Ultraviolet vision in anemonefish improves colour discrimination.

In many animals, ultraviolet (UV) vision guides navigation, foraging, and communication, but few studies have addressed the contribution of UV signals to colour vision, or measured UV discrimination thresholds using behavioural experiments. Here, we tested UV colour vision in an anemonefish (Amphiprion ocellaris) using a five-channel (RGB-V-UV) LED display. We first determined that the maximal sensitivity of the A. ocellaris UV cone was ∼386 nm using microspectrophotometry. Three additional cone spectral sensitivities had maxima at ∼497, 515 and ∼535 nm. We then behaviourally measured colour discrimination thresholds by training anemonefish to distinguish a coloured target pixel from grey distractor pixels of varying intensity. Thresholds were calculated for nine sets of colours with and without UV signals. Using a tetrachromatic vision model, we found that anemonefish were better (i.e. discrimination thresholds were lower) at discriminating colours when target pixels had higher UV chromatic contrast. These colours caused a greater stimulation of the UV cone relative to other cone types. These findings imply that a UV component of colour signals and cues improves their detectability, which likely increases the prominence of anemonefish body patterns for communication and the silhouette of zooplankton prey.

In primary visual cortex fMRI responses to chromatic and achromatic stimuli are interdependent and predict contrast detection thresholds.

Chromatic and achromatic signals in primary visual cortex have historically been considered independent of each other but have since shown evidence of interdependence. Here, we investigated the combination of two components of a stimulus; an achromatic dynamically changing check background and a chromatic (L-M or S cone) target grating. We found that combinations of chromatic and achromatic signals in primary visual cortex were interdependent, with the dynamic range of responses to chromatic contrast decreasing as achromatic contrast increased. A contrast detection threshold study also revealed interdependence of background and target, with increasing chromatic contrast detection thresholds as achromatic background contrast increased. A model that incorporated a normalising effect of achromatic contrast on chromatic responses, but not vice versa, best predicted our V1 data as well as behavioural thresholds. Further along the visual hierarchy, the dynamic range of chromatic responses was maintained when compared to achromatic responses, which became increasingly compressive.

Release from response interference in color-word contingency learning.

In identifying the print colors of words when some combinations of color and word occur more frequently than others, people quickly show evidence of learning these associations. This contingency learning effect is evident in faster and more accurate responses to high-contingency combinations than to low-contingency combinations. Across four experiments, we systematically varied the number of response-irrelevant word stimuli connected to response-relevant colors. In each experiment, one group experienced the typical contingency learning paradigm with three colors linked to three words; other groups saw more words (six or twelve) linked to the same three colors. All four experiments disconfirmed a central prediction derived from the Parallel Episodic Processing (PEP 2.0) model (Schmidt et al., 2016)-that the magnitude of the contingency learning effect should remain stable as more words are added to the response-irrelevant dimension, as long as the color-word contingency ratios are maintained. Responses to high-contingency items did slow down numerically as the number of words increased between groups, consistent with the prediction from PEP 2.0, but these changes were unreliable. Inconsistent with PEP 2.0, however, overall response time did not slow down and responses to low-contingency items actually sped up as the number of words increased across groups. These findings suggest that the PEP 2.0 model should be modified to incorporate response interference caused by high-probability associations when responding to low-probability combinations.

Rapid color categorization revealed by frequency-tagging-based EEG.

There has been much debate on whether color categories affect how we perceive color. Recent theories have put emphasis on the role of top-down influence on color perception that the original continuous color space in the visual cortex may be transformed into categorical encoding due to top-down modulation. To test the influence of color categories on color perception, we adopted an RSVP paradigm, where color stimuli were presented at a fast speed of 100 ms per stimulus and were forward and backward masked by the preceding and following stimuli. Moreover, no explicit color naming or categorization was required. In theory, backward masking with such a short interval in a passive viewing task should constrain top-down influence from higher-level brain areas. To measure any potentially subtle differences in brain response elicited by different color categories, we embedded a sensitive frequency-tagging-based EEG paradigm within the RSVP stimuli stream where the oddball color stimuli were encoded with a different frequency from the base color stimuli. We showed that EEG responses to cross-category oddball colors at the frequency where the oddball stimuli were presented was significantly larger than the responses to within-category oddball colors. Our study suggested that the visual cortex can automatically and implicitly encode color categories when color stimuli are presented rapidly.

Assessing perceptual chromatic equiluminance using a reflexive pupillary response.

