Brightness Discrimination Research Articles

TACS phase-specifically biases brightness perception of flickering light

BackgroundVisual phenomena like brightness illusions impressively demonstrate the highly constructive nature of perception. In addition to physical illumination, the subjective experience of brightness is related to temporal neural dynamics in visual cortex. ObjectiveHere, we asked whether biasing the temporal pattern of neural excitability in visual cortex by transcranial alternating current stimulation (tACS) modulates brightness perception of concurrent rhythmic visual stimuli. MethodsParticipants performed a brightness discrimination task of two flickering lights, one of which was targeted by same-frequency electrical stimulation at varying phase shifts. tACS was applied with an occipital and a periorbital active control montage, based on simulations of electrical currents using finite element head models. ResultsExperimental results reveal that flicker brightness perception is modulated dependent on the phase shift between sensory and electrical stimulation, solely under occipital tACS. Phase-specific modulatory effects by tACS were dependent on flicker-evoked neural phase stability at the tACS-targeted frequency, recorded prior to electrical stimulation. Further, the optimal timing of tACS application leading to enhanced brightness perception was correlated with the neural phase delay of the cortical flicker response. ConclusionsOur results corroborate the role of temporally coordinated neural activity in visual cortex for brightness perception of rhythmic visual input in humans. Phase-specific behavioral modulations by tACS emphasize its efficacy to transfer perceptually relevant temporal information to the cortex. These findings provide an important step towards understanding the basis of visual perception and further confirm electrical stimulation as a tool for advancing controlled modulations of neural activity and related behavior.

Relative contributions of melanopsin to brightness discrimination when hue and luminance also vary.

A large number of studies have shown the effect of melanopsin-dependent retinal ganglion cells on humans performing brightness discrimination tasks. These studies often utilized targets that only differ in their melanopsin activation levels, and not in their luminance or hue, which are both factors that make large contributions to brightness discrimination. The purpose of the present study was to evaluate the relative contribution of melanopsin activation to brightness discrimination when luminance and hue are also varying in addition to melanopsin activation. Using an apparatus consisting of three separate high luminance projectors, we were able to manipulate melanopsin-isolating stimulation, and L-, M-, and S-cone stimulation separately, thus allowing us to vary stimuli in their melanopsin activation, luminance, and hue category independently. We constructed three sets of target stimuli with three different levels of melanopsin activation (100%, 131%, and 167% relative melanopsin excitation) and five levels of luminance. We then had subjects do a two-alternative forced choice task where they compared the previously described target stimuli set to a set of four comparison stimuli that varied in their hue category but had identical luminances. We found that in our stimuli set the overall contribution of melanopsin activity to brightness discrimination was small (an average of 6% increase in likelihood to call a high melanopsin activity stimulus brighter compared to a low melanopsin activity stimulus) when luminance and hue also varied. However, a significant interaction showed that when the comparison was between stimuli differing only in melanopsin stimulation (with luminance and hue unchanged) the contribution of melanopsin to brightness judgments was about 3 times larger (an average of 18% increase in likelihood to call a high melanopsin activity stimulus brighter compared to a low melanopsin activity stimulus). This suggests that although luminance and hue have large effects on brightness discrimination such that the melanopsin contribution can become hard to detect, when there are minimal cone-dependent signals available, melanopsin can make a large contribution to brightness discrimination.

The Eye of the Common Octopus (Octopus vulgaris).

Octopus vulgaris, well-known from temperate waters of the Mediterranean Sea and a well-cited model species among the cephalopods, has large eyes with which it scans its environment actively and which allow the organism to discriminate objects easily. On cursory examination, the single-chambered eyes of octopus with their spherical lenses resemble vertebrate eyes. However there are also apparent differences. For example, the retina of the octopus is everted instead of inverted, and it is equipped with primary rhabdomeric photoreceptors rather than secondary ciliary variety found in the retina of the vertebrate eye. The eyes of octopus are well adapted to the habitat and lifestyle of the species; the pupil closes quickly as a response to sudden light stimuli mimicking a situation in which the octopus leaves its den in shallow water during daytime. Although the many general anatomical and physiological features of octopus vision have been described elsewhere, our review reveals that a lot of information is still missing. Investigations that remain to be undertaken include a detailed examination of the dioptric apparatus or the visual functions such as brightness discrimination as well as a conclusive test for a faculty analogous to, or in lieu of, color vision. For a better understanding of the octopus eye and the functions mediated by it, we suggest that future studies focus on knowledge gaps that we outline in the present review.

