Americans weigh an attended emotion more than Koreans in overall mood judgments.

Face ensemble coding is the perceptual ability to create a quick and overall impression of a group of faces, triggering social and behavioral motivations towards other people (approaching friendly people or avoiding an angry mob). Cultural differences in this ability have been reported, such that Easterners are better at face ensemble coding than Westerners are. The underlying mechanism has been attributed to differences in processing styles, with Easterners allocating attention globally, and Westerners focusing on local parts. However, the remaining question is how such default attention mode is influenced by salient information during ensemble perception. We created visual displays that resembled a real-world social setting in which one individual in a crowd of different faces drew the viewer's attention while the viewer judged the overall emotion of the crowd. In each trial, one face in the crowd was highlighted by a salient cue, capturing spatial attention before the participants viewed the entire group. American participants' judgment of group emotion more strongly weighed the attended individual face than Korean participants, suggesting a greater influence of local information on global perception. Our results showed that different attentional modes between cultural groups modulate social-emotional processing underlying people's perceptions and attributions.

The effect of masks on the emotion perception of a facial crowd.

The present study investigated the effect of facial masks on people's ability to perceive emotions in crowds. We presented faces with the bottom halves occluded by masks or full faces without occlusion. In two sequentially presented crowds, we varied the number of faces, emotional valence, and intensity of facial expressions, examining the impact of masks on the perception of crowd emotion. Participants reported which of the two crowds they would avoid based on the crowds' average emotions. The participants' ability to judge the average emotion of a crowd, especially a crowd expressing happiness, was impaired when the crowd wore masks. For faces covered by masks, crowd emotion judgments were more negatively biased than those without masks. However, participants could still distinguish the emotional intensities of a crowd wearing masks above chance. Additionally, participants responded more quickly to a crowd with more people without compromising accuracy, despite the perceptual challenges imposed by facial masks. Our results suggest that under ambiguous social situations in which individuals' emotions are partially hidden by masks, a large group may provide stronger social cues than a small group, thereby promoting communication and regulating social behaviors.

Inconsistent attentional contexts impair relearning following gradual visuomotor adaptation.

One of the brain's primary functions is to promote actions in dynamic, distracting environments. Because distractions divert attention from our primary goals, we must learn to maintain accurate actions under sensory and cognitive distractions. Visuomotor adaptation is a learning process that restores performance when sensorimotor capacities or environmental conditions are abruptly or gradually altered. Prior work showed that learning to counteract an abrupt perturbation under a particular single- or dual-task setting (i.e., attentional context) was associated with better recall under the same conditions. This suggested that the attentional context was encoded during adaptation and used as a recall cue. The current study investigated whether the attentional context (i.e., single vs. dual task) also affected adaptation and recall to a gradual perturbation, which limited awareness of movement errors. During adaptation, participants moved a cursor to a target while learning to counteract a visuomotor rotation that increased from 0° to 45° by 0.3° each trial, with or without performing a secondary task. Relearning was impaired when the attentional context was different between adaptation and recall (experiment 1), even when the exposure to the attentional context was limited to the early or late half of adaptation (experiment 2). Changing the secondary task did not affect relearning, indicating that the attentional context, rather than specific stimuli or tasks, was associated with better recall performance (experiment 3). These findings highlight the importance of cognitive factors, such as attention, in visuomotor adaptation and have implications for learning and rehabilitation paradigms.NEW & NOTEWORTHY Adaptation acquired under single- or dual-task setting, which created an undivided or divided attentional context, respectively, was impaired when relearning occurred under different conditions (i.e., shifting from a dual to single task). Changes to the attentional context impaired relearning when the initial adaptation was to a gradual perturbation. Explicit awareness of the perturbation was not necessary for this effect to be robust, nor was the effect attributable to changes in the secondary task requirements.

Differential neurodynamics and connectivity in the dorsal and ventral visual pathways during perception of emotional crowds and individuals: a MEG study.

