Developmental changes in colour constancy in a naturalistic object selection task.

When the illumination falling on a surface change, so does the reflected light. Despite this, adult observers are good at perceiving surfaces as relatively unchanging-an ability termed colour constancy. Very few studies have investigated colour constancy in infants, and even fewer in children. Here we asked whether there is a difference in colour constancy between children and adults; what the developmental trajectory is between six and 11 years; and whether the pattern of constancy across illuminations and reflectances differs between adults and children. To this end, we developed a novel, child-friendly computer-based object selection task. In this, observers saw a dragon's favourite sweet under a neutral illumination and picked the matching sweet from an array of eight seen under a different illumination (blue, yellow, red, or green). This set contained a reflectance match (colour constant; perfect performance) and a tristimulus match (colour inconstant). We ran two experiments, with two-dimensional scenes in one and three-dimensional renderings in the other. Twenty-six adults and 33 children took part in the first experiment; 26 adults and 40 children took part in the second. Children performed better than adults on this task, and their performance decreased with age in both experiments. We found differences across illuminations and sweets, but a similar pattern across both age groups. This unexpected finding might reflect a real decrease in colour constancy from childhood to adulthood, explained by developmental changes in the perceptual and cognitive mechanisms underpinning colour constancy, or differences in task strategies between children and adults. HIGHLIGHTS: Six- to 11-year-old children demonstrated better performance than adults on a colour constancy object selection task. Performance decreased with age over childhood. These findings may indicate development of cognitive strategies used to overcome automatic colour constancy mechanisms.

Correctly establishing evidence for cue combination via gains in sensory precision: Why the choice of comparator matters.

Studying how sensory signals from different sources (sensory cues) are integrated within or across multiple senses allows us to better understand the perceptual computations that lie at the foundation of adaptive behaviour. As such, determining the presence of precision gains - the classic hallmark of cue combination - is important for characterising perceptual systems, their development and functioning in clinical conditions. However, empirically measuring precision gains to distinguish cue combination from alternative perceptual strategies requires careful methodological considerations. Here, we note that the majority of existing studies that tested for cue combination either omitted this important contrast, or used an analysis approach that, unknowingly, strongly inflated false positives. Using simulations, we demonstrate that this approach enhances the chances of finding significant cue combination effects in up to 100% of cases, even when cues are not combined. We establish how this error arises when the wrong cue comparator is chosen and recommend an alternative analysis that is easy to implement but has only been adopted by relatively few studies. By comparing combined-cue perceptual precision with the best single-cue precision, determined for each observer individually rather than at the group level, researchers can enhance the credibility of their reported effects. We also note that testing for deviations from optimal predictions alone is not sufficient to ascertain whether cues are combined. Taken together, to correctly test for perceptual precision gains, we advocate for a careful comparator selection and task design to ensure that cue combination is tested with maximum power, while reducing the inflation of false positives.

Internal biases are linked to disrupted cue combination in children and adults.

Cue combination describes the use of two sensory cues together to increase perceptual precision. Internal relative bias describes a situation in which two cues to the same state of the world are perceived as signaling different states of the world on average. Current theory and evidence have difficulty accounting for many instances where cue combination is absent, such as in children under 10 years old, and in a variety of tasks. Here we show that internal relative biases between cues could be a key explanatory factor. Experiment 1, studying children's three-dimensional (slant) perception via disparity and texture, found a negative cross-sectional correlation between internal relative bias and cue combination behavior in 7- to 10-year-olds. Strikingly, children who had below-median levels of internal relative bias were able to combine cues, unlike the typical result for that age range. Experiment 2, studying adults' visual-auditory localization, found that cue combination behavior increased after an intervention designed to decrease internal relative bias. We interpret this as strong but preliminary evidence that internal relative bias can disrupt cue combination behavior. This provides a plausible mechanism to explain why children under 10 generally do not combine cues and why the audiovisual cue combination is so inconsistent in adults. Moving forward, we suggest that researchers who fail to find an expected cue combination effect should further investigate the possibility of issues with internal relative bias. Decreasing internal relative bias may also be an important goal for rehabilitation and sensory substitution or augmentation approaches to promoting efficient multisensory perception.

Newly learned shape-color associations show signatures of reliability-weighted averaging without forced fusion or a memory color effect.