The transfer of perceptual learning to a physically and orientation different stimulus requires triple training

Perceptual learning becomes completely transferrable with double training when the observers receive additional exposure of the transfer stimulus via an irrelevant task. However, we noticed that Vernier learning transferred to a new location after double training only when the training and transfer stimuli shared the same orientation. A second exposure to a suprathreshold orthogonal Vernier was necessary to enable cross-orientation learning transfer (Wang_et_al._2012). Here we investigated whether orientation learning transfer between orthogonal gratings and symmetric dot patterns also requires such triple training (training plus double exposures). Observers learned orientation discrimination of a grating or symmetric dot pattern, and the learning transfer to the other, orthogonal, stimulus was tested. We found (1) Baseline: Grating orientation learning (35°/125°) had little effect on orthogonal dot-pattern orientation discrimination; (2) Double training: Additional exposure to the orthogonal dot patterns via a brightness discrimination task produced incomplete learning transfer to orthogonal dot patterns, since further dot-pattern orientation training induced extra performance gain; (3) Triple training: Observers received exposure of both new orientation and new stimulus via two separate tasks: grating brightness discrimination at the orthogonal orientation, and dot pattern brightness discrimination at the same or orthogonal orientation. After triple training, dot-pattern orthogonal orientation discrimination was improved, and continued training produced no further gains, indicating complete transfer of grating orientation learning to the orthogonal symmetric dot patterns. (4) Control: The above double exposures by themselves had little impact on dot-pattern orientation discrimination. (5) The same patterns of learning transfer results were replicated from dot-patterns to gratings. These results suggest that the brain performs two remapping to transfer perceptual learning. One remaps learning to an orthogonal orientation of the same stimulus, which is enabled by exposure of the same but orthogonal-oriented stimulus. The other remaps learning to a physically distinct stimulus by exposure of the new stimulus.

Clarifying frequency-dependent brightness enhancement: delta- and theta-band flicker, not alpha-band flicker, consistently seen as brightest.

Frequency-dependent brightness enhancement, a perceptual illusion in which a flickering light can appear twice as bright as a constant light, has historically been reported to produce maximum effects at a flicker rate within the alpha (8-12Hz) band (Bartley in J Exp Psychol 23(3):313-319, 1938). Our recent examinations of this phenomenon using brightness discrimination between two flickering stimuli, however, have instead revealed the brightest percepts from theta-band (4-7Hz) flicker (Bertrand et al. in Sci Rep8(1):6152, 2018). Two primary questions arise from these seemingly contradictory findings: first, could task differences between these studies have caused recruitment of discrete oscillatory processes? Second, could the reported theta-band flicker enhancement be the result of an aliased alpha rhythm, sequentially sampling two stimulus locations, resulting in an ~ 5Hz half-alpha rhythm? Here, we investigated these questions with two experiments: one replicating Bartley's (1938) adjustment paradigm, and one containing both Bartley's adjustment task and Bertrand's (2018) discrimination task, but presenting stimuli only sequentially (rather than concurrently). Examination of a range of frequencies (2-12Hz) revealed the greatest brightness enhancement arising from flicker in the delta- and theta-band across all conditions, regardless of the spatial or temporal configuration of the stimuli. We speculate that these slower rhythms play an integral role in complex visual operations (e.g., a discrimination decision) where the entrainment of the endogenous neural rhythm to matched exogenous rhythmic stimulation promotes more efficient processing of visual information and thus produces perceptual biases as seen in frequency-dependent brightness enhancement.

