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.

Different types of uncertainty in multisensory perceptual decision making.

Efficient decision-making requires accounting for sources of uncertainty (noise, or variability). Many studies have shown how the nervous system is able to account for perceptual uncertainty (noise, variability) that arises from limitations in its own abilities to encode perceptual stimuli. However, many other sources of uncertainty exist, reflecting for example variability in the behaviour of other agents or physical processes. Here we review previous studies on decision making under uncertainty as a function of the different types of uncertainty that the nervous system encounters, showing that noise that is intrinsic to the perceptual system can often be accounted for near-optimally (i.e. not statistically different from optimally), whereas accounting for other types of uncertainty can be much more challenging. As an example, we present a study in which participants made decisions about multisensory stimuli with both intrinsic (perceptual) and extrinsic (environmental) uncertainty and show that the nervous system accounts for these differently when making decisions: they account for internal uncertainty but under-account for external. Human perceptual systems may be well equipped to account for intrinsic (perceptual) uncertainty because, in principle, they have access to this. Accounting for external uncertainty is more challenging because this uncertainty must be learned. This article is part of the theme issue 'Decision and control processes in multisensory perception'.

Sensory augmentation for a rapid motor task in a multisensory environment.

BACKGROUND: Sensory substitution and augmentation systems (SSASy) seek to either replace or enhance existing sensory skills by providing a new route to access information about the world. Tests of such systems have largely been limited to untimed, unisensory tasks. OBJECTIVE: To test the use of a SSASy for rapid, ballistic motor actions in a multisensory environment. METHODS: Participants played a stripped-down version of air hockey in virtual reality with motion controls (Oculus Touch). They were trained to use a simple SASSy (novel audio cue) for the puck's location. They were tested on ability to strike an oncoming puck with the SASSy, degraded vision, or both. RESULTS: Participants coordinated vision and the SSASy to strike the target with their hand more consistently than with the best single cue alone, t(13) = 9.16, p <.001, Cohen's d = 2.448. CONCLUSIONS: People can adapt flexibly to using a SSASy in tasks that require tightly timed, precise, and rapid body movements. SSASys can augment and coordinate with existing sensorimotor skills rather than being limited to replacement use cases - in particular, there is potential scope for treating moderate vision loss. These findings point to the potential for augmenting human abilities, not only for static perceptual judgments, but in rapid and demanding perceptual-motor tasks.

Multisensory perception and decision-making with a new sensory skill.

It is clear that people can learn a new sensory skill-a new way of mapping sensory inputs onto world states. It remains unclear how flexibly a new sensory skill can become embedded in multisensory perception and decision-making. To address this, we trained typically sighted participants (N = 12) to use a new echo-like auditory cue to distance in a virtual world, together with a noisy visual cue. Using model-based analyses, we tested for key markers of efficient multisensory perception and decision-making with the new skill. We found that 12 of 14 participants learned to judge distance using the novel auditory cue. Their use of this new sensory skill showed three key features: (a) It enhanced the speed of timed decisions; (b) it largely resisted interference from a simultaneous digit span task; and (c) it integrated with vision in a Bayes-like manner to improve precision. We also show some limits following this relatively short training: Precision benefits were lower than the Bayes-optimal prediction, and there was no forced fusion of signals. We conclude that people already embed new sensory skills in flexible multisensory perception and decision-making after a short training period. A key application of these insights is to the development of sensory augmentation systems that can enhance human perceptual abilities in novel ways. The limitations we reveal (sub-optimality, lack of fusion) provide a foundation for further investigations of the limits of these abilities and their brain basis. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

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.

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.

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 novel cues to location are combined with familiar cues but not always with each other.

Mature perceptual systems can learn new arbitrary sensory signals (novel cues) to properties of the environment, but little is known about the extent to which novel cues are integrated into normal perception. In normal perception, multiple uncertain familiar cues are combined, often near-optimally (reliability-weighted averaging), to increase perceptual precision. We trained observers to use abstract novel cues to estimate horizontal locations of hidden objects on a monitor. In experiment 1, 4 groups of observers each learned to use a different novel cue. All groups benefited from a suboptimal but significant gain in precision using novel and familiar cues together after short-term training (3 ∼1.5 hr sessions), extending previous reports of novel-familiar cue combination. In experiment 2, we tested whether 2 novel cues may also be combined with each other. One pair of novel cues could be combined to improve precision but the other could not, at least not after 3 sessions of repeated training. Overall, our results provide extensive evidence that novel cues can be learned and combined with familiar cues to enhance perception, but mixed evidence for whether perceptual and decision-making systems can extend this ability to the combination of multiple novel cues with only short-term training. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

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.

An adaptive cue selection model of allocentric spatial reorientation.

After becoming disoriented, an organism must use the local environment to reorient and recover vectors to important locations. A new theory, adaptive combination, suggests that the information from different spatial cues is combined with Bayesian efficiency during reorientation. To test this further, we modified the standard reorientation paradigm to be more amenable to Bayesian cue combination analyses while still requiring reorientation in an allocentric (i.e., world-based, not egocentric) frame. Twelve adults and 20 children at ages 5 to 7 years old were asked to recall locations in a virtual environment after a disorientation. Results were not consistent with adaptive combination. Instead, they are consistent with the use of the most useful (nearest) single landmark in isolation. We term this adaptive selection. Experiment 2 suggests that adults also use the adaptive selection method when they are not disoriented but are still required to use a local allocentric frame. This suggests that the process of recalling a location in the allocentric frame is typically guided by the single most useful landmark rather than a Bayesian combination of landmarks. These results illustrate that there can be important limits to Bayesian theories of the cognition, particularly for complex tasks such as allocentric recall. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

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.

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.