Reliability-weighted averaging of multiple perceptual estimates (or cues) can improve precision. Research suggests that newly learned statistical associations can be rapidly integrated in this way for efficient decision-making. Yet, it remains unclear if the integration of newly learned statistics into decision-making can directly influence perception, rather than taking place only at the decision stage. In two experiments, we implicitly taught observers novel associations between shape and color. Observers made color matches by adjusting the color of an oval to match a simultaneously presented reference. As the color of the oval changed across trials, so did its shape according to a novel mapping of axis ratio to color. Observers showed signatures of reliability-weighted averaging-a precision improvement in both experiments and reweighting of the newly learned shape cue with changes in uncertainty in Experiment 2. To ask whether this was accompanied by perceptual effects, Experiment 1 tested for forced fusion by measuring color discrimination thresholds with and without incongruent novel cues. Experiment 2 tested for a memory color effect, observers adjusting the color of ovals with different axis ratios until they appeared gray. There was no evidence for forced fusion and the opposite of a memory color effect. Overall, our results suggest that the ability to quickly learn novel cues and integrate them with familiar cues is not immediately (within the short duration of our experiments and in the domain of color and shape) accompanied by common perceptual effects.

Central tendency biases must be accounted for to consistently capture Bayesian cue combination in continuous response data.

Observers in perceptual tasks are often reported to combine multiple sensory cues in a weighted average that improves precision-in some studies, approaching statistically optimal (Bayesian) weighting, but in others departing from optimality, or not benefitting from combined cues at all. To correctly conclude which combination rules observers use, it is crucial to have accurate measures of their sensory precision and cue weighting. Here, we present a new approach for accurately recovering these parameters in perceptual tasks with continuous responses. Continuous responses have many advantages, but are susceptible to a central tendency bias, where responses are biased towards the central stimulus value. We show that such biases lead to inaccuracies in estimating both precision gains and cue weightings, two key measures used to assess sensory cue combination. We introduce a method that estimates sensory precision by regressing continuous responses on targets and dividing the variance of the residuals by the squared slope of the regression line, "correcting-out" the error introduced by the central bias and increasing statistical power. We also suggest a complementary analysis that recovers the sensory cue weights. Using both simulations and empirical data, we show that the proposed methods can accurately estimate sensory precision and cue weightings in the presence of central tendency biases. We conclude that central tendency biases should be (and can easily be) accounted for to consistently capture Bayesian cue combination in continuous response data.

Merging familiar and new senses to perceive and act in space.

Our experience of the world seems to unfold seamlessly in a unitary 3D space. For this to be possible, the brain has to merge many disparate cognitive representations and sensory inputs. How does it do so? I discuss work on two key combination problems: coordinating multiple frames of reference (e.g. egocentric and allocentric), and coordinating multiple sensory signals (e.g. visual and proprioceptive). I focus on two populations whose spatial processing we can observe at a crucial stage of being configured and optimised: children, whose spatial abilities are still developing significantly, and naïve adults learning new spatial skills, such as sensing distance using auditory cues. The work uses a model-based approach to compare participants' behaviour with the predictions of alternative information processing models. This lets us see when and how-during development, and with experience-the perceptual-cognitive computations underpinning our experiences in space change. I discuss progress on understanding the limits of effective spatial computation for perception and action, and how lessons from the developing spatial cognitive system can inform approaches to augmenting human abilities with new sensory signals provided by technology.

Coding Locations Relative to One or Many Landmarks in Childhood.

Cognitive development studies how information processing in the brain changes over the course of development. A key part of this question is how information is represented and stored in memory. This study examined allocentric (world-based) spatial memory, an important cognitive tool for planning routes and interacting with the space around us. This is typically theorized to use multiple landmarks all at once whenever it operates. In contrast, here we show that allocentric spatial memory frequently operates over a limited spatial window, much less than the full proximal scene, for children between 3.5 and 8.5 years old. The use of multiple landmarks increases gradually with age. Participants were asked to point to a remembered target location after a change of view in immersive virtual reality. A k-fold cross-validation model-comparison selected a model where young children usually use the target location's vector to the single nearest landmark and rarely take advantage of the vectors to other nearby landmarks. The comparison models, which attempt to explain the errors as generic forms of noise rather than encoding to a single spatial cue, did not capture the distribution of responses as well. Parameter fits of this new single- versus multi-cue model are also easily interpretable and related to other variables of interest in development (age, executive function). Based on this, we theorize that spatial memory in humans develops through three advancing levels (but not strict stages): most likely to encode locations egocentrically (relative to the self), then allocentrically (relative to the world) but using only one landmark, and finally, most likely to encode locations relative to multiple parts of the scene.

Bayesian transfer in a complex spatial localization task.