Neurovisualization of multidimensional psychophysiological structure of consciousness during the elementary process of brightness discrimination

This chapter will introduce a series of methodologies which can be applied to determine the most influential factors in the study of quality in public transport. These methodologies provide important information highlighting the points that need addressing to make improvements in quality. The methods used to determine the importance of the main factors affecting quality in public transport are described and supported by the development of two methods: factorial analysis and importance-performance analysis. This is followed by the presentation of a series of indexes and procedures which have been used to study public transport service quality: SERVQUAL, SERVPERF, and Quattro. These indexes also reveal the attributes considered to be most representative of service quality, forming the starting point for the studies that are developed later.

Entrainment of theta, not alpha, oscillations is predictive of the brightness enhancement of a flickering stimulus

Frequency-dependent brightness enhancement, where a flickering light can appear twice as bright as an equiluminant constant light, has been reported to exist within the alpha (8–12 Hz) band. Could oscillatory neural activity be driving this perceptual effect? Here, in two experiments, human subjects reported which of two flickering stimuli were brighter. Strikingly, 4 Hz stimuli were reported as brighter more than 80% of the time when compared to all other tested frequencies, even though all stimuli were equiluminant and of equal temporal length. Electroencephalography recordings showed that inter-trial phase coherence (ITC) of theta (4 Hz) was: (1) Significantly greater than alpha, contralateral to the flickering stimulus; (2) Enhanced by the presence of a second ipsilateral 4 Hz flickering stimulus; and (3) Uniquely lateralized, unlike the alpha band. Importantly, on trials with two identical stimuli (i.e. 4 Hz vs 4 Hz), the brightness discrimination judgment could be predicted by the hemispheric balance in the amount of 4 Hz ITC. We speculate that the theta rhythm plays a distinct information transfer role, where its ability to share information between hemispheres via entrainment promotes a better processing of visual information to inform a discrimination decision.

Melanopsin- and L-cone–induced pupil constriction is inhibited by S- and M-cones in humans

The human retina contains five photoreceptor types: rods; short (S)-, mid (M)-, and long (L)-wavelength-sensitive cones; and melanopsin-expressing ganglion cells. Recently, it has been shown that selective increments in M-cone activation are paradoxically perceived as brightness decrements, as opposed to L-cone increments. Here we show that similar effects are also observed in the pupillary light response, whereby M-cone or S-cone increments lead to pupil dilation whereas L-cone or melanopic illuminance increments resulted in pupil constriction. Additionally, intermittent photoreceptor activation increased pupil constriction over a 30-min interval. Modulation of L-cone or melanopic illuminance within the 0.25-4-Hz frequency range resulted in more sustained pupillary constriction than light of constant intensity. Opposite results were found for S-cone and M-cone modulations (2 Hz), mirroring the dichotomy observed in the transient responses. The transient and sustained pupillary light responses therefore suggest that S- and M-cones provide inhibitory input to the pupillary control system when selectively activated, whereas L-cones and melanopsin response fulfill an excitatory role. These findings provide insight into functional networks in the human retina and the effect of color-coding in nonvisual responses to light, and imply that nonvisual and visual brightness discrimination may share a common pathway that starts in the retina.

Do you see what I see? The difference between dog and human visual perception may affect the outcome of experiments

The visual sense of dogs is in many aspects different than that of humans. Unfortunately, authors do not explicitly take into consideration dog-human differences in visual perception when designing their experiments. With an image manipulation program we altered stationary images, according to the present knowledge about dog-vision. Besides the effect of dogs’ dichromatic vision, the software shows the effect of the lower visual acuity and brightness discrimination, too. Fifty adult humans were tested with pictures showing a female experimenter pointing, gazing or glancing to the left or right side. Half of the pictures were shown after they were altered to a setting that approximated dog vision. Participants had difficulty to find out the direction of glancing when the pictures were in dog-vision mode. Glances in dog-vision setting were followed less correctly and with a slower response time than other cues. Our results are the first that show the visual performance of humans under circumstances that model how dogs’ weaker vision would affect their responses in an ethological experiment. We urge researchers to take into consideration the differences between perceptual abilities of dogs and humans, by developing visual stimuli that fit more appropriately to dogs’ visual capabilities.

