Individual differences in human gaze behavior generalize from faces to objects.

Individuals differ in where they fixate on a face, with some looking closer to the eyes while others prefer the mouth region. These individual biases are highly robust, generalize from the lab to the outside world, and have been associated with social cognition and associated disorders. However, it is unclear, whether these biases are specific to faces or influenced by domain-general mechanisms of vision. Here, we juxtaposed these hypotheses by testing whether individual face fixation biases generalize to inanimate objects. We analyzed >1.8 million fixations toward faces and objects in complex natural scenes from 405 participants tested in multiple labs. Consistent interindividual differences in fixation positions were highly inter-correlated across faces and objects in all samples. Observers who fixated closer to the eye region also fixated higher on inanimate objects and vice versa. Furthermore, the inter-individual spread of fixation positions scaled with target size in precisely the same, non-linear manner for faces and objects. These findings contradict a purely domain-specific account of individual face gaze. Instead, they suggest significant domain-general contributions to the individual way we look at faces, a finding with potential relevance for basic vision, face perception, social cognition, and associated clinical conditions.

Faces in scenes attract rapid saccades.

During natural vision, the human visual system has to process upcoming eye movements in parallel to currently fixated stimuli. Saccades targeting isolated faces are known to have lower latency and higher velocity, but it is unclear how this generalizes to the natural cycle of saccades and fixations during free-viewing of complex scenes. To which degree can the visual system process high-level features of extrafoveal stimuli when they are embedded in visual clutter and compete with concurrent foveal input? Here, we investigated how free-viewing dynamics vary as a function of an upcoming fixation target while controlling for various low-level factors. We found strong evidence that face- versus inanimate object-directed saccades are preceded by shorter fixations and have higher peak velocity. Interestingly, the boundary conditions for these two effects are dissociated. The effect on fixation duration was limited to face saccades, which were small and followed the trajectory of the preceding one, early in a trial. This is reminiscent of a recently proposed model of perisaccadic retinotopic shifts of attention. The effect on saccadic velocity, however, extended to very large saccades and increased with trial duration. These findings suggest that multiple, independent mechanisms interact to process high-level features of extrafoveal targets and modulate the dynamics of natural vision.

Free viewing biases for complex scenes in preschoolers and adults.

Adult gaze behaviour towards naturalistic scenes is highly biased towards semantic object classes. Little is known about the ontological development of these biases, nor about group-level differences in gaze behaviour between adults and preschoolers. Here, we let preschoolers (n = 34, age 5 years) and adults (n = 42, age 18-59 years) freely view 40 complex scenes containing objects with different semantic attributes to compare their fixation behaviour. Results show that preschool children allocate a significantly smaller proportion of dwell time and first fixations on Text and instead fixate Faces, Touched objects, Hands and Bodies more. A predictive model of object fixations controlling for a range of potential confounds suggests that most of these differences can be explained by drastically reduced text salience in pre-schoolers and that this effect is independent of low-level salience. These findings are in line with a developmental attentional antagonism between text and body parts (touched objects and hands in particular), which resonates with recent findings regarding 'cortical recycling'. We discuss this and other potential mechanisms driving salience differences between children and adults.

Reading the mind in the nose.

Humans infer mental states and traits from faces and their expressions. Previous research focused on the role of eyes and mouths in this process, even though most observers fixate somewhere in between. Here, we report that ratings of the nose region are surprisingly consistent with those for the full face and even with subjective feelings of the nose bearer. We propose the nose as central to faces and their perception.

Quick, eyes! Isolated upper face regions but not artificial features elicit rapid saccades.

Human faces elicit faster saccades than objects or animals, resonating with the great importance of faces for our species. The underlying mechanisms are largely unclear. Here, we test two hypotheses based on previous findings. First, ultra-rapid saccades toward faces may not depend on the presence of the whole face, but the upper face region containing the eye region. Second, ultra-rapid saccades toward faces (and possibly face parts) may emerge from our extensive experience with this stimulus and thus extend to glasses and masks - artificial features frequently encountered as part of a face. To test these hypotheses, we asked 43 participants to complete a saccadic choice task, which contrasted images of whole, upper and lower faces, face masks, and glasses with car images. The resulting data confirmed ultra-rapid saccades for isolated upper face regions, but not for artificial facial features.