Prior knowledge can help observers in various situations. Adults can simultaneously learn two location priors and integrate these with sensory information to locate hidden objects. Importantly, observers weight prior and sensory (likelihood) information differently depending on their respective reliabilities, in line with principles of Bayesian inference. Yet, there is limited evidence that observers actually perform Bayesian inference, rather than a heuristic, such as forming a look-up table. To distinguish these possibilities, we ask whether previously learned priors will be immediately integrated with a new, untrained likelihood. If observers use Bayesian principles, they should immediately put less weight on the new, less reliable, likelihood ("Bayesian transfer"). In an initial experiment, observers estimated the position of a hidden target, drawn from one of two distinct distributions, using sensory and prior information. The sensory cue consisted of dots drawn from a Gaussian distribution centered on the true location with either low, medium, or high variance; the latter introduced after block three of five to test for evidence of Bayesian transfer. Observers did not weight the cue (relative to the prior) significantly less in the high compared to medium variance condition, counter to Bayesian predictions. However, when explicitly informed of the different prior variabilities, observers placed less weight on the new high variance likelihood ("Bayesian transfer"), yet, substantially diverged from ideal. Much of this divergence can be captured by a model that weights sensory information, according only to internal noise in using the cue. These results emphasize the limits of Bayesian models in complex tasks.

Specular highlights improve color constancy when other cues are weakened.

Previous studies suggest that to achieve color constancy, the human visual system makes use of multiple cues, including a priori assumptions about the illumination ("daylight priors"). Specular highlights have been proposed to aid constancy, but the evidence for their usefulness is mixed. Here, we used a novel cue-combination approach to test whether the presence of specular highlights or the validity of a daylight prior improves illumination chromaticity estimates, inferred from achromatic settings, to determine whether and under which conditions either cue contributes to color constancy. Observers made achromatic settings within three-dimensional rendered scenes containing matte or glossy shapes, illuminated by either daylight or nondaylight illuminations. We assessed both the variability of these settings and their accuracy, in terms of the standard color constancy index (CCI). When a spectrally uniform background was present, neither CCIs nor variability improved with specular highlights or daylight illuminants (Experiment 1). When a Mondrian background was introduced, CCIs decreased overall but were higher for scenes containing glossy, as opposed to matte, shapes (Experiments 2 and 3). There was no overall reduction in variability of settings and no benefit for scenes illuminated by daylights. Taken together, these results suggest that the human visual system indeed uses specular highlights to improve color constancy but only when other cues, such as from the local surround, are weakened.

Prefrontal Dynamics Associated with Efficient Detours and Shortcuts: A Combined Functional Magnetic Resonance Imaging and Magnetoencenphalography Study.

Central to the concept of the "cognitive map" is that it confers behavioral flexibility, allowing animals to take efficient detours, exploit shortcuts, and avoid alluring, but unhelpful, paths. The neural underpinnings of such naturalistic and flexible behavior remain unclear. In two neuroimaging experiments, we tested human participants on their ability to navigate to a set of goal locations in a virtual desert island riven by lava, which occasionally spread to block selected paths (necessitating detours) or receded to open new paths (affording real shortcuts or false shortcuts to be avoided). Detours activated a network of frontal regions compared with shortcuts. Activity in the right dorsolateral PFC specifically increased when participants encountered tempting false shortcuts that led along suboptimal paths that needed to be differentiated from real shortcuts. We also report modulation in event-related fields and theta power in these situations, providing insight to the temporal evolution of response to encountering detours and shortcuts. These results help inform current models as to how the brain supports navigation and planning in dynamic environments.

Backtracking during navigation is correlated with enhanced anterior cingulate activity and suppression of alpha oscillations and the 'default-mode' network.

Successful navigation can require realizing the current path choice was a mistake and the best strategy is to retreat along the recent path: 'back-track'. Despite the wealth of studies on the neural correlates of navigation little is known about backtracking. To explore the neural underpinnings of backtracking we tested humans during functional magnetic resonance imaging on their ability to navigate to a set of goal locations in a virtual desert island riven by lava which constrained the paths that could be taken. We found that on a subset of trials, participants spontaneously chose to backtrack and that the majority of these choices were optimal. During backtracking, activity increased in frontal regions and the dorsal anterior cingulate cortex, while activity was suppressed in regions associated with the core default-mode network. Using the same task, magnetoencephalography and a separate group of participants, we found that power in the alpha band was significantly decreased immediately prior to such backtracking events. These results highlight the importance for navigation of brain networks previously identified in processing internally-generated errors and that such error-detection responses may involve shifting the brain from default-mode states to aid successful spatial orientation.

Long-term effect of gene therapy on Leber's congenital amaurosis.