Reading the prevailing emotion of groups of people ("crowd emotion") is critical to understanding their overall intention and disposition. It alerts us to potential dangers, such as angry mobs or panicked crowds, giving us time to escape. A critical aspect of processing crowd emotion is that it must occur rapidly, because delays often are costly. Although knowing the timing of neural events is crucial for understanding how the brain guides behaviors using coherent signals from a glimpse of multiple faces, this information is currently lacking in the literature on face ensemble coding. Therefore, we used magnetoencephalography to examine the neurodynamics in the dorsal and ventral visual streams and the periamygdaloid cortex to compare perception of groups of faces versus individual faces. Forty-six participants compared two groups of four faces or two individual faces with varying emotional expressions and chose which group or individual they would avoid. We found that the dorsal stream was activated as early as 68 msec after the onset of stimuli containing groups of faces. In contrast, the ventral stream was activated later and predominantly for individual face stimuli. The latencies of the dorsal stream activation peaks correlated with participants' response times for facial crowds. We also found enhanced connectivity earlier between the periamygdaloid cortex and the dorsal stream regions for crowd emotion perception. Our findings suggest that ensemble coding of facial crowds proceeds rapidly and in parallel by engaging the dorsal stream to mediate adaptive social behaviors, via a distinct route from single face perception.

An explicit investigation of the roles that feature distributions play in rapid visual categorization.

Ensemble representations are often described as efficient tools when summarizing features of multiple similar objects as a group. However, it can sometimes be more useful not to compute a single summary description for all of the objects if they are substantially different, for example when they belong to entirely different categories. It was proposed that the visual system can efficiently use the distributional information of ensembles to decide whether simultaneously displayed items belong to single or several different categories. Here we directly tested how the feature distribution of items in a visual array affects an ability to discriminate individual items (Experiment 1) and sets (Experiments 2-3) when participants were instructed explicitly to categorize individual objects based on the median of size distribution. We varied the width (narrow or fat) as well as the shape (smooth or two-peaked) of distributions in order to manipulate the ease of ensemble extraction from the items. We found that observers unintentionally relied on the grand mean as a natural categorical boundary and that their categorization accuracy increased as a function of the size differences among individual items and a function of their separation from the grand mean. For ensembles drawn from two-peaked size distributions, participants showed better categorization performance. They were more accurate at judging within-category ensemble properties in other dimensions (centroid and orientation) and less biased by superset statistics. This finding corroborates the idea that the two-peaked feature distributions support the "segmentability" of spatially intermixed sets of objects. Our results emphasize important roles of ensemble statistics (mean, range, distribution shape) in explicit visual categorization.

Fast saccadic and manual responses to faces presented to the koniocellular visual pathway.

The parallel pathways of the human visual system differ in their tuning to luminance, color, and spatial frequency. These attunements recently have been shown to propagate to differential processing of higher-order stimuli, facial threat cues, in the magnocellular (M) and parvocellular (P) pathways, with greater sensitivity to clear and ambiguous threat, respectively. The role of the third, koniocellular (K) pathway in facial threat processing, however, remains unknown. To address this gap in knowledge, we briefly presented peripheral face stimuli psychophysically biased towards M, P, or K pathways. Observers were instructed to report via a key-press whether the face was angry or neutral while their eye movements and manual responses were recorded. We found that short-latency saccades were made more frequently to faces presented in the K channel than to P or M channels. Saccade latencies were not significantly modulated by expressive and identity cues. In contrast, manual response latencies and accuracy were modulated by both pathway biasing and by interactions of facial expression with facial masculinity, such that angry male faces elicited the fastest, and angry female faces, the least accurate, responses. We conclude that face stimuli can evoke fast saccadic and manual responses when projected to the K pathway.

Differential magnocellular versus parvocellular pathway contributions to the combinatorial processing of facial threat.

