discrimination of a static stimulus improved consistently following theta-burst stimulation of the occipital lobe. Using a reaction-time task, we found that these improvements occurred throughout the visual field and lasted beyond one hour post-rTMS. Low-frequency (1 Hz) stimulation yielded similar improvements. In contrast, we did not find consistent effects of rTMS on performance in a fine orientation discrimination task.

Conclusions/Significance

Overall our results suggest that rTMS generally improves or has no effect on visual acuity, with the nature of the effect depending on the type of stimulation and the task. We interpret our results in the context of an ideal-observer model of visual perception.