BACKGROUND: Mutations in RPE65 cause Leber's congenital amaurosis, a progressive retinal degenerative disease that severely impairs sight in children. Gene therapy can result in modest improvements in night vision, but knowledge of its efficacy in humans is limited. METHODS: We performed a phase 1-2 open-label trial involving 12 participants to evaluate the safety and efficacy of gene therapy with a recombinant adeno-associated virus 2/2 (rAAV2/2) vector carrying the RPE65 complementary DNA, and measured visual function over the course of 3 years. Four participants were administered a lower dose of the vector, and 8 were administered a higher dose. In a parallel study in dogs, we investigated the relationship among vector dose, visual function, and electroretinography (ERG) findings. RESULTS: Improvements in retinal sensitivity were evident, to varying extents, in six participants for up to 3 years, peaking at 6 to 12 months after treatment and then declining. No associated improvement in retinal function was detected by means of ERG. Three participants had intraocular inflammation, and two had clinically significant deterioration of visual acuity. The reduction in central retinal thickness varied among participants. In dogs, RPE65 gene therapy with the same vector at lower doses improved vision-guided behavior, but only higher doses resulted in improvements in retinal function that were detectable with the use of ERG. CONCLUSIONS: Gene therapy with rAAV2/2 RPE65 vector improved retinal sensitivity, albeit modestly and temporarily. Comparison with the results obtained in the dog model indicates that there is a species difference in the amount of RPE65 required to drive the visual cycle and that the demand for RPE65 in affected persons was not met to the extent required for a durable, robust effect. (Funded by the National Institute for Health Research and others; ClinicalTrials.gov number, NCT00643747.).

Development of allocentric spatial recall from new viewpoints in virtual reality.

Using landmarks and other scene features to recall locations from new viewpoints is a critical skill in spatial cognition. In an immersive virtual reality task, we asked children 3.5-4.5 years old to remember the location of a target using various cues. On some trials they could use information from their own self-motion. On some trials they could use a view match. In the very hardest kind of trial, they were 'teleported' to a new viewpoint and could only use an allocentric spatial representation. This approach provides a strict test for allocentric coding (without either a matching viewpoint or self-motion information) while avoiding additional task demands in previous studies (it does not require them to deal with a small table-top environment or to manage stronger cue conflicts). Both the younger and older groups were able to point back at the target location better than chance when they could use view matching and/or self-motion, but allocentric recall was only seen in the older group (4.0-4.5). In addition, we only obtained evidence for a specific kind of allocentric recall in the older group: they tracked one major axis of the space significantly above chance, r(158) = .28, but not the other, r(158) = -.01. We conclude that there is a major qualitative change in coding for spatial recall around the fourth birthday, potentially followed by further development towards fully flexible recall from new viewpoints.

Effects of two-dimensional versus three-dimensional landmark geometry and layout on young children's recall of locations from new viewpoints.

Spatial memory is an important aspect of adaptive behavior and experience, providing both content and context to the perceptions and memories that we form in everyday life. Young children's abilities in this realm shift from mainly egocentric (self-based) to include allocentric (world-based) codings at around 4 years of age. However, information about the cognitive mechanisms underlying acquisition of these new abilities is still lacking. We examined allocentric spatial recall in 4.5- to 8.5-year-olds, looking for continuity with navigation as previously studied in 2- to 4-year-olds and other species. We specifically predicted an advantage for three-dimensional landmarks over two-dimensional ones and for recalling targets "in the middle" versus elsewhere. However, we did not find compelling evidence for either of these effects, and indeed some analyses even support the opposite of each of these conclusions. There were also no significant interactions with age. These findings highlight the incompleteness of our overall theories of the development of spatial cognition in general and allocentric spatial recall in particular. They also suggest that allocentric spatial recall involves processes that have separate behavioral characteristics from other cognitive systems involved in navigation earlier in life and in other species.

Observer models of perceptual development.

We agree with Rahnev & Denison (R&D) that to understand perception at a process level, we must investigate why performance sometimes deviates from idealised decision models. Recent research reveals that such deviations from optimality are pervasive during perceptual development. We argue that a full understanding of perception requires a model of how perceptual systems become increasingly optimised during development.

The impact of semantically congruent and incongruent visual information on auditory object recognition across development.