Recently, speed of presentation of facially expressive stimuli was found to influence the processing of compound threat cues (e.g., anger/fear/gaze). For instance, greater amygdala responses were found to clear (e.g., direct gaze anger/averted gaze fear) versus ambiguous (averted gaze anger/direct gaze fear) combinations of threat cues when rapidly presented (33 and 300ms), but greater to ambiguous versus clear threat cues when presented for more sustained durations (1, 1.5, and 2s). A working hypothesis was put forth (Adams et al., 2012) that these effects were due to differential magnocellular versus parvocellular pathways contributions to the rapid versus sustained processing of threat, respectively. To test this possibility directly here, we restricted visual stream processing in the fMRI environment using facially expressive stimuli specifically designed to bias visual input exclusively to the magnocellular versus parvocellular pathways. We found that for magnocellular-biased stimuli, activations were predominantly greater to clear versus ambiguous threat-gaze pairs (on par with that previously found for rapid presentations of threat cues), whereas activations to ambiguous versus clear threat-gaze pairs were greater for parvocellular-biased stimuli (on par with that previously found for sustained presentations). We couch these findings in an adaptive dual process account of threat perception and highlight implications for other dual process models within psychology.

Magnocellular and parvocellular pathway contributions to facial threat cue processing.

Human faces evolved to signal emotions, with their meaning contextualized by eye gaze. For instance, a fearful expression paired with averted gaze clearly signals both presence of threat and its probable location. Conversely, direct gaze paired with facial fear leaves the source of the fear-evoking threat ambiguous. Given that visual perception occurs in parallel streams with different processing emphases, our goal was to test a recently developed hypothesis that clear and ambiguous threat cues would differentially engage the magnocellular (M) and parvocellular (P) pathways, respectively. We employed two-tone face images to characterize the neurodynamics evoked by stimuli that were biased toward M or P pathways. Human observers (N = 57) had to identify the expression of fearful or neutral faces with direct or averted gaze while their magnetoencephalogram was recorded. Phase locking between the amygdaloid complex, orbitofrontal cortex (OFC) and fusiform gyrus increased early (0-300 ms) for M-biased clear threat cues (averted-gaze fear) in the ß-band (13-30 Hz) while P-biased ambiguous threat cues (direct-gaze fear) evoked increased θ (4-8 Hz) phase locking in connections with OFC of the right hemisphere. We show that M and P pathways are relatively more sensitive toward clear and ambiguous threat processing, respectively, and characterize the neurodynamics underlying emotional face processing in the M and P pathways.

Spatial and feature-based attention to expressive faces.

Facial emotion is an important cue for deciding whether an individual is potentially helpful or harmful. However, facial expressions are inherently ambiguous and observers typically employ other cues to categorize emotion expressed on the face, such as race, sex, and context. Here, we explored the effect of increasing or reducing different types of uncertainty associated with a facial expression that is to be categorized. On each trial, observers responded according to the emotion and location of a peripherally presented face stimulus and were provided with either: (1) no information about the upcoming face; (2) its location; (3) its expressed emotion; or (4) both its location and emotion. While cueing emotion or location resulted in faster response times than cueing unpredictive information, cueing face emotion alone resulted in faster responses than cueing face location alone. Moreover, cueing both stimulus location and emotion resulted in a superadditive reduction of response times compared with cueing location or emotion alone, suggesting that feature-based attention to emotion and spatially selective attention interact to facilitate perception of face stimuli. While categorization of facial expressions was significantly affected by stable identity cues (sex and race) in the face, we found that these interactions were eliminated when uncertainty about facial expression, but not spatial uncertainty about stimulus location, was reduced by predictive cueing. This demonstrates that feature-based attention to facial expression greatly attenuates the need to rely on stable identity cues to interpret facial emotion.

Sex-related differences in behavioral and amygdalar responses to compound facial threat cues.