Individual fixation tendencies in person viewing generalize from images to videos.

Fixation behavior toward persons in static scenes varies considerably between individuals. However, it is unclear whether these differences generalize to dynamic stimuli. Here, we examined individual differences in the distribution of gaze across seven person features (i.e. body and face parts) in static and dynamic scenes. Forty-four participants freely viewed 700 complex static scenes followed by eight director-cut videos (28,925 frames). We determined the presence of person features using hand-delineated pixel masks (images) and Deep Neural Networks (videos). Results replicated highly consistent individual differences in fixation tendencies for all person features in static scenes and revealed that these tendencies generalize to videos. Individual fixation behavior for both, images and videos, fell into two anticorrelated clusters representing the tendency to fixate faces versus bodies. These results corroborate a low-dimensional space for individual gaze biases toward persons and show they generalize from images to videos.

Individual differences in looking at persons in scenes.

Individuals freely viewing complex scenes vary in their fixation behavior. The most prominent and reliable dimension of such individual differences is the tendency to fixate faces. However, much less is known about how observers distribute fixations across other body parts of persons in scenes and how individuals may vary in this regard. Here, we aimed to close this gap. We expanded a popular annotated stimulus set (Xu, Jiang, Wang, Kankanhalli, & Zhao, 2014) with 6,365 hand-delineated pixel masks for the body parts of 1,136 persons embedded in 700 complex scenes, which we publish with this article (https://osf.io/ynujz/). This resource allowed us to analyze the person-directed fixations of 103 participants freely viewing these scenes. We found large and reliable individual differences in the distribution of fixations across person features. Individual fixation tendencies formed two anticorrelated clusters, one for the eyes, head, and the inner face and one for body features (torsi, arms, legs, and hands). Interestingly, the tendency to fixate mouths was independent of the face cluster. Finally, our results show that observers who tend to avoid person fixations in general, particularly do so for the face region. These findings underscore the role of individual differences in fixation behavior and reveal underlying dimensions. They are further in line with a recently proposed push-pull relationship between cortical tuning for faces and bodies. They may also aid the comparison of special populations to general variation.

Pitfalls in post hoc analyses of population receptive field data.

Data binning involves grouping observations into bins and calculating bin-wise summary statistics. It can cope with overplotting and noise, making it a versatile tool for comparing many observations. However, data binning goes awry if the same observations are used for binning (selection) and contrasting (selective analysis). This creates circularity, biasing noise components and resulting in artifactual changes in the form of regression towards the mean. Importantly, these artifactual changes are a statistical necessity. Here, we use (null) simulations and empirical repeat data to expose this flaw in the scope of post hoc analyses of population receptive field data. In doing so, we reveal that the type of data analysis, data properties, and circular data cleaning are factors shaping the appearance of such artifactual changes. We furthermore highlight that circular data cleaning and circular sorting of change scores are selection practices that result in artifactual changes even without circular data binning. These pitfalls might have led to erroneous claims about changes in population receptive fields in previous work and can be mitigated by using independent data for selection purposes. Our evaluations highlight the urgency for us researchers to make the validation of analysis pipelines standard practice.

Characteristic fixation biases in Super-Recognizers.

Neurotypical observers show large and reliable individual differences in gaze behavior along several semantic object dimensions. Individual gaze behavior toward faces has been linked to face identity processing, including that of neurotypical observers. Here, we investigated potential gaze biases in Super-Recognizers (SRs), individuals with exceptional face identity processing skills. Ten SRs, identified with a novel conservative diagnostic framework, and 43 controls freely viewed 700 complex scenes depicting more than 5000 objects. First, we tested whether SRs and controls differ in fixation biases along four semantic dimensions: faces, text, objects being touched, and bodies. Second, we tested potential group differences in fixation biases toward eyes and mouths. Finally, we tested whether SRs fixate closer to the theoretical optimal fixation point for face identification. SRs showed a stronger gaze bias toward faces and away from text and touched objects, starting from the first fixation onward. Further, SRs spent a significantly smaller proportion of first fixations and dwell time toward faces on mouths but did not differ in dwell time or first fixations devoted to eyes. Face fixation of SRs also fell significantly closer to the theoretical optimal fixation point for identification, just below the eyes. Our findings suggest that reliable superiority for face identity processing is accompanied by early fixation biases toward faces and preferred saccadic landing positions close to the theoretical optimum for face identification. We discuss future directions to investigate the functional basis of individual fixation behavior and face identity processing ability.

