The Inversion Effect in Biological Motion Perception: Evidence for a “Life Detector”?

If biological-motion point-light displays are presented upside down, adequate perception is strongly impaired. Reminiscent of the inversion effect in face recognition, it has been suggested that the inversion effect in biological motion is due to impaired configural processing in a highly trained expert system. Here, we present data that are incompatible with this view. We show that observers can readily retrieve information about direction from scrambled point-light displays of humans and animals. Even though all configural information is entirely disrupted, perception of these displays is still subject to a significant inversion effect. Inverting only parts of the display reveals that the information about direction, as well as the associated inversion effect, is entirely carried by the local motion of the feet. We interpret our findings in terms of a visual filter that is tuned to the characteristic motion of the limbs of an animal in locomotion and hypothesize that this mechanism serves as a general detection system for the presence of articulated terrestrial animals.     

Perception of Social Interactions for Spatially Scrambled Biological Motion

It is vitally important for humans to detect living creatures in the environment and to analyze their behavior to facilitate action understanding and high-level social inference. The current study employed naturalistic point-light animations to examine the ability of human observers to spontaneously identify and discriminate socially interactive behaviors between two human agents. Specifically, we investigated the importance of global body form, intrinsic joint movements, extrinsic whole-body movements, and critically, the congruency between intrinsic and extrinsic motions. Motion congruency is hypothesized to be particularly important because of the constraint it imposes on naturalistic action due to the inherent causal relationship between limb movements and whole body motion. Using a free response paradigm in Experiment 1, we discovered that many naïve observers (55%) spontaneously attributed animate and/or social traits to spatially-scrambled displays of interpersonal interaction. Total stimulus motion energy was strongly correlated with the likelihood that an observer would attribute animate/social traits, as opposed to physical/mechanical traits, to the scrambled dot stimuli. In Experiment 2, we found that participants could identify interactions between spatially-scrambled displays of human dance as long as congruency was maintained between intrinsic/extrinsic movements. Violating the motion congruency constraint resulted in chance discrimination performance for the spatially-scrambled displays. Finally, Experiment 3 showed that scrambled point-light dancing animations violating this constraint were also rated as significantly less interactive than animations with congruent intrinsic/extrinsic motion. These results demonstrate the importance of intrinsic/extrinsic motion congruency for biological motion analysis, and support a theoretical framework in which early visual filters help to detect animate agents in the environment based on several fundamental constraints. Only after satisfying these basic constraints could stimuli be evaluated for high-level social content. In this way, we posit that perceptual animacy may serve as a gateway to higher-level processes that support action understanding and social inference.     

Integrating biological motion: the role of grouping in the perception of point-light actions

The human visual system is highly sensitive to biological motion and manages to organize even a highly reduced point-light stimulus into a vivid percept of human action. The current study investigated to what extent the origin of this saliency of point-light displays is related to its intrinsic Gestalt qualities. In particular, we studied whether biological motion perception is facilitated when the elements can be grouped according to good continuation and similarity as Gestalt principles of perceptual organization. We found that both grouping principles enhanced biological motion perception but their effects differed when stimuli were inverted. These results provide evidence that Gestalt principles of good continuity and similarity also apply to more complex and dynamic meaningful stimuli.     

Biological Motion Primes the Animate/Inanimate Distinction in Infancy

Given that biological motion is both detected and preferred early in life, we tested the hypothesis that biological motion might be instrumental to infants’ differentiation of animate and inanimate categories. Infants were primed with either point-light displays of realistic biological motion, random motion, or schematic biological motion of an unfamiliar shape. After being habituated to these displays, 12-month-old infants categorized animals and vehicles as well as furniture and vehicles with the sequential touching task. The findings indicated that infants primed with point-light displays of realistic biological motion showed better categorization of animates than those exposed to random or schematic biological motion. These results suggest that human biological motion might be one of the motion cues that provide the building blocks for infants’ concept of animacy.     

Patterns of fMRI activity dissociate overlapping functional brain areas that respond to biological motion

Accurate perception of the actions and intentions of other people is essential for successful interactions in a social environment. Several cortical areas that support this process respond selectively in fMRI to static and dynamic displays of human bodies and faces. Here we apply pattern-analysis techniques to arrive at a new understanding of the neural response to biological motion. Functionally defined body-, face-, and motion-selective visual areas all responded significantly to "point-light" human motion. Strikingly, however, only body selectivity was correlated, on a voxel-by-voxel basis, with biological motion selectivity. We conclude that (1) biological motion, through the process of structure-from-motion, engages areas involved in the analysis of the static human form; (2) body-selective regions in posterior fusiform gyrus and posterior inferior temporal sulcus overlap with, but are distinct from, face- and motion-selective regions; (3) the interpretation of region-of-interest findings may be substantially altered when multiple patterns of selectivity are considered.     