During face perception, we integrate facial expression and eye gaze to take advantage of their shared signals. For example, fear with averted gaze provides a congruent avoidance cue, signaling both threat presence and its location, whereas fear with direct gaze sends an incongruent cue, leaving threat location ambiguous. It has been proposed that the processing of different combinations of threat cues is mediated by dual processing routes: reflexive processing via magnocellular (M) pathway and reflective processing via parvocellular (P) pathway. Because growing evidence has identified a variety of sex differences in emotional perception, here we also investigated how M and P processing of fear and eye gaze might be modulated by observer's sex, focusing on the amygdala, a structure important to threat perception and affective appraisal. We adjusted luminance and color of face stimuli to selectively engage M or P processing and asked observers to identify emotion of the face. Female observers showed more accurate behavioral responses to faces with averted gaze and greater left amygdala reactivity both to fearful and neutral faces. Conversely, males showed greater right amygdala activation only for M-biased averted-gaze fear faces. In addition to functional reactivity differences, females had proportionately greater bilateral amygdala volumes, which positively correlated with behavioral accuracy for M-biased fear. Conversely, in males only the right amygdala volume was positively correlated with accuracy for M-biased fear faces. Our findings suggest that M and P processing of facial threat cues is modulated by functional and structural differences in the amygdalae associated with observer's sex.

Neurodynamics and connectivity during facial fear perception: The role of threat exposure and signal congruity.

Fearful faces convey threat cues whose meaning is contextualized by eye gaze: While averted gaze is congruent with facial fear (both signal avoidance), direct gaze (an approach signal) is incongruent with it. We have previously shown using fMRI that the amygdala is engaged more strongly by fear with averted gaze during brief exposures. However, the amygdala also responds more to fear with direct gaze during longer exposures. Here we examined previously unexplored brain oscillatory responses to characterize the neurodynamics and connectivity during brief (~250 ms) and longer (~883 ms) exposures of fearful faces with direct or averted eye gaze. We performed two experiments: one replicating the exposure time by gaze direction interaction in fMRI (N = 23), and another where we confirmed greater early phase locking to averted-gaze fear (congruent threat signal) with MEG (N = 60) in a network of face processing regions, regardless of exposure duration. Phase locking to direct-gaze fear (incongruent threat signal) then increased significantly for brief exposures at ~350 ms, and at ~700 ms for longer exposures. Our results characterize the stages of congruent and incongruent facial threat signal processing and show that stimulus exposure strongly affects the onset and duration of these stages.

Cross-cultural and hemispheric laterality effects on the ensemble coding of emotion in facial crowds.

In many social situations, we make a snap judgment about crowds of people relying on their overall mood (termed "crowd emotion"). Although reading crowd emotion is critical for interpersonal dynamics, the sociocultural aspects of this process have not been explored. The current study examined how culture modulates the processing of crowd emotion in Korean and American observers. Korean and American (non-East Asian) participants were briefly presented with two groups of faces that were individually varying in emotional expressions and asked to choose which group between the two they would rather avoid. We found that Korean participants were more accurate than American participants overall, in line with the framework on cultural viewpoints: Holistic versus analytic processing in East Asians versus Westerners. Moreover, we found a speed advantage for other-race crowds in both cultural groups. Finally, we found different hemispheric lateralization patterns: American participants were more accurate to perceive the facial crowd to be avoided when it was presented in the left visual field than the right visual field, indicating a right hemisphere advantage for processing crowd emotion of both European American and Korean facial crowds. However, Korean participants showed weak or nonexistent laterality effects, with a slight right hemisphere advantage for European American facial crowds and no advantage in perceiving Korean facial crowds. Instead, Korean participants showed positive emotion bias for own-race faces. This work suggests that culture plays a role in modulating our crowd emotion perception of groups of faces and responses to them.

Differential hemispheric and visual stream contributions to ensemble coding of crowd emotion.

In crowds, where scrutinizing individual facial expressions is inefficient, humans can make snap judgments about the prevailing mood by reading "crowd emotion". We investigated how the brain accomplishes this feat in a set of behavioral and fMRI studies. Participants were asked to either avoid or approach one of two crowds of faces presented in the left and right visual hemifields. Perception of crowd emotion was improved when crowd stimuli contained goal-congruent cues and was highly lateralized to the right hemisphere. The dorsal visual stream was preferentially activated in crowd emotion processing, with activity in the intraparietal sulcus and superior frontal gyrus predicting perceptual accuracy for crowd emotion perception, whereas activity in the fusiform cortex in the ventral stream predicted better perception of individual facial expressions. Our findings thus reveal significant behavioral differences and differential involvement of the hemispheres and the major visual streams in reading crowd versus individual face expressions.