Cholinergic Potentiation of Restoration of Visual Function after Optic Nerve Damage in Rats.

Enhancing cortical plasticity and brain connectivity may improve residual vision following a visual impairment. Since acetylcholine plays an important role in attention and neuronal plasticity, we explored whether potentiation of the cholinergic transmission has an effect on the visual function restoration. To this end, we evaluated for 4 weeks the effect of the acetylcholinesterase inhibitor donepezil on brightness discrimination, visually evoked potentials, and visual cortex reactivity after a bilateral and partial optic nerve crush in adult rats. Donepezil administration enhanced brightness discrimination capacity after optic nerve crush compared to nontreated animals. The visually evoked activation of the primary visual cortex was not restored, as measured by evoked potentials, but the cortical neuronal activity measured by thallium autometallography was not significantly affected four weeks after the optic nerve crush. Altogether, the results suggest a role of the cholinergic system in postlesion cortical plasticity. This finding agrees with the view that restoration of visual function may involve mechanisms beyond the area of primary damage and opens a new perspective for improving visual rehabilitation in humans.

2D histogram equalisation based on the human visual system

Histogram equalisation (HE) methods using the 2D histogram (2DH) have achieved a great success in contrast enhancement. The 2DH is constructed by using the occurrence of a local pixel pair (LPP) consisting of each pixel and its surrounding pixels. However, the 2DH-based methods often produce over-stretching artefacts because the low-textured regions primarily existing in the image induce a spike at some LPPs in the 2DH. To solve this problem, the 2DH is constructed by employing two properties of the human visual system (HVS) as follows: the HVS has the better brightness discrimination in the dark region according to Weber's law, and the HVS is less sensitive to visual artefacts in the higher-textured regions. To create a spike-free 2DH, a weighting function reflecting these two properties of the HVS is designed for the LPP. As compared with the popular 2DH-based methods, the HE with the proposed 2DH can effectively enhance the image contrast while achieving the best perceptual similarity score between the input and output images.

Rendering of HDR images to improve brightness discrimination

The impression of quality of images can be enhanced on a high dynamic range (HDR) displays. Generally, a conventional 8-bit image can be processed to an HDR image by inverse tone mapping operators. Among the operators, brightness discrimination mapping by applying brightness adaptation model attempted to mimic the human visual system. In this paper, we use a brightness adaptation model to derive a brightness discrimination mapping algorithm for HDR displays. The proposed algorithm maximizes a function, which represents the local and global brightness discrimination range by exploiting characteristics of the human visual system. Enhancement of details is verified by visualizing HDR images from dark to bright regions. Improvement of dynamic range is quantified by measuring increased discrimination ratio.

Why do seals have cones? Behavioural evidence for colour-blindness in harbour seals.

All seals and cetaceans have lost at least one of two ancestral cone classes and should therefore be colour-blind. Nevertheless, earlier studies showed that these marine mammals can discriminate colours and a colour vision mechanism has been proposed which contrasts signals from cones and rods. However, these earlier studies underestimated the brightness discrimination abilities of these animals, so that they could have discriminated colours using brightness only. Using a psychophysical discrimination experiment, we showed that a harbour seal can solve a colour discrimination task by means of brightness discrimination alone. Performing a series of experiments in which two harbour seals had to discriminate the brightness of colours, we also found strong evidence for purely scotopic (rod-based) vision at light levels that lead to mesopic (rod–cone-based) vision in other mammals. This finding speaks against rod–cone-based colour vision in harbour seals. To test for colour-blindness, we used a cognitive approach involving a harbour seal trained to use a concept of same and different. We tested this seal with pairs of isoluminant stimuli that were either same or different in colour. If the seal had perceived colour, it would have responded to colour differences between stimuli. However, the seal responded with “same”, providing strong evidence for colour-blindness.Electronic supplementary materialThe online version of this article (doi:10.1007/s10071-014-0823-3) contains supplementary material, which is available to authorized users.