The ability to use different sensory signals in conjunction confers numerous advantages on perception. Multisensory perception in adults is influenced by factors beyond low-level stimulus properties such as semantic congruency. Sensitivity to semantic relations has been shown to emerge early in development; however, less is known about whether implementation of these associations changes with development or whether development in the representations themselves might modulate their influence. Here, we used a Stroop-like paradigm that requires participants to identify an auditory stimulus while ignoring a visual stimulus. Prior research shows that in adults visual distractors have more impact on processing of auditory objects than vice versa; however, this pattern appears to be inverted early in development. We found that children from 8years of age (and adults) gain a speed advantage from semantically congruent visual information and are disadvantaged by semantically incongruent visual information. At 6years of age, children gain a speed advantage for semantically congruent visual information but are not disadvantaged by semantically incongruent visual information (as compared with semantically unrelated visual information). Both children and adults were influenced by associations between auditory and visual stimuli, which they had been exposed to on only 12 occasions during the learning phase of the study. Adults showed a significant speed advantage over children for well-established associations but showed no such advantage for newly acquired pairings. This suggests that the influence of semantic associations on multisensory processing does not change with age but rather these associations become more robust and, in turn, more influential.

Risky visuomotor choices during rapid reaching in childhood.

Many everyday actions are implicit gambles because imprecisions in our visuomotor systems place probabilities on our success or failure. Choosing optimal action strategies involves weighting the costs and gains of potential outcomes by their corresponding probabilities, and requires stable representations of one's own imprecisions. How this ability is acquired during development in childhood when visuomotor skills change drastically is unknown. In a rewarded rapid reaching task, 6- to 11-year-old children followed 'risk-seeking' strategies leading to overly high point-loss. Adults' performance, in contrast, was close to optimal. Children's errors were not explained by distorted estimates of value or probability, but may reflect different action selection criteria or immature integration of value and probability information while planning movements. These findings provide a starting point for understanding children's risk-taking in everyday visuomotor situations when suboptimal choices can be dangerous. Moreover, children's risky visuomotor decisions mirror those reported for non-motor gambles, raising the possibility that common processes underlie development across decision-making domains.

Integration of audio-visual information for spatial decisions in children and adults.

In adults, decisions based on multisensory information can be faster and/or more accurate than those relying on a single sense. However, this finding varies significantly across development. Here we studied speeded responding to audio-visual targets, a key multisensory function whose development remains unclear. We found that when judging the locations of targets, children aged 4 to 12 years and adults had faster and less variable response times given auditory and visual information together compared with either alone. Comparison of response time distributions with model predictions indicated that children at all ages were integrating (pooling) sensory information to make decisions but that both the overall speed and the efficiency of sensory integration improved with age. The evidence for pooling comes from comparison with the predictions of Miller's seminal 'race model', as well as with a major recent extension of this model and a comparable 'pooling' (coactivation) model. The findings and analyses can reconcile results from previous audio-visual studies, in which infants showed speed gains exceeding race model predictions in a spatial orienting task (Neil et al., 2006) but children below 7 years did not in speeded reaction time tasks (e.g. Barutchu et al., 2009). Our results provide new evidence for early and sustained abilities to integrate visual and auditory signals for spatial localization from a young age.

Digital LED Pixels: Instructions for use and a characterization of their properties.

This article details how to control light emitting diodes (LEDs) using an ordinary desktop computer. By combining digitally addressable LEDs with an off-the-shelf microcontroller (Arduino), multiple LEDs can be controlled independently and with a high degree of temporal, chromatic, and luminance precision. The proposed solution is safe (can be powered by a 5-V battery), tested (has been used in published research), inexpensive (∼ $60 + $2 per LED), highly interoperable (can be controlled by any type of computer/operating system via a USB or Bluetooth connection), requires no prior knowledge of electrical engineering (components simply require plugging together), and uses widely available components for which established help forums already exist. Matlab code is provided, including a 'minimal working example' of use suitable for use by beginners. Properties of the recommended LEDs are also characterized, including their response time, luminance profile, and color gamut. Based on these, it is shown that the LEDs are highly stable in terms of both luminance and chromaticity, and do not suffer from issues of warm-up, chromatic shift, and slow response times associated with traditional CRT and LCD monitor technology.

Hearing Where the Eyes See: Children Use an Irrelevant Visual Cue When Localizing Sounds.

To reduce sensory uncertainty, humans combine cues from multiple senses. However, in everyday life, many co-occurring cues are irrelevant to the task at hand. How do humans know which cues to ignore? And does this ability change with development? This study shows the ability to ignore cross-modal irrelevant information develops late in childhood. Participants performed a sound discrimination task, with or without an irrelevant visual flash, presented synchronously in front of them. Adults ignored the irrelevant visual information, while 7- to 10-year-olds' responses were biased toward the flash location. The findings show that acquiring mature cue combination mechanisms is a multifaceted process that includes learning to ignore irrelevant cues, as well as to optimally combine relevant cues.