Observer's anxiety facilitates magnocellular processing of clear facial threat cues, but impairs parvocellular processing of ambiguous facial threat cues.

Facial expression and eye gaze provide a shared signal about threats. While a fear expression with averted gaze clearly points to the source of threat, direct-gaze fear renders the source of threat ambiguous. Separable routes have been proposed to mediate these processes, with preferential attunement of the magnocellular (M) pathway to clear threat, and of the parvocellular (P) pathway to threat ambiguity. Here we investigated how observers' trait anxiety modulates M- and P-pathway processing of clear and ambiguous threat cues. We scanned subjects (N = 108) widely ranging in trait anxiety while they viewed fearful or neutral faces with averted or directed gaze, with the luminance and color of face stimuli calibrated to selectively engage M- or P-pathways. Higher anxiety facilitated processing of clear threat projected to M-pathway, but impaired perception of ambiguous threat projected to P-pathway. Increased right amygdala reactivity was associated with higher anxiety for M-biased averted-gaze fear, while increased left amygdala reactivity was associated with higher anxiety for P-biased, direct-gaze fear. This lateralization was more pronounced with higher anxiety. Our findings suggest that trait anxiety differentially affects perception of clear (averted-gaze fear) and ambiguous (direct-gaze fear) facial threat cues via selective engagement of M and P pathways and lateralized amygdala reactivity.

Long lasting attentional-context dependent visuomotor memory.

Using a dual-task paradigm, we recently reported that visuomotor adaptation acquired under distraction of a secondary attention-demanding discrimination task could be remembered only when a similar distraction was present. In contrast, when tested without the distracting task, performance reverted to untrained levels (Song & Bédard, 2015). Here, we demonstrated that this newfound paradoxical benefit of consistent dual-task context lasts over 1 day, such that visuomotor memory retrieval is enhanced under conditions where it is more difficult to engage in attentional selection of the motor task. Furthermore, this long-term effect was evident even when the task type or sensory modality of the secondary task differed between initial adaptation and the delayed recall on the next day. We conclude that attentional diversion by performing a dual-task forms a long-term vital context for visuomotor memory independent of external contexts without taxing capacity limited attention. (PsycINFO Database Record

PsiMLE: A maximum-likelihood estimation approach to estimating psychophysical scaling and variability more reliably, efficiently, and flexibly.

A simple and popular psychophysical model-usually described as overlapping Gaussian tuning curves arranged along an ordered internal scale-is capable of accurately describing both human and nonhuman behavioral performance and neural coding in magnitude estimation, production, and reproduction tasks for most psychological dimensions (e.g., time, space, number, or brightness). This model traditionally includes two parameters that determine how a physical stimulus is transformed into a psychological magnitude: (1) an exponent that describes the compression or expansion of the physical signal into the relevant psychological scale (ß), and (2) an estimate of the amount of inherent variability (often called internal noise) in the Gaussian activations along the psychological scale (σ). To date, linear slopes on log-log plots have traditionally been used to estimate ß, and a completely separate method of averaging coefficients of variance has been used to estimate σ. We provide a respectful, yet critical, review of these traditional methods, and offer a tutorial on a maximum-likelihood estimation (MLE) and a Bayesian estimation method for estimating both ß and σ [PsiMLE(ß,σ)], coupled with free software that researchers can use to implement it without a background in MLE or Bayesian statistics (R-PsiMLE). We demonstrate the validity, reliability, efficiency, and flexibility of this method through a series of simulations and behavioral experiments, and find the new method to be superior to the traditional methods in all respects.

Grouping by proximity and the visual impression of approximate number in random dot arrays.