Characterization of the role of adult neurogenesis in touch-screen discrimination learning.

Recent theories posit that adult neurogenesis supports dentate gyrus pattern separation and hence is necessary for some types of discrimination learning. Using an inducible transgenic mouse model, we investigated the contribution of adult-born neurons to spatial and nonspatial touch-screen discriminations of varying levels of difficulty. Arresting neurogenesis caused a modest but statistically significant impairment in a position discrimination task. However, the effect was present only on trials after a learned discrimination was reversed, suggesting that neurogenesis supports cognitive flexibility rather than spatial discrimination per se. The deficit was present 4-10 weeks after the arrest of neurogenesis but not immediately after, consistent with previous evidence that the behavioral effects of arresting neurogenesis arise because of the depletion of adult-born neurons at least 1 month old. The arrest of neurogenesis failed to affect a nonspatial brightness discrimination task that was equal in difficulty to the spatial task. The data suggest that adult neurogenesis is not strictly necessary for spatial or perceptual discrimination learning and instead implicate adult neurogenesis in factors related to reversal learning, such as cognitive flexibility or proactive interference.

Automatic spatial association for luminance

In the present study, we investigated whether luminance and the side of response execution are associated, showing a SNARC-like effect (faster responses with the left hand for dark stimuli, and vice versa for light stimuli). A total of 30 participants were tested in two experiments. In Experiment1, the association between space and the luminance of chromatic stimuli was directly tested (brightness discrimination). In Experiment2, the same spatial association was tested indirectly (hue discrimination). The results showed that participants responded faster with their left hand to hues with lower luminance, and with their right hand to hues with higher luminance, in either the direct or the indirect task. The consistency of this association in both tasks demonstrates the automaticity of the SNARC-like effect for luminance.

Measuring psychometric functions with the diffusion model.

The diffusion decision model (Ratcliff, 1978) was used to examine discrimination for a range of perceptual tasks: numerosity discrimination, number discrimination, brightness discrimination, motion discrimination, speed discrimination, and length discrimination. The model produces a measure of the quality of the information that drives decision processes, a measure termed drift rate in the model. As drift rate varies across experimental conditions that differ in difficulty, a psychometric function that plots drift rate against difficulty can be constructed. Psychometric functions for the tasks in this article usually plot accuracy against difficulty, but for some levels of difficulty, accuracy can be at ceiling. The diffusion model extends the range of difficulty that can be evaluated because drift rates depend on response times (RTs) as well as accuracy, and when RTs decrease across conditions that are all at ceiling in accuracy, then drift rates will distinguish among the conditions. Signal detection theory assumes that the variable driving performance is the z-transform of the accuracy value, and, somewhat surprisingly, this closely matches drift rate extracted from the diffusion model when accuracy is not at ceiling, but not sometimes when accuracy is high. Even though the functions are similar in the middle of the range, the interpretations of the variability in the models (e.g., perceptual variability, decision process variability) are incompatible.

Brightness discrimination in the day- and night-adapted wandering spider Cupiennius salei.

Cupiennius salei is a nocturnal spider with eight eyes, which undergo a remarkable circadian cycle: the rhabdomeric membrane of the photoreceptor cells is dismantled during the day and rebuilt at the beginning of the night. Such drastic changes might influence the brightness discrimination ability. We tested this hypothesis by presenting square-shaped flickering stimuli with certain luminances on stationary backgrounds with other luminances to spiders with day- or night-adapted eyes. When the spider, through its three pairs of so-called secondary eyes, perceives a visible contrast between the stimulus and the background, its principal eye muscle activity should increase. We therefore recorded this activity in vivo to assess the brightness discrimination ability of Cupiennius salei. Our results show that this spider has good brightness discrimination ability, which is significantly better with dark-adapted eyes. A Michelson contrast of 0.1 to 0.2 at night, and of 0.2 to 0.3 for day-adapted eyes, is sufficient to elicit a significant response, except below a critical value of luminance (~16 cd m(-2)), where the minimal perceivable contrast needs to be higher. In the Discussion we compare these performances with those of other animals, in particular with jumping spiders.