Equiluminant stimuli help assess the integrity of colour perception and the relationship of colour to other visual features. As a result of individual variation, it is necessary to calibrate experimental visual stimuli to suit each individual's unique equiluminant ratio. Most traditional methods rely on training observers to report their subjective equiluminance point. Such paradigms cannot easily be implemented on pre-verbal or non-verbal observers. Here, we present a novel Pupil Frequency-Tagging Method (PFTM) for detecting a participant's unique equiluminance point without verbal instruction and with minimal training. PFTM analyses reflexive pupil oscillations induced by slow (< 2 Hz) temporal alternations between coloured stimuli. Two equiluminant stimuli will induce a similar pupil dilation response regardless of colour; therefore, an observer's equiluminant point can be identified as the luminance ratio between two colours for which the oscillatory amplitude of the pupil at the tagged frequency is minimal. We compared pupillometry-based equiluminance ratios to those obtained with two established techniques in humans: minimum flicker and minimum motion. In addition, we estimated the equiluminance point in non-human primates, demonstrating that this new technique can be successfully employed in non-verbal subjects.

Effector-independent Representations Guide Sequential Target Selection Biases in Action.

Previous work shows that automatic attention biases toward recently selected target features transfer across action and perception and even across different effectors such as the eyes and hands on a trial-by-trial basis. Although these findings suggest a common neural representation of selection history across effectors, the extent to which information about recently selected target features is encoded in overlapping versus distinct brain regions is unknown. Using fMRI and a priming of pop-out task where participants selected unpredictable, uniquely colored targets among homogeneous distractors via reach or saccade, we show that color priming is driven by shared, effector-independent underlying representations of recent selection history. Consistent with previous work, we found that the intraparietal sulcus (IPS) was commonly activated on trials where target colors were switched relative to those where the colors were repeated; however, the dorsal anterior insula exhibited effector-specific activation related to color priming. Via multivoxel cross-classification analyses, we further demonstrate that fine-grained patterns of activity in both IPS and the medial temporal lobe encode information about selection history in an effector-independent manner, such that ROI-specific models trained on activity patterns during reach selection could predict whether a color was repeated or switched on the current trial during saccade selection and vice versa. Remarkably, model generalization performance in IPS and medial temporal lobe also tracked individual differences in behavioral priming sensitivity across both types of action. These results represent a first step to clarify the neural substrates of experience-driven selection biases in contexts that require the coordination of multiple actions.

Induced awareness of synesthetic sensations in synesthetically predisposed "Borderline Non-synesthetes".

A long-standing issue concerning synesthesia is whether the trait is continuous or discontinuous with ordinary perception. Here, we found that a substantial proportion of non-synesthetes (>10 % out of >200 unselected participants) spontaneously became aware of their synesthesia by participating in an online survey that forced them to select colors for stimuli that evoke color sensations in synesthetes. Notably, the test-retest consistencies of color sensation in these non-synesthetes were comparable to those in self-claimed synesthetes, revealing their strong though latent synesthetic dispositions. The effect was absent or weak in a matched control survey that did not include the color-picking test. Therefore, the color-picking task likely provided the predisposed "borderline non-synesthetes" with an opportunity to dwell on their tendency toward synesthesia and allowed their subconscious sensations to become conscious ones. The finding suggests that the general population has a continuum of synesthetic disposition that encompasses both synesthetes and non-synesthetes.

Chromatic discrimination in fixed saturation levels from tufted capuchin monkeys with different color vision genotypes.

Recent research has proposed new approaches to investigate color vision in Old World Monkeys by measuring suprathreshold chromatic discrimination. In this study, we aimed to extend this approach to New World Monkeys with different color vision genotypes by examining their performance in chromatic discrimination tasks along different fixed chromatic saturation axes. Four tufted capuchin monkeys were included in the study, and their color vision genotypes were one classical protanope, one classical deuteranope, one non-classical protanope, and a normal trichromat. During the experiments, the monkeys were required to perform a chromatic discrimination task using pseudoisochromatic stimuli with varying target saturations of 0.06, 0.04, 0.03, and 0.02 u'v' units. The number of errors made by the monkeys along different chromatic axes was recorded, and their performance was quantified using the binomial probability of their hits during the tests. Our results showed that dichromatic monkeys made more errors near the color confusion lines associated with their specific color vision genotypes, while the trichromatic monkey did not demonstrate any systematic errors. At high chromatic saturation, the trichromatic monkey had significant hits in the chromatic axes around the 180° chromatic axis, whereas the dichromatic monkeys had errors in colors around the color confusion lines. At lower saturation, the performance of the dichromatic monkeys became more challenging to differentiate among the three types, but it was still distinct from that of the trichromatic monkey. In conclusion, our findings suggest that high saturation conditions can be used to identify the color vision dichromatic phenotype of capuchin monkeys, while low chromatic saturation conditions enable the distinction between trichromats and dichromats. These results extend the understanding of color vision in New World Monkeys and highlight the usefulness of suprathreshold chromatic discrimination measures in exploring color vision in non-human primates.