Investigating the status of biological stimuli as objects of attention in multiple object tracking

Background

Humans are able to track multiple simultaneously moving objects. A number of factors have been identified that can influence the ease with which objects can be attended and tracked. Here, we explored the possibility that object tracking abilities may be specialized for tracking biological targets such as people.

Methodology/Principal Findings

We used the Multiple Object Tracking (MOT) paradigm to explore whether the high-level biological status of the targets affects the efficiency of attentional selection and tracking. In Experiment 1, we assessed the tracking of point-light biological motion figures. As controls, we used either the same stimuli or point-light letters, presented in upright, inverted or scrambled configurations. While scrambling significantly affected performance for both letters and point-light figures, there was an effect of inversion restricted to biological motion, inverted figures being harder to track. In Experiment 2, we found that tracking performance was equivalent for natural point-light walkers and ‘moon-walkers’, whose implied direction was incongruent with their actual direction of motion. In Experiment 3, we found higher tracking accuracy for inverted faces compared with upright faces. Thus, there was a double dissociation between inversion effects for biological motion and faces, with no inversion effect for our non-biological stimuli (letters, houses).

Conclusions/Significance

MOT is sensitive to some, but not all naturalistic aspects of biological stimuli. There does not appear to be a highly specialized role for tracking people. However, MOT appears constrained by principles of object segmentation and grouping, where effectively grouped, coherent objects, but not necessarily biological objects, are tracked most successfully.     

Perception of biological motion in schizophrenia and healthy individuals: a behavioral and FMRI study

Background

Anomalous visual perception is a common feature of schizophrenia plausibly associated with impaired social cognition that, in turn, could affect social behavior. Past research suggests impairment in biological motion perception in schizophrenia. Behavioral and functional magnetic resonance imaging (fMRI) experiments were conducted to verify the existence of this impairment, to clarify its perceptual basis, and to identify accompanying neural concomitants of those deficits.

Methodology/Findings

In Experiment 1, we measured ability to detect biological motion portrayed by point-light animations embedded within masking noise. Experiment 2 measured discrimination accuracy for pairs of point-light biological motion sequences differing in the degree of perturbation of the kinematics portrayed in those sequences. Experiment 3 measured BOLD signals using event-related fMRI during a biological motion categorization task.Compared to healthy individuals, schizophrenia patients performed significantly worse on both the detection (Experiment 1) and discrimination (Experiment 2) tasks. Consistent with the behavioral results, the fMRI study revealed that healthy individuals exhibited strong activation to biological motion, but not to scrambled motion in the posterior portion of the superior temporal sulcus (STSp). Interestingly, strong STSp activation was also observed for scrambled or partially scrambled motion when the healthy participants perceived it as normal biological motion. On the other hand, STSp activation in schizophrenia patients was not selective to biological or scrambled motion.

Conclusion

Schizophrenia is accompanied by difficulties discriminating biological from non-biological motion, and associated with those difficulties are altered patterns of neural responses within brain area STSp. The perceptual deficits exhibited by schizophrenia patients may be an exaggerated manifestation of neural events within STSp associated with perceptual errors made by healthy observers on these same tasks. The present findings fit within the context of theories of delusion involving perceptual and cognitive processes.     

The N170 component is sensitive to face-like stimuli: a study of Chinese Peking opera makeup

The N170 component is considered a neural marker of face-sensitive processing. In the present study, the face-sensitive N170 component of event-related potentials (ERPs) was investigated with a modified oddball paradigm using a natural face (the standard stimulus), human- and animal-like makeup stimuli, scrambled control images that mixed human- and animal-like makeup pieces, and a grey control image. Nineteen participants were instructed to respond within 1000 ms by pressing the ‘F’ or ‘J’ key in response to the standard or deviant stimuli, respectively. We simultaneously recorded ERPs, response accuracy, and reaction times. The behavioral results showed that the main effect of stimulus type was significant for reaction time, whereas there were no significant differences in response accuracies among stimulus types. In relation to the ERPs, N170 amplitudes elicited by human-like makeup stimuli, animal-like makeup stimuli, scrambled control images, and a grey control image progressively decreased. A right hemisphere advantage was observed in the N170 amplitudes for human-like makeup stimuli, animal-like makeup stimuli, and scrambled control images but not for grey control image. These results indicate that the N170 component is sensitive to face-like stimuli and reflect configural processing in face recognition.     