The influence of familiarity on memory for faces and mask wearing.

During the COVID-19 pandemic, the wearing of face masks became mandatory in public areas or at workplaces in many countries. While offering protection, the coverage of large parts of our face (nose, mouth and chin) may have consequences for face recognition. This seems especially important in the context of contact tracing which can require memory of familiar and unfamiliar contacts and whether they were wearing a mask. In this study, we tested how well participants perform at remembering faces and whether they wore a mask, and if this depends on familiarity. Our results show that: (a) Participants remembered familiar faces better than unfamiliar ones, regardless of mask wearing. (b) Participants remembered unmasked faces better than masked faces, regardless of familiarity. (c) Participants were significantly worse at remembering whether an unfamiliar face was wearing a mask or not-even if they indicated remembering the face. (d) Participants showed a bias to indicate no memory of unfamiliar faces. (e) Participants showed a bias to indicate that unfamiliar faces wore a mask, even if they did not. In sum, it was harder to remember both, the identity of unfamiliar faces and whether they wore a mask. These findings have practical relevance for contact tracing and epidemic control.

Heritable functional architecture in human visual cortex.

How much of the functional organization of our visual system is inherited? Here we tested the heritability of retinotopic maps in human visual cortex using functional magnetic resonance imaging. We demonstrate that retinotopic organization shows a closer correspondence in monozygotic (MZ) compared to dizygotic (DZ) twin pairs, suggesting a partial genetic determination. Using population receptive field (pRF) analysis to examine the preferred spatial location and selectivity of these neuronal populations, we estimate a heritability around 10-20% for polar angle preferences and spatial selectivity, as quantified by pRF size, in extrastriate areas V2 and V3. Our findings are consistent with heritability in both the macroscopic arrangement of visual regions and stimulus tuning properties of visual cortex. This could constitute a neural substrate for variations in a range of perceptual effects, which themselves have been found to be at least partially genetically determined. These findings also add convergent evidence for the hypothesis that functional map topology is linked with cortical morphology.

Inferior Occipital Gyrus Is Organized along Common Gradients of Spatial and Face-Part Selectivity.

The ventral visual stream of the human brain is subdivided into patches with categorical stimulus preferences, like faces or scenes. However, the functional organization within these areas is less clear. Here, we used functional magnetic resonance imaging and vertex-wise tuning models to independently probe spatial and face-part preferences in the inferior occipital gyrus (IOG) of healthy adult males and females. The majority of responses were well explained by Gaussian population tuning curves for both retinotopic location and the preferred relative position within a face. Parameter maps revealed a common gradient of spatial and face-part selectivity, with the width of tuning curves drastically increasing from posterior to anterior IOG. Tuning peaks clustered more idiosyncratically but were also correlated across maps of visual and face space. Preferences for the upper visual field went along with significantly increased coverage of the upper half of the face, matching recently discovered biases in human perception. Our findings reveal a broad range of neural face-part selectivity in IOG, ranging from narrow to "holistic." IOG is functionally organized along this gradient, which in turn is correlated with retinotopy.SIGNIFICANCE STATEMENT Brain imaging has revealed a lot about the large-scale organization of the human brain and visual system. For example, occipital cortex contains map-like representations of the visual field, while neurons in ventral areas cluster into patches with categorical preferences, like faces or scenes. Much less is known about the functional organization within these areas. Here, we focused on a well established face-preferring area-the inferior occipital gyrus (IOG). A novel neuroimaging paradigm allowed us to map the retinotopic and face-part tuning of many recording sites in IOG independently. We found a steep posterior-anterior gradient of decreasing face-part selectivity, which correlated with retinotopy. This suggests the functional role of ventral areas is not uniform and may follow retinotopic "protomaps."

Practice modality of motor sequences impacts the neural signature of motor imagery.