Action, emotion, and music-colour synaesthesia: an examination of sensorimotor and emotional responses in synaesthetes and non-synaesthetes.

Synaesthesia has been conceptualised as a joining of sensory experiences. Taking a holistic, embodied perspective, we investigate in this paper the role of action and emotion, testing hypotheses related to (1) changes to action-related qualities of a musical stimulus affect the resulting synaesthetic experience; (2) a comparable relationship exists between music, sensorimotor and emotional responses in synaesthetes and the general population; and (3) sensorimotor responses are more strongly associated with synaesthesia than emotion. 29 synaesthetes and 33 non-synaesthetes listened to 12 musical excerpts performed on a musical instrument they had first-hand experience playing, an instrument never played before, and a deadpan performance generated by notation software, i.e., a performance without expression. They evaluated the intensity of their experience of the music using a list of dimensions that relate to sensorimotor, emotional or synaesthetic sensations. Results demonstrated that the intensity of listeners' responses was most strongly influenced by whether or not music is performed by a human, more so than familiarity with a particular instrument. Furthermore, our findings reveal a shared relationship between emotional and sensorimotor responses among both synaesthetes and non-synaesthetes. Yet it was sensorimotor intensity that was shown to be fundamentally associated with the intensity of the synaesthetic response. Overall, the research argues for, and gives first evidence of a key role of action in shaping the experiences of music-colour synaesthesia.

Resting Stroop task: Evidence of task conflict in trials with no required response.

In the typical Stroop task, participants are presented with color words written in different ink colors and are asked to respond to their color. It has been suggested that the Stroop task consists of two main conflicts: information conflict (color vs. word naming) and task conflict (respond to color vs. read the word). In the current study, we developed a novel task that includes both Response trials (i.e., trials in which a response is required) and Rest trials (i.e., trials in which no response is required or available) and investigated the existence of both information and task conflicts in Rest trials. We found evidence for task conflict in Response and also in Rest trials, while evidence for information conflict was only observed in Response trials. These results are in line with a model of task conflict that occurs independently of and prior to information conflict in the Stroop task.

The basis of S-R learning: associations between individual stimulus features and responses.

Three experiments tested the hypothesis that response selection skill involves associations between individual stimulus features and responses. The Orientation group in Experiments 1 and 2 first practiced responding to the orientation of a line stimulus while ignoring its color, and the Color group practiced responding to the color of the line while disregarding its orientation. When in the ensuing test conditions the Orientation group responded to the color of the line, RTs and errors increased when the then irrelevant line orientation was inconsistent with practice. This confirmed that during practice, Orientation participants had developed orientation feature-response associations they could not fully inhibit. Yet, evidence for color feature-response associations was not observed in the Color group. This was attributed to orientation identification being faster than color identification, even after having practiced responding to colors. Experiment 3 involved practicing to three line stimuli with unique orientation and color combinations. It showed evidence for the independent development of orientation feature-response associations and color feature-response associations. Together, these results indicate that the typical RT reduction with practice in response selection tasks is caused in part by the capacity of participants to learn selecting responses on the basis of the stimulus feature that becomes available first.

Tomatoes Are Red: The Perception of Achromatic Objects Elicits Retrieval of Associated Color Knowledge.

When preparing to name an object, semantic knowledge about the object and its attributes is activated, including perceptual properties. It is unclear, however, whether semantic attribute activation contributes to lexical access or is a consequence of activating a concept irrespective of whether that concept is to be named or not. In this study, we measured neural responses using fMRI while participants named objects that are typically green or red, presented in black line drawings. Furthermore, participants underwent two other tasks with the same objects, color naming and semantic judgment, to see if the activation pattern we observe during picture naming is (a) similar to that of a task that requires accessing the color attribute and (b) distinct from that of a task that requires accessing the concept but not its name or color. We used representational similarity analysis to detect brain areas that show similar patterns within the same color category, but show different patterns across the two color categories. In all three tasks, activation in the bilateral fusiform gyri ("Human V4") correlated with a representational model encoding the red-green distinction weighted by the importance of color feature for the different objects. This result suggests that when seeing objects whose color attribute is highly diagnostic, color knowledge about the objects is retrieved irrespective of whether the color or the object itself have to be named.