Perceptual grouping of biological motion promotes binocular rivalry

Investigation of perceptual rivalry between conflicting stimuli presented one to each eye can further understanding of the neural underpinnings of conscious visual perception. During rivalry, visual awareness fluctuates between perceptions of the two stimuli. Here, we demonstrate that high-level perceptual grouping can promote rivalry between stimulus pairs that would otherwise be perceived as nonrivalrous. Perceptual grouping was generated with point-light walker stimuli that simulate human motion, visible only as lights placed on the joints. Although such walking figures are unrecognizable when stationary, recognition judgments as complex as gender and identity can accurately be made from animated displays, demonstrating the efficiency with which our visual system can group dynamic local signals into a globally coherent walking figure. We find that point-light walker stimuli presented one to each eye and in different colors and configurations results in strong rivalry. However, rivalry is minimal when the two walkers are split between the eyes or both presented to one eye. This pattern of results suggests that processing animated walker figures promotes rivalry between signals from the two eyes rather than between higher-level representations of the walkers. This leads us to hypothesize that awareness during binocular rivalry involves the integrated activity of high-level perceptual mechanisms in conjunction with lower-level ocular suppression modulated via cortical feedback.     

Biological Motion Coding in the Brain: Analysis of Visually Driven EEG Functional Networks

Herein, we address the time evolution of brain functional networks computed from electroencephalographic activity driven by visual stimuli. We describe how these functional network signatures change in fast scale when confronted with point-light display stimuli depicting biological motion (BM) as opposed to scrambled motion (SM). Whereas global network measures (average path length, average clustering coefficient, and average betweenness) computed as a function of time did not discriminate between BM and SM, local node properties did. Comparing the network local measures of the BM condition with those of the SM condition, we found higher degree and betweenness values in the left frontal (F7) electrode, as well as a higher clustering coefficient in the right occipital (O2) electrode, for the SM condition. Conversely, for the BM condition, we found higher degree values in central parietal (Pz) electrode and a higher clustering coefficient in the left parietal (P3) electrode. These results are discussed in the context of the brain networks involved in encoding BM versus SM.     

Do dogs mind the dots? Investigating domestic dogs' (Canis familiaris) preferential looking at human-shaped point-light figures

It is generally assumed that dogs show increased attention towards humans. A major part of this includes attention towards visual cues such as bodily gestures. We tested empirically whether dogs are visually attentive towards human body movement. Based on methods from visual perception research in humans, we used point-light figures (PLFs) to investigate whether dogs attend to human body movement compared to other forms of motion. We investigated dogs' attentiveness towards vocalisation-paired PLFs by adopting a preferential looking paradigm. Results indicate that dogs show increased attention towards vocalisation-paired human-shaped PLFs in comparison with inverted and non-inverted scrambled configurations of the same PLFs. This increased attention, however, was only present in the case of PLFs that simulated a human in frontal orientation but not for PLFs in lateral orientation. We conclude that dogs prefer to look at PLFs of socially relevant (i.e. frontally facing) human body (i.e. natural) movement rather than scrambled (i.e. unnatural) displays. Our results indicate that PLFs may function as a promising tool to investigate dogs' visual perceptual preferences and mechanisms.     

Pre-Exposure to Moving Form Enhances Static Form Sensitivity

Background

Motion-defined form can seem to persist briefly after motion ceases, before seeming to gradually disappear into the background. Here we investigate if this subjective persistence reflects a signal capable of improving objective measures of sensitivity to static form.

Methodology/Principal Findings

We presented a sinusoidal modulation of luminance, masked by a background noise pattern. The sinusoidal luminance modulation was usually subjectively invisible when static, but visible when moving. We found that drifting then stopping the waveform resulted in a transient subjective persistence of the waveform in the static display. Observers'' objective sensitivity to the position of the static waveform was also improved after viewing moving waveforms, compared to viewing static waveforms for a matched duration. This facilitation did not occur simply because movement provided more perspectives of the waveform, since performance following pre-exposure to scrambled animations did not match that following pre-exposure to smooth motion. Observers did not simply remember waveform positions at motion offset, since removing the waveform before testing reduced performance.

Conclusions/Significance

Motion processing therefore interacts with subsequent static visual inputs in a way that can improve performance in objective sensitivity measures. We suggest that the brief subjective persistence of motion-defined forms that can occur after motion offsets is a consequence of the decay of a static form signal that has been transiently enhanced by motion processing.     