Motor imagery is conceptualized as an internal simulation that uses motor-related parts of the brain as its substrate. Many studies have investigated this sharing of common neural resources between the two modalities of motor imagery and motor execution. They have shown overlapping but not identical activation patterns that thereby result in a modality-specific neural signature. However, it is not clear how far this neural signature depends on whether the imagined action has previously been practiced physically or only imagined. The present study aims to disentangle whether the neural imprint of an imagined manual pointing sequence within cortical and subcortical motor areas is determined by the nature of this prior practice modality. Each participant practiced two sequences physically, practiced two other sequences mentally, and did a behavioural pre-test without any further practice on a third pair of sequences. After a two-week practice intervention, participants underwent fMRI scans while imagining all six sequences. Behavioural data demonstrated practice-related effects as well as very good compliance with instructions. Functional MRI data confirmed the previously known motor imagery network. Crucially, we found that mental and physical practice left a modality-specific footprint during mental motor imagery. In particular, activation within the right posterior cerebellum was stronger when the imagined sequence had previously been practiced physically. We conclude that cerebellar activity is shaped specifically by the nature of the prior practice modality.

OSIEshort: A small stimulus set can reliably estimate individual differences in semantic salience.

Recent findings revealed consistent individual differences in fixation tendencies among observers free-viewing complex scenes. The present study aimed at (1) replicating these differences, and (2) testing whether they can be estimated using a shorter test. In total, 103 participants completed two eye-tracking sessions. The first session was a direct replication of the original study, but the second session used a smaller subset of images, optimized to capture individual differences efficiently. The first session replicated the large and consistent individual differences along five semantic dimensions observed in the original study. The second session showed that these differences can be estimated using about 40 to 100 images (depending on the tested dimension). Additional analyses revealed that only the first 2 seconds of viewing duration seem to be informative regarding these differences. Taken together, our findings suggest that reliable individual differences in semantic salience can be estimated with a test totaling less than 2 minutes of viewing duration.

Neural correlates of top-down modulation of haptic shape versus roughness perception.

Exploring an object's shape by touch also renders information about its surface roughness. It has been suggested that shape and roughness are processed distinctly in the brain, a result based on comparing brain activation when exploring objects that differed in one of these features. To investigate the neural mechanisms of top-down control on haptic perception of shape and roughness, we presented the same multidimensional objects but varied the relevance of each feature. Specifically, participants explored two objects that varied in shape (oblongness of cuboids) and surface roughness. They either had to compare the shape or the roughness in an alternative-forced-choice-task. Moreover, we examined whether the activation strength of the identified brain regions as measured by functional magnetic resonance imaging (fMRI) can predict the behavioral performance in the haptic discrimination task. We observed a widespread network of activation for shape and roughness perception comprising bilateral precentral and postcentral gyrus, cerebellum, and insula. Task-relevance of the object's shape increased activation in the right supramarginal gyrus (SMG/BA 40) and the right precentral gyrus (PreCG/BA 44) suggesting that activation in these areas does not merely reflect stimulus-driven processes, such as exploring shape, but also entails top-down controlled processes driven by task-relevance. Moreover, the strength of the SMG/PreCG activation predicted individual performance in the shape but not in the roughness discrimination task. No activation was found for the reversed contrast (roughness > shape). We conclude that macrogeometric properties, such as shape, can be modulated by top-down mechanisms whereas roughness, a microgeometric feature, seems to be processed automatically.

Individual differences in visual salience vary along semantic dimensions.

What determines where we look? Theories of attentional guidance hold that image features and task demands govern fixation behavior, while differences between observers are interpreted as a "noise-ceiling" that strictly limits predictability of fixations. However, recent twin studies suggest a genetic basis of gaze-trace similarity for a given stimulus. This leads to the question of how individuals differ in their gaze behavior and what may explain these differences. Here, we investigated the fixations of >100 human adults freely viewing a large set of complex scenes containing thousands of semantically annotated objects. We found systematic individual differences in fixation frequencies along six semantic stimulus dimensions. These differences were large (>twofold) and highly stable across images and time. Surprisingly, they also held for first fixations directed toward each image, commonly interpreted as "bottom-up" visual salience. Their perceptual relevance was documented by a correlation between individual face salience and face recognition skills. The set of reliable individual salience dimensions and their covariance pattern replicated across samples from three different countries, suggesting they reflect fundamental biological mechanisms of attention. Our findings show stable individual differences in salience along a set of fundamental semantic dimensions and that these differences have meaningful perceptual implications. Visual salience reflects features of the observer as well as the image.