We address the challenges of how to model human perceptual grouping in random dot arrays and how perceptual grouping affects human number estimation in these arrays. We introduce a modeling approach relying on a modified k-means clustering algorithm to formally describe human observers' grouping behavior. We found that a default grouping window size of approximately 4° of visual angle describes human grouping judgments across a range of random dot arrays (i.e., items within 4° are grouped together). This window size was highly consistent across observers and images, and was also stable across stimulus durations, suggesting that the k-means model captured a robust signature of perceptual grouping. Further, the k-means model outperformed other models (e.g., CODE) at describing human grouping behavior. Next, we found that the more the dots in a display are clustered together, the more human observers tend to underestimate the numerosity of the dots. We demonstrate that this effect is independent of density, and the modified k-means model can predict human observers' numerosity judgments and underestimation. Finally, we explored the robustness of the relationship between clustering and dot number underestimation and found that the effects of clustering remain, but are greatly reduced, when participants receive feedback on every trial. Together, this work suggests some promising avenues for formal models of human grouping behavior, and it highlights the importance of a 4° window of perceptual grouping. Lastly, it reveals a robust, somewhat plastic, relationship between perceptual grouping and number estimation.

Encoding attentional states during visuomotor adaptation.

We recently showed that visuomotor adaptation acquired under attentional distraction is better recalled under a similar level of distraction compared to no distraction. This paradoxical effect suggests that attentional state (e.g., divided or undivided) is encoded as an internal context during visuomotor learning and should be reinstated for successful recall (Song & Bédard, 2015). To investigate if there is a critical temporal window for encoding attentional state in visuomotor memory, we manipulated whether participants performed the secondary attention-demanding task concurrently in the early or late phase of visuomotor learning. Recall performance was enhanced when the attentional states between recall and the early phase of visuomotor learning were consistent. However, it reverted to untrained levels when tested under the attentional state of the late-phase learning. This suggests that attentional state is primarily encoded during the early phase of learning before motor errors decrease and reach an asymptote. Furthermore, we demonstrate that when divided and undivided attentional states were mixed during visuomotor adaptation, only divided attention was encoded as an internal cue for memory retrieval. Therefore, a single attentional state appears to be primarily integrated with visuomotor memory while motor error reduction is in progress during learning.

Mean size as a unit of visual working memory.

Visual environments often contain multiple elements, some of which are similar to one another or spatially grouped together. In the current study we investigated how one can use perceptual groups in representing ensemble features of the groups. In experiment 1 we found that participants' performance improved when items were easily segmented by a grouping cue based on proximity, suggesting that spatial grouping facilitates extracting and remembering ensemble representations from visual arrays consisting of multiple elements. In experiment 2 we found that spatial grouping improved performance only when the grouped subsets were tested for the memory task, whereas it impaired performance when other subsets that were not grouped were tested, suggesting that the benefit from grouping may come from better extraction for storage, rather than later decision processes such as accessibility. Taken together, our results suggest that perceptual grouping of multiple items by proximity facilitates extraction of ensemble statistics from groups of items, enhancing visual memory of the ensembles in a visual array.

The effects of sampling and internal noise on the representation of ensemble average size.

Increasing numbers of studies have explored human observers' ability to rapidly extract statistical descriptions from collections of similar items (e.g., the average size and orientation of a group of tilted Gabor patches). Determining whether these descriptions are generated by mechanisms that are independent from object-based sampling procedures requires that we investigate how internal noise, external noise, and sampling affect subjects' performance. Here we systematically manipulated the external variability of ensembles and used variance summation modeling to estimate both the internal noise and the number of samples that affected the representation of ensemble average size. The results suggest that humans sample many more than one or two items from an array when forming an estimate of the average size, and that the internal noise that affects ensemble processing is lower than the noise that affects the processing of single objects. These results are discussed in light of other recent modeling efforts and suggest that ensemble processing of average size relies on a mechanism that is distinct from segmenting individual items. This ensemble process may be more similar to texture processing.