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Psychophysical evidence for the number sense, Philos Trans R Soc Lond B Biol Sci, 1740 (373).

Psychophysical evidence for the number sense, Philos Trans R Soc Lond B Biol Sci, 1740 (373). 

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It is now clear that most animals, including humans, possess an ability to rapidly estimate number. Some have questioned whether this ability arises from dedicated numerosity mechanisms, or is derived indirectly from judgements of density or other attributes. We describe a series of psychophysical experiments, largely using adaptation techniques, which demonstrate clearly the existence of a number sense in humans. The number sense is truly general, extending over space, time and sensory modality, and is closely linked with action. We further show that when multiple cues are present, numerosity emerges as the natural dimension for discrimination. However, when element density increases past a certain level, the elements become too crowded to parse, and the scene is perceived as a texture rather than array of elements. The two different regimes are psychophysically discriminable in that they follow distinct psychophysical laws, and show different dependencies on eccentricity, luminance levels and effects of perceptual grouping. The distinction is important, as the ability to discriminate numerosity, but not texture, correlates with formal maths skills.This article is part of the discussion meeting issue ‘The origins of numerical abilities’.

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Spatiotopic coding during dynamic head tilt, J Neurophysiol, 2 (117), 808-817.

Spatiotopic coding during dynamic head tilt, J Neurophysiol, 2 (117), 808-817.

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Humans maintain a stable representation of the visual world effortlessly, despite constant movements of the eyes, head, and body, across multiple planes. Whereas visual stability in the face of saccadic eye movements has been intensely researched, fewer studies have investigated retinal image transformations induced by head movements, especially in the frontal plane. Unlike head rotations in the horizontal and sagittal planes, tilting the head in the frontal plane is only partially counteracted by torsional eye movements and consequently induces a distortion of the retinal image to which we seem to be completely oblivious. One possible mechanism aiding perceptual stability is an active reconstruction of a spatiotopic map of the visual world, anchored in allocentric coordinates. To explore this possibility, we measured the positional motion aftereffect (PMAE; the apparent change in position after adaptation to motion) with head tilts of approximately 42 degrees between adaptation and test (to dissociate retinal from allocentric coordinates). The aftereffect was shown to have both a retinotopic and spatiotopic component. When tested with unpatterned Gaussian blobs rather than sinusoidal grating stimuli, the retinotopic component was greatly reduced, whereas the spatiotopic component remained. The results suggest that perceptual stability may be maintained at least partially through mechanisms involving spatiotopic coding.

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Active Vision: Dynamic Reformatting of Visual Information by the Saccade-Drift Cycle, Curr Biol, 9 (27), R341-R344.

 Active Vision: Dynamic Reformatting of Visual Information by the Saccade-Drift Cycle, Curr Biol, 9 (27), R341-R344.

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Visual processing depends on rapid parsing of global features followed by analysis of fine detail. A new study suggests that this transformation is enabled by a cycle of saccades and fixational drifts, which reformat visual input to match the spatiotemporal sensitivity of fast and slow neuronal pathways.

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Audio-visual temporal perception in children with restored hearing, Neuropsychologia, (99), 350-359.

Audio-visual temporal perception in children with restored hearing, Neuropsychologia, (99), 350-359.

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It is not clear how audio-visual temporal perception develops in children with restored hearing. In this study we measured temporal discrimination thresholds with an audio-visual temporal bisection task in 9 deaf children with restored audition, and 22 typically hearing children. In typically hearing children, audition was more precise than vision, with no gain in multisensory conditions (as previously reported in Gori et al. (2012b)). However, deaf children with restored audition showed similar thresholds for audio and visual thresholds and some evidence of gain in audio-visual temporal multisensory conditions. Interestingly, we found a strong correlation between auditory weighting of multisensory signals and quality of language: patients who gave more weight to audition had better language skills. Similarly, auditory thresholds for the temporal bisection task were also a good predictor of language skills. This result supports the idea that the temporal auditory processing is associated with language development.

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The light-from-above prior is intact in autistic children, J Exp Child Psychol, (161), 113-125.

The light-from-above prior is intact in autistic children, J Exp Child Psychol, (161), 113-125.

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Sensory information is inherently ambiguous. The brain disambiguates this information by anticipating or predicting the sensory environment based on prior knowledge. Pellicano and Burr (2012) proposed that this process may be atypical in autism and that internal assumptions, or “priors,” may be underweighted or less used than in typical individuals. A robust internal assumption used by adults is the “light-from-above” prior, a bias to interpret ambiguous shading patterns as if formed by a light source located above (and slightly to the left) of the scene. We investigated whether autistic children (n=18) use this prior to the same degree as typical children of similar age and intellectual ability (n=18). Children were asked to judge the shape (concave or convex) of a shaded hexagon stimulus presented in 24 rotations. We estimated the relation between the proportion of convex judgments and stimulus orientation for each child and calculated the light source location most consistent with those judgments. Children behaved similarly to adults in this task, preferring to assume that the light source was from above left, when other interpretations were compatible with the shading evidence. Autistic and typical children used prior assumptions to the same extent to make sense of shading patterns. Future research should examine whether this prior is as adaptable (i.e., modifiable with training) in autistic children as it is in typical adults.