The hare and the tortoise: Emphasizing speed can change the evidence used to make decisions.

Decision-makers effortlessly balance the need for urgency against the need for caution. Theoretical and neurophysiological accounts have explained this tradeoff solely in terms of the quantity of evidence required to trigger a decision (the "threshold"). This explanation has also been used as a benchmark test for evaluating new models of decision making, but the explanation itself has not been carefully tested against data. We rigorously test the assumption that emphasizing decision speed versus decision accuracy selectively influences only decision thresholds. In data from a new brightness discrimination experiment we found that emphasizing decision speed over decision accuracy not only decreases the amount of evidence required for a decision but also decreases the quality of information being accumulated during the decision process. This result was consistent for 2 leading decision-making models and in a model-free test. We also found the same model-based results in archival data from a lexical decision task (reported by Wagenmakers, Ratcliff, Gomez, & McKoon, 2008) and new data from a recognition memory task. We discuss discuss implications for theoretical development and applications.

Saccadic brightness decisions do not use a difference model

Eye movements are the most frequent (∼3 per second), shortest-latency (∼150-250 ms), and biomechanically simplest (1 joint, no inertial complexities) voluntary motor behavior in primates, providing a model sensorimotor decision-making system. Current computational "difference" models of choice behavior utilize a single decision variable encoding the difference between two alternate signals, often implemented as a log-likelihood ratio. Alternatively, the oculomotor literature describes a "race" mechanism, in which two separate decision variables encoding the two alternate signals race against one another independently. These two models make two qualitatively distinct predictions, which can be tested empirically with a two-alternative forced-choice task. Unlike the race model, a decision variable based upon a differencing operation predicts strong mirror image correlations between response time (RT) and the signal strengths of the selected and unselected stimuli (because differencing creates equal and opposite correlations). In a saccadic brightness discrimination task, we observed positive correlations between response rate (1/RT) and the strength of both the selected and unselected stimulus, a simple qualitative prediction of race models that applies to any 2AFC task but which is fundamentally at odds with the most basic prediction of any difference model. Our data are, however, qualitatively consistent with a mechanism in which two competing motor plans co-exist and their two corresponding neural decision variables race to a threshold to drive the saccadic decision.

Ultraviolet reflection enhances the risk of predation in a vertebrate

Abstract Many animals are sensitive to ultraviolet light and also possess UV-reflective regions on their body surface. Individuals reflecting UV have been shown to be preferred during social interactions such as mate choice or shoaling decisions. However, whether those body UV-reflections enhance also the conspicuousness to UV-sensitive predators and therefore entail costs for its bearer is less well documented. Two size-matched three-spined sticklebacks Gasterosteus aculeatus, one enclosed in a UV-transmitting (UV+) and another in a UV-blocking (UV-) chamber, were simultaneously presented to individual brown trout Salmo trutta. “yearlings”. Brown trout of this age are sensitive to the UV part of the electromagnetic spectrum and are natural predators of three-spined sticklebacks. The stickleback that was attacked first as well as the subsequent number of attacks was recorded. Sticklebacks enclosed in the UV-transmitting chamber were attacked first significantly more often compared to sticklebacks enclosed in the UV-blocking chamber. Control experiments using neutral density filters revealed that this was more likely due to UV having an influence on hue perception rather than brightness discrimination. The difference in attack probability corresponded to the difference in chromatic contrasts between sticklebacks and the experimental background calculated for both the UV+ and UVconditions in a physiological model of trout colour vision. UV reflections seem to be costly by enhancing the risk of predation due to an increased conspicuousness of prey. This is the first study in a vertebrate, to our knowledge, demonstrating direct predation risk due to UV wavelengths.