Combining the senses: The role of experience- and task-dependent mechanisms in the development of audiovisual simultaneity perception.

The brain's ability to integrate information from the different senses is essential for decreasing sensory uncertainty and ultimately limiting errors. Temporal correspondence is one of the key processes that determines whether information from different senses will be integrated and is influenced by both experience- and task-dependent mechanisms in adults. Here we investigated the development of both task- and experience-dependent temporal mechanisms by testing 7-8-year-old children, 10-11-year-old children, and adults in two tasks (simultaneity judgment, temporal order judgment) using audiovisual stimuli with differing degrees of association based on prior experience (low for beep-flash vs. high for face-voice). By fitting an independent channels model to the data, we found that while the experience-dependent mechanism of audiovisual simultaneity perception is already adult-like in 10-11-year-old children, the task-dependent mechanism is still not. These results indicate that differing maturation rates of experience-dependent and task-dependent mechanisms underlie the development of multisensory integration. Understanding this development has important implications for clinical and educational interventions. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

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.

The Difficulty of Effectively Using Allocentric Prior Information in a Spatial Recall Task.

Prior information represents the long-term statistical structure of an environment. For example, colds develop more often than throat cancer, making the former a more likely diagnosis for a sore throat. There is ample evidence for effective use of prior information during a variety of perceptual tasks, including the ability to recall locations using an egocentric (self-based) frame. However, it is not yet known if people can use prior information effectively when using an allocentric (world-based) frame. Forty-eight adults were shown sixty sets of three target locations in a sparse virtual environment with three beacons. The targets were drawn from one of four prior distributions. They were then asked to point to the targets after a delay and a change in perspective. While searches were biased towards the beacons, we did not find any evidence that participants successfully exploited the prior distributions of targets. These results suggest that allocentric reasoning does not conform to normative Bayesian models: we saw no evidence for use of priors in our cognitively-complex (allocentric) task, unlike in previous, simpler (egocentric) recall tasks. It is possible that this reflects the high biological cost of processing precise allocentric information.

Boundaries in spatial cognition: Looking like a boundary is more important than being a boundary.

Large walls and other typical boundaries strongly influence neural activity related to navigation and the representations of spatial layouts. They are also major aids to reliable navigation behavior in young children and nonhuman animals. Is this because they are physical boundaries (barriers to movement), or because they present certain visual features, such as visually extended 3D surfaces? Here, these 2 factors were dissociated by using immersive virtual reality and real boundaries. Eighty adults recalled target locations in 1 of 4 environments: plywood, where a virtual wall coincided with a large piece of real plywood; pass through, where the virtual wall coincided with empty space and participants could pass through it; pass over, where the virtual wall was projected downward to be visible underneath a transparent floor; and cones, where the walls were replaced with traffic cones. One condition had features that were boundaries and looked like boundaries (plywood); 2 had features that were not boundaries but looked like boundaries (pass over/through); and 1 had features that were not boundaries and did not look like boundaries (cones). The precision and bias of responses changed only as a function of looking like a boundary. This suggests that variations in spatial coding are more closely linked to the visual properties of environmental layouts than to whether they contain physical boundaries (barriers to movement). (PsycInfo Database Record (c) 2020 APA, all rights reserved).

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.

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.

Sensory cue combination in children under 10 years of age.

Cue combination occurs when two independent noisy perceptual estimates are merged together as a weighted average, creating a unified estimate that is more precise than either single estimate alone. Surprisingly, this effect has not been demonstrated compellingly in children under the age of 10 years, in contrast with the array of other multisensory skills that children show even in infancy. Instead, across a wide variety of studies, precision with both cues is no better than the best single cue - and sometimes worse. Here we provide the first consistent evidence of cue combination in children from 7 to 10 years old. Across three experiments, participants showed evidence of a bimodal precision advantage (Experiments 1a and 1b) and the majority were best-fit by a combining model (Experiment 2). The task was to localize a target horizontally with a binaural audio cue and a noisy visual cue in immersive virtual reality. Feedback was given as well, which could both (a) help participants judge how reliable each cue is and (b) help correct between-cue biases that might prevent cue combination. Crucially, our results show cue combination when feedback is only given on single cues - therefore, combination itself was not a strategy learned via feedback. We suggest that children at 7-10 years old are capable of cue combination in principle, but must have sufficient representations of reliabilities and biases in their own perceptual estimates as relevant to the task, which can be facilitated through task-specific feedback.

Efficient visual information sampling develops late in childhood.

It is often unclear which course of action gives the best outcome. We can reduce this uncertainty by gathering more information, but gathering information always comes at a cost. For example, a sports player waiting too long to judge a ball's trajectory will run out of time to intercept it. Efficient samplers must therefore optimize a trade-off: when the costs of collecting further information exceed the expected benefits, they should stop sampling and start acting. In visually guided tasks, adults can make these trade-offs efficiently, correctly balancing any reductions in visuomotor uncertainty against cost factors associated with increased sampling. To investigate how this ability develops during childhood, we tested 6- to 11-year-olds, adolescents, and adults on a visual localization task in which the costs and benefits of sampling were formalized in a quantitative framework. This allowed us to compare participants to each other and to an ideal observer who maximizes expected reward. Visual sampling became substantially more efficient between 6 and 11 years, converging onto adult performance in adolescence. Younger children systematically undersampled information relative to the ideal observer and varied their sampling strategy more. Further analyses suggested that young children used a suboptimal decision rule that insufficiently accounted for the chance of task failure, in line with a late developing ability to compute with probabilities and costs. We therefore propose that late development of efficient information sampling, a crucial element of real-world decision-making under risk, may form an important component of suboptimality in child perception, action, and decision-making. (PsycINFO Database Record (c) 2019 APA, all rights reserved).