Subjective vividness of motor imagery has a neural signature in human premotor and parietal cortex.

Motor imagery (MI) is the process in which subjects imagine executing a body movement with a strong kinesthetic component from a first-person perspective. The individual capacity to elicit such mental images is not universal but varies within and between subjects. Neuroimaging studies have shown that these inter-as well as intra-individual differences in imagery quality mediate the amplitude of neural activity during MI on a group level. However, these analyses were not sensitive to forms of representation that may not map onto a simple modulation of overall amplitude. Therefore, the present study asked how far the subjective impression of motor imagery vividness is reflected by a spatial neural code, and how patterns of neural activation in different motor regions relate to specific imagery impressions. During fMRI scanning, 20 volunteers imagined three different types of right-hand actions. After each imagery trial, subjects were asked to evaluate the perceived vividness of their imagery. A correlation analysis compared the rating differences and neural dissimilarity values of the rating groups separately for each region of interest. Results showed a significant positive correlation in the left vPMC and right IPL, indicating that these regions particularly reflect perceived imagery vividness in that similar rated trials evoke more similar neural patterns. A decoding analysis revealed that the vividness of the motor image related systematically to the action specificity of neural activation patterns in left vPMC and right SPL. Imagined actions accompanied by higher vividness ratings were significantly more distinguishable within these areas. Altogether, results showed that spatial patterns of neural activity within the human motor cortices reflect the individual vividness of imagined actions. Hence, the findings reveal a link between the subjective impression of motor imagery vividness and objective physiological markers.

How to Enhance the Power to Detect Brain-Behavior Correlations With Limited Resources.

Neuroscience has been diagnosed with a pervasive lack of statistical power and, in turn, reliability. One remedy proposed is a massive increase of typical sample sizes. Parts of the neuroimaging community have embraced this recommendation and actively push for a reallocation of resources toward fewer but larger studies. This is especially true for neuroimaging studies focusing on individual differences to test brain-behavior correlations. Here, I argue for a more efficient solution. Ad hoc simulations show that statistical power crucially depends on the choice of behavioral and neural measures, as well as on sampling strategy. Specifically, behavioral prescreening and the selection of extreme groups can ascertain a high degree of robust in-sample variance. Due to the low cost of behavioral testing compared to neuroimaging, this is a more efficient way of increasing power. For example, prescreening can achieve the power boost afforded by an increase of sample sizes from n = 30 to n = 100 at ∼5% of the cost. This perspective article briefly presents simulations yielding these results, discusses the strengths and limitations of prescreening and addresses some potential counter-arguments. Researchers can use the accompanying online code to simulate the expected power boost of prescreening for their own studies.

The optimal experimental design for multiple alternatives perceptual search.

Perceptual bias is inherent to all our senses, particularly in the form of visual illusions and aftereffects. However, many experiments measuring perceptual biases may be susceptible to nonperceptual factors, such as response bias and decision criteria. Here, we quantify how robust multiple alternative perceptual search (MAPS) is for disentangling estimates of perceptual biases from these confounding factors. First, our results show that while there are considerable response biases in our four-alternative forced-choice design, these are unrelated to perceptual biases estimates, and these response biases are not produced by the response modality (keyboard vs. mouse). We also show that perceptual bias estimates are reduced when feedback is given on each trial, likely due to feedback enabling observers to partially (and actively) correct for perceptual biases. However, this does not impact the reliability with which MAPS detects the presence of perceptual biases. Finally, our results show that MAPS can detect actual perceptual biases and is not a decisional bias towards choosing the target in the middle of the candidate stimulus distribution. In summary, researchers conducting a MAPS experiment should use a constant reference stimulus, but consider varying the mean of the candidate distribution. Ideally, they should not employ trial-wise feedback if the magnitude of perceptual biases is of interest.