Pupil responses to colorfulness are selectively reduced in healthy older adults.

The alignment between visual pathway signaling and pupil dynamics offers a promising non-invasive method to further illuminate the mechanisms of human color perception. However, only limited research has been done in this area and the effects of healthy aging on pupil responses to the different color components have not been studied yet. Here we aim to address this by modelling the effects of color lightness and chroma (colorfulness) on pupil responses in young and older adults, in a closely controlled passive viewing experiment with 26 broad-spectrum digital color fields. We show that pupil responses to color lightness and chroma are independent from each other in both young and older adults. Pupil responses to color lightness levels are unaffected by healthy aging, when correcting for smaller baseline pupil sizes in older adults. Older adults exhibit weaker pupil responses to chroma increases, predominantly along the Green-Magenta axis, while relatively sparing the Blue-Yellow axis. Our findings complement behavioral studies in providing physiological evidence that colors fade with age, with implications for color-based applications and interventions both in healthy aging and later-life neurodegenerative disorders.

Brain activity characteristics of RGB stimulus: an EEG study.

The perception of color is a fundamental cognitive feature of our psychological experience, with an essential role in many aspects of human behavior. Several studies used magnetoencephalography, functional magnetic resonance imaging, and electroencephalography (EEG) approaches to investigate color perception. Their methods includes the event-related potential and spectral power activity of different color spaces, such as Derrington-Krauskopf-Lennie and red-green-blue (RGB), in addition to exploring the psychological and emotional effects of colors. However, we found insufficient studies in RGB space that considered combining all aspects of EEG signals. Thus, in the present study, focusing on RGB stimuli and using a data-driven approach, we investigated significant differences in the perception of colors. Our findings show that beta oscillation of green compared to red and blue colors occurs in early sensory periods with a latency shifting in the occipital region. Furthermore, in the occipital region, the theta power of the blue color decreases noticeably compared to the other colors. Concurrently, in the prefrontal area, we observed an increase in phase consistency in response to the green color, while the blue color showed a decrease. Therefore, our results can be used to interpret the brain activity mechanism of color perception in RGB color space and to choose suitable colors for more efficient performance in cognitive activities.

Mental imagery and visual attentional templates: A dissociation.

There is a growing interest in the relationship between mental images and attentional templates as both are considered pictorial representations that involve similar neural mechanisms. Here, we investigated the role of mental imagery in the automatic implementation of attentional templates and their effect on involuntary attention. We developed a novel version of the contingent capture paradigm designed to encourage the generation of a new template on each trial and measure contingent spatial capture by a template-matching visual feature (color). Participants were required to search at four different locations for a specific object indicated at the start of each trial. Immediately prior to the search display, color cues were presented surrounding the potential target locations, one of which matched the target color (e.g., red for strawberry). Across three experiments, our task induced a robust contingent capture effect, reflected by faster responses when the target appeared in the location previously occupied by the target-matching cue. Contrary to our predictions, this effect remained consistent regardless of self-reported individual differences in visual mental imagery (Experiment 1, N = 216) or trial-by-trial variation of voluntary imagery vividness (Experiment 2, N = 121). Moreover, contingent capture was observed even among aphantasic participants, who report no imagery (Experiment 3, N = 91). The magnitude of the effect was not reduced in aphantasics compared to a control sample of non-aphantasics, although the two groups reported substantial differences in their search strategy and exhibited differences in overall speed and accuracy. Our results hence establish a dissociation between the generation and implementation of attentional templates for a visual feature (color) and subjectively experienced imagery.

The advantage of being a synesthete: The behavioral benefits of ticker-tape synesthesia.

As first described by Francis Galton, some persons perceive vividly and automatically in their mind's eye the written form of words that they are hearing. This phenomenon, labeled ticker-tape synesthesia (TTS), is thought to reflect an abnormally strong influence of speech processing in language areas on to orthographic representations in the visual cortex. Considering the relevance of TTS for the study of reading acquisition, we looked for objective behavioral advantages or impairments in 22 synesthetes, as compared to 22 matched control participants. In three auditory tasks relying on orthographic working memory (letters counting, backward spelling, and letter shape decision), we predicted and observed better performance in synesthetes than in controls. In two visual tasks (lexical decision and letter decision) with a concurrent auditory stimulation, we predicted that synesthetes should suffer from a larger interference by irrelevant speech than controls, but eventually found no difference between the groups. Those results, which we discuss in relation to orthographic processing, mental imagery, and working memory, promote TTS from pure subjectivity to an experimentally measurable phenomenon.