Impaired Recognition of Communicative Interactions from Biological Motion in Schizophrenia

BackgroundPatients with schizophrenia are deficient in multiple aspects of social cognition, including biological motion perception. In the present study we investigated the ability to read social information from point-light stimuli in schizophrenia.Conclusions/SignificanceThese findings are consistent with theories of “overmentalizing” (excessive attribution of intentionality) in schizophrenia, and suggest that processing social information from biological motion does rely on social cognition abilities.     

Rapid Perceptual Switching of a Reversible Biological Figure

Certain visual stimuli can give rise to contradictory perceptions. In this paper we examine the temporal dynamics of perceptual reversals experienced with biological motion, comparing these dynamics to those observed with other ambiguous structure from motion (SFM) stimuli. In our first experiment, naïve observers monitored perceptual alternations with an ambiguous rotating walker, a figure that randomly alternates between walking in clockwise (CW) and counter-clockwise (CCW) directions. While the number of reported reversals varied between observers, the observed dynamics (distribution of dominance durations, CW/CCW proportions) were comparable to those experienced with an ambiguous kinetic depth cylinder. In a second experiment, we compared reversal profiles with rotating and standard point-light walkers (i.e. non-rotating). Over multiple test repetitions, three out of four observers experienced consistently shorter mean percept durations with the rotating walker, suggesting that the added rotational component may speed up reversal rates with biomotion. For both stimuli, the drift in alternation rate across trial and across repetition was minimal. In our final experiment, we investigated whether reversals with the rotating walker and a non-biological object with similar global dimensions (rotating cuboid) occur at random phases of the rotation cycle. We found evidence that some observers experience peaks in the distribution of response locations that are relatively stable across sessions. Using control data, we discuss the role of eye movements in the development of these reversal patterns, and the related role of exogenous stimulus characteristics. In summary, we have demonstrated that the temporal dynamics of reversal with biological motion are similar to other forms of ambiguous SFM. We conclude that perceptual switching with biological motion is a robust bistable phenomenon.     

Unaffected perceptual thresholds for biological and non-biological form-from-motion perception in autism spectrum conditions

Background

Perception of biological motion is linked to the action perception system in the human brain, abnormalities within which have been suggested to underlie impairments in social domains observed in autism spectrum conditions (ASC). However, the literature on biological motion perception in ASC is heterogeneous and it is unclear whether deficits are specific to biological motion, or might generalize to form-from-motion perception.

Methodology and Principal Findings

We compared psychophysical thresholds for both biological and non-biological form-from-motion perception in adults with ASC and controls. Participants viewed point-light displays depicting a walking person (Biological Motion), a translating rectangle (Structured Object) or a translating unfamiliar shape (Unstructured Object). The figures were embedded in noise dots that moved similarly and the task was to determine direction of movement. The number of noise dots varied on each trial and perceptual thresholds were estimated adaptively. We found no evidence for an impairment in biological or non-biological object motion perception in individuals with ASC. Perceptual thresholds in the three conditions were almost identical between the ASC and control groups.

Discussion and Conclusions

Impairments in biological motion and non-biological form-from-motion perception are not across the board in ASC, and are only found for some stimuli and tasks. We discuss our results in relation to other findings in the literature, the heterogeneity of which likely relates to the different tasks performed. It appears that individuals with ASC are unaffected in perceptual processing of form-from-motion, but may exhibit impairments in higher order judgments such as emotion processing. It is important to identify more specifically which processes of motion perception are impacted in ASC before a link can be made between perceptual deficits and the higher-level features of the disorder.     

Direction of Biological Motion Affects Early Brain Activation: A Link with Social Cognition

A number of EEG studies have investigated the time course of brain activation for biological movement over this last decade, however the temporal dynamics of processing are still debated. Moreover, the role of direction of movement has not received much attention even though it is an essential component allowing us to determine the intentions of the moving agent, and thus permitting the anticipation of potential social interactions.In this study, we examined event-related responses (ERPs) in 15 healthy human participants to light point walkers and their scrambled counterparts, whose movements occurred either in the radial or in the lateral plane. Compared to scrambled motion (SM), biological motion (BM) showed an enhanced negativity between 210 and 360ms. A source localization algorithm (sLORETA) revealed that this was due to an increase in superior and middle temporal lobe activity. Regarding direction, we found that radial BM produced an enhanced P1 compared to lateral BM, lateral SM and radial SM. This heightened P1 was due to an increase in activity in extrastriate regions, as well as in superior temporal, medial parietal and medial prefrontal areas. This network is known to be involved in decoding the underlying intentionality of the movement and in the attribution of mental states. The social meaning signaled by the direction of biological motion therefore appears to trigger an early response in brain activity.     