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Autism is associated with reduced ability to interpret grasping actions of others, Sci Rep, 1 (7), 12687.

Autism is associated with reduced ability to interpret grasping actions of others, Sci Rep, 1 (7), 12687.

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We investigated the ability of children with ASD to discriminate a small cylinder from a large cube by observing a point-light movie of an actor grasping the object, either from an allocentric or egocentric viewpoint (observing action of others or self). Compared with typically developing controls, high functioning autistic children showed a strong selective impairment in this task, but only with the allocentric viewpoint, where thresholds were twice as high: egocentric thresholds were similar to age- and ability-matched controls. The magnitude of the impairment correlated strongly with the degree of symptomology (R2 = 0.5). The results suggest that children with ASD might be impaired in their ability to predict and infer the consequences of others’ movements, which could be related to the social-communicative deficits often reported in autism.

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Evidence for a number sense, Behavioral and Brain Sciences, (40), 18-19.

Evidence for a number sense, Behavioral and Brain Sciences, (40), 18-19.

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Numerosity is inherently confounded by related stimulus attributes such as density and area, and many studies have reported interactions of various strengths between area, density, and numerosity. However, direct measurements of sensitivity within the area-density-numerosity space show that numerosity emerges as the most spontaneous and sensitive dimension, strongly supporting the existence of a dedicated number sense.

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Area Prostriata in the Human Brain, Curr Biol, 19 (27), 3056-3060 e3053.

Area Prostriata in the Human Brain, Curr Biol, 19 (27), 3056-3060 e3053.

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Area prostriata is a cortical area at the fundus of the calcarine sulcus, described anatomically in humans [ 1–5 ] and other primates [ 6–9 ]. It is lightly myelinated and lacks the clearly defined six-layer structure evident throughout the cerebral cortex, with a thinner layer 4 and thicker layer 2 [ 10 ], characteristic of limbic cortex [ 11 ]. In the marmoset and rhesus monkey, area prostriata has cortical connections with MT+ [ 12 ], the cingulate motor cortex [ 8 ], the auditory cortex [ 13 ], the orbitofrontal cortex, and the frontal polar cortices [ 14 ]. Here we use functional magnetic resonance together with a wide-field projection system to study its functional properties in humans. With population receptive field mapping [ 15 ], we show that area prostriata has a complete representation of the visual field, clearly distinct from the adjacent area V1. As in the marmoset, the caudal-dorsal border of human prostriata—abutting V1—represents the far peripheral visual field, with eccentricities decreasing toward its rostral boundary. Area prostriata responds strongly to very fast motion, greater than 500°/s. The functional properties of area prostriata suggest that it may serve to alert the brain quickly to fast visual events, particularly in the peripheral visual field.

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Auditory Sensitivity and Decision Criteria Oscillate at Different Frequencies Separately for the Two Ears, Current Biology,

Auditory Sensitivity and Decision Criteria Oscillate at Different Frequencies Separately for the Two Ears, Current Biology,

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Many behavioral measures of visual perception fluctuate continually in a rhythmic manner, reflecting the influence of endogenous brain oscillations, particularly theta (~4–7 Hz) and alpha (~8–12 Hz) rhythms. However, it is unclear whether these oscillations are unique to vision or whether auditory performance also oscillates. Several studies report no oscillatory modulation in audition, while those with positive findings suffer from confounds relating to neural entrainment. Here, we used a bilateral pitch-identification task to investigate rhythmic fluctuations in auditory performance separately for the two ears and applied signal detection theory (SDT) to test for oscillations of both sensitivity and criterion (changes in decision boundary). Using uncorrelated dichotic white noise to induce a phase reset of oscillations, we demonstrate that, as with vision, both auditory sensitivity and criterion showed strong oscillations over time, at different frequencies: ~6 Hz (theta range) for sensitivity and ~8 Hz (low alpha range) for criterion, implying distinct underlying sampling mechanisms. The modulation in sensitivity in left and right ears was in antiphase, suggestive of attention-like mechanisms sampling alternatively from the two ears.

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Connecting Visual Objects Reduces Perceived Numerosity and Density for Sparse but not Dense Patterns, Journal of Numerical Cognition, 4 (3).

Connecting Visual Objects Reduces Perceived Numerosity and Density for Sparse but not Dense Patterns, Journal of Numerical Cognition, 4 (3).

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How is numerosity encoded by the visual system? – directly, or derived indirectly from texture density? We recently suggested that the numerosity of sparse patterns is encoded directly by dedicated mechanisms (which have been described as the “Approximate Number System” ANS). However, at high dot densities, where items become “crowded” and difficult to segregate, “texture-density” mechanisms come into play. Here we tested the importance of item segmentation on numerosity and density perception at various stimulus densities, by measuring the effect of connecting visual objects with thin lines. The results confirmed many previous studies showing that connecting items robustly reduces the apparent numerosity of patterns of moderate density. We further showed that the apparent density of moderate-density patterns is also reduced by connecting the dots. Crucially, we found that both these effects are strongly reduced at higher numerosities. Indeed for density judgments, the effect reverses, so connecting dots in dense patterns increases the apparent density (as expected from the physical characteristics). The results provide clear support for the three-regime framework of number perception, and suggest that for moderately sparse stimuli, numerosity – but not texture-density – is perceived directly.