Do We Prepare for What We Predict? How Target Expectations Affect Preparatory Attentional Templates and Target Selection in Visual Search.

Visual search is guided by representations of target-defining features (attentional templates) that are activated in a preparatory fashion. Here, we investigated whether these template activation processes are modulated by probabilistic expectations about upcoming search targets. We tracked template activation while observers prepared to search for one or two possible color-defined targets by measuring N2pc components (markers of attentional capture) to task-irrelevant color probes flashed every 200 msec during the interval between search displays. These probes elicit N2pcs only if the corresponding color template is active at the time when the probe appears. Probe N2pcs emerged from about 600 msec before search display onset. They did not differ between one-color and two-color search, indicating that two color templates can be activated concurrently. Critically, probe N2pcs measured during two-color search were identical for probes matching an expected or unexpected color (target color probability: 80% vs. 20%), or one of two equally likely colors. This strongly suggests that probabilistic target color expectations had no impact on search preparation. In marked contrast, subsequent target selection processes were strongly affected by these expectations. We discuss possible explanations for this clear dissociation in the effects of expectations on preparatory search template activation and search target selection, respectively.

Color Vision in the Mountains.

This Lessons from History article uses science, aviation, medicine, and mountaineering sources to describe some of the effects of hypoxia, illumination, and other environmental conditions on the eye, the central nervous system, and light and color perception. The historical perspective is augmented by an analysis of an informal observation of the altered perception of red color on a deck of playing cards while climbing Mera Peak in the Himalaya. The appearance of a grayer red color on the cards was initially attributed to the effects of hypoxia alone. Instead, analysis of cards in combination with the low incidence of protan color vision defects at altitude indicated that glare and contrast effects in the extremely bright lighting environment combined with hypoxia likely caused the perception of a grayer red. The incident provides an educational opportunity for review, analysis, and commentary about some of the complex elements that impact color vision.

In paired preference tests, domestic chicks innately choose the colour green over red, and the shape of a frog over a sphere when both stimuli are green.

Many animals express unlearned colour preferences that depend on the context in which signals are encountered. These colour biases may have evolved in response to the signalling system to which they relate. For example, many aposematic animals advertise their unprofitability with red warning signals. Predators' innate biases against these warning colours have been suggested as one of the potential explanations for the initial evolution of aposematism. It is unclear, however, whether unlearned colour preferences reported in a number of species is truly an innate behaviour or whether it is based on prior experience. We tested the spontaneous colour and shape preferences of dark-hatched, unfed, and visually naive domestic chicks (Gallus gallus). In four experiments, we presented chicks with a choice between either red (a colour typically associated with warning patterns) or green (a colour associated with palatable cryptic prey), volume-matched spheres (representing a generalised fruit shape) or frogs (representing an aposematic animal's shape). Chicks innately preferred green stimuli and avoided red. Chicks also preferred the shape of a frog over a sphere when both stimuli were green. However, no preference for frogs over spheres was present when stimuli were red. Male chicks that experienced a bitter taste of quinine immediately before the preference test showed a higher preference for green frog-shaped stimuli. Our results suggest that newly hatched chicks innately integrate colour and shape cues during decision making, and that this can be augmented by other sensory experiences. Innate and experience-based behaviour could confer a fitness advantage to novel aposematic prey, and favour the initial evolution of conspicuous colouration.

Weighted Geometric Mean (WGM) method: A new chromatic adaptation model.

The human visual system has undergone evolutionary changes to develop sophisticated mechanisms that enable stable color perception under varying illumination. These mechanisms are known as chromatic adaptation, a fundamental aspect of color vision. Chromatic adaptation can be divided into two categories: sensory adaptation, which involves automatic adjustments in the visual system, such as retinal gain control, in response to changes in the stimulus, and cognitive adaptation, which depends on the observer's knowledge of the scene and context. The geometric mean has been suggested to be the fundamental mathematical relationship that governs peripheral sensory adaptation. This paper proposes the WGM model, an advanced chromatic adaptation model based on a weighted geometric mean approach that can anticipate incomplete adaptation as it moves along the Planckian or Daylight locus. Compared with two other chromatic adaptation models (CAT16 and vK20), the WGM model is tested with different corresponding color data sets and found to be a significantly improvement while also predicting degree of adaptation (sensory and cognitive adaptation) in a physiologically plausible manner.