Further Explorations of the Facing Bias in Biological Motion Perception: Perspective Cues,Observer Sex,and Response Times

The human visual system has evolved to be highly sensitive to visual information about other persons and their movements as is illustrated by the effortless perception of point-light figures or ‘biological motion’. When presented orthographically, a point-light walker is interpreted in two anatomically plausible ways: As ‘facing the viewer’ or as ‘facing away’ from the viewer. However, human observers show a ‘facing bias’: They perceive such a point-light walker as facing towards them in about 70-80% of the cases. In studies exploring the role of social and biological relevance as a possible account for the facing bias, we found a ‘figure gender effect’: Male point-light figures elicit a stronger facing bias than female point-light figures. Moreover, we also found an ‘observer gender effect’: The ‘figure gender effect’ was stronger for male than for female observers. In the present study we presented to 11 males and 11 females point-light walkers of which, very subtly, the perspective information was manipulated by modifying the earlier reported ‘perspective technique’. Proportions of ‘facing the viewer’ responses and reaction times were recorded. Results show that human observers, even in the absence of local shape or size cues, easily pick up on perspective cues, confirming recent demonstrations of high visual sensitivity to cues on whether another person is potentially approaching. We also found a consistent difference in how male and female observers respond to stimulus variations (figure gender or perspective cues) that cause variations in the perceived in-depth orientation of a point-light walker. Thus, the ‘figure gender effect’ is possibly caused by changes in the relative locations and motions of the dots that the perceptual system tends to interpret as perspective cues. Third, reaction time measures confirmed the existence of the facing bias and recent research showing faster detection of approaching than receding biological motion.     

Perception of ‘Back-Channeling’ Nonverbal Feedback in Musical Duo Improvisation

In witnessing face-to-face conversation, observers perceive authentic communication according to the social contingency of nonverbal feedback cues (‘back-channeling’) by non-speaking interactors. The current study investigated the generality of this function by focusing on nonverbal communication in musical improvisation. A perceptual experiment was conducted to test whether observers can reliably identify genuine versus fake (mismatched) duos from musicians’ nonverbal cues, and how this judgement is affected by observers’ musical background and rhythm perception skill. Twenty-four musicians were recruited to perform duo improvisations, which included solo episodes, in two styles: standard jazz (where rhythm is based on a regular pulse) or free improvisation (where rhythm is non-pulsed). The improvisations were recorded using a motion capture system to generate 16 ten-second point-light displays (with audio) of the soloist and the silent non-soloing musician (‘back-channeler’). Sixteen further displays were created by splicing soloists with back-channelers from different duos. Participants (N = 60) with various musical backgrounds were asked to rate the point-light displays as either real or fake. Results indicated that participants were sensitive to the real/fake distinction in the free improvisation condition independently of musical experience. Individual differences in rhythm perception skill did not account for performance in the free condition, but were positively correlated with accuracy in the standard jazz condition. These findings suggest that the perception of back-channeling in free improvisation is not dependent on music-specific skills but is a general ability. The findings invite further study of the links between interpersonal dynamics in conversation and musical interaction.     

Self-recognition of avatar motion: how do I know it's me?

When motion is isolated from form cues and viewed from third-person perspectives, individuals are able to recognize their own whole body movements better than those of friends. Because we rarely see our own bodies in motion from third-person viewpoints, this self-recognition advantage may indicate a contribution to perception from the motor system. Our first experiment provides evidence that recognition of self-produced and friends' motion dissociate, with only the latter showing sensitivity to orientation. Through the use of selectively disrupted avatar motion, our second experiment shows that self-recognition of facial motion is mediated by knowledge of the local temporal characteristics of one's own actions. Specifically, inverted self-recognition was unaffected by disruption of feature configurations and trajectories, but eliminated by temporal distortion. While actors lack third-person visual experience of their actions, they have a lifetime of proprioceptive, somatosensory, vestibular and first-person-visual experience. These sources of contingent feedback may provide actors with knowledge about the temporal properties of their actions, potentially supporting recognition of characteristic rhythmic variation when viewing self-produced motion. In contrast, the ability to recognize the motion signatures of familiar others may be dependent on configural topographic cues.