Perceiving human locomotion: priming effects in direction discrimination

During the perception of biological motion, the available stimulus information is confined to a small number of lights attached to the major joints of a moving actor. Despite this drastic impoverishment of the stimulus, the human visual apparatus organizes the swarm of moving dots in a vivid percept of a human figure. In addition, observers effortlessly identify the action the figure is involved in. After a historical introduction and a short walk through the literature, data from a priming experiment are presented. In a serial two-choice reaction-time task, participants were presented with a point-light walker, facing either to the right or to the left and walking either forward or backward on a treadmill. Subjects had to identify the direction of articulatory movements. Reliable priming effects were established in consecutive trials, but these effects were tempered by the relation between priming and primed walker. The reaction time to a walker was shorter when the walker in the preceding trial moved in the same direction and was facing in the same direction. The findings are discussed in relation to recent data from neuropsychological case studies, neuroimaging, and single-cell recording.     

Adaptation to biological motion leads to a motion and a form aftereffect

Recent models have proposed a two-stage process of biological motion recognition. First, template or snapshot neurons estimate the body form. Then, motion is estimated from body form change. This predicts separate aftereffects for body form and body motion. We tested this prediction. Observers viewing leftward- or rightward-facing point-light walkers that walked forward or backward subsequently experienced oppositely directed aftereffects in stimuli ambiguous in the facing or the walking direction. These aftereffects did not originate from adaptation to the motion of the individual light points, because they occurred for limited-lifetime stimuli that restrict local motion. They also occurred when the adaptor displayed a random sequence of body postures that did not induce the walking motion percept. We thus conclude that biological motion gives rise to separate form and motion aftereffects and that body form representations are involved in biological motion perception.     

Motion induction from biological motion

A new type of motion illusion is described in which ambiguous motion becomes unidirectional on superimposition of a human figure walking on a treadmill. A point-light walker in profile was superimposed on a vertical counterphase grating backdrop. Eleven na?ve observers judged the apparent direction of motion against the grating as left or right in a two-alternative forced-choice task and found that the grating appeared to drift in a direction opposite to the walking. The illusion disappeared when the point lights moved in scrambled configurations. This indicates that the illusion is caused by biological motion that provides recognition of gaits. A human figure walking backwards produced no illusion because of the difficulty in identifying the gait. This indicates that the illusion is determined by translational motion rather than form represented from biological motion.     

Learned prediction affects body perception

Learning to recognize objects appears to depend critically on extended observation of appearance over time. Specifically, temporal association between dissimilar views of an object has been proposed as a tool for learning invariant representations for recognition. We examined heretofore untested aspects of the temporal association hypothesis using a familiar dynamic object, the human body. Specifically, we examined the role of appearance prediction (temporal asymmetry) in temporal association. In our task, observers performed a change detection task using upright and inverted images of a walking body either with or without previous exposure to a motion stimulus depicting an upright walker. Observers who were exposed to the dynamic stimulus were further divided into two groups dependent on whether the observed motion depicted forward or backward walking. We find that the effect of the motion stimulus on sensitivity is highly dependent on whether the observed motion is consistent with past experience.     

Biological motion cues trigger reflexive attentional orienting

The human visual system is extremely sensitive to biological signals around us. In the current study, we demonstrate that biological motion walking direction can induce robust reflexive attentional orienting. Following a brief presentation of a central point-light walker walking towards either the left or right direction, observers' performance was significantly better on a target in the walking direction compared with that in the opposite direction even when participants were explicitly told that walking direction was not predictive of target location. Interestingly, the effect disappeared when the walker was shown upside-down. Moreover, the reflexive attentional orienting could be extended to motions of other biological entities but not inanimate objects, and was not due to the viewpoint effect of the point-light figure. Our findings provide strong evidence that biological motion cues can trigger reflexive attentional orienting, and highlight the intrinsic sensitivity of the human visual attention system to biological signals.     

Backward-walking biological motion orients attention to moving away instead of moving toward

Walking direction is an important attribute of biological motion because it carries key information, such as the specific intention of the walker. Although it is known that spatial attention is guided by walking direction, it remains unclear whether this attentional shift is reflexive (i.e., constantly shifts to the walking direction) or not. A richer interpretation of this effect is that attention is guided to seek the information that is necessary to understand the motion. To investigate this issue, we examined how backward-walking biological motion orients attention because the intention of walking backward is usually to avoid something that walking forward would encounter. The results showed that attention was oriented to the walking-away direction of biological motion instead of the walking-toward direction (Experiment 1), and this effect was not due to the gaze direction of biological motion (Experiment 2). Our findings suggest that the attentional shift triggered by walking direction is not reflexive, thus providing support for the rich interpretation of these attentional effects.     

Bistability and biasing effects in the perception of ambiguous point-light walkers

The perceptually bistable character of point-light walkers has been examined in three experiments. A point-light figure without explicit depth cues constitutes a perfectly ambiguous stimulus: from all viewpoints, multiple interpretations are possible concerning the depth orientation of the figure. In the first experiment, it is shown that non-lateral views of the walker are indeed interpreted in two orientations, either as facing towards the viewer or as facing away from the viewer, but that the interpretation in which the walker is oriented towards the viewer is reported more frequently. In the second experiment the point-light figure was walking backwards, making the global orientation of the point-light figure opposite to the direction of global motion. The interpretation in which the walker was facing the viewer was again reported more frequently. The robustness of these findings was examined in the final experiment, in which the effects of disambiguating the stimulus by introducing a local depth cue (occlusion) or a more global depth cue (applying perspective projection) were explored.     

Intact perception of biological motion in the face of profound spatial deficits: Williams syndrome

Williams syndrome (WS) is a rare genetic disorder that results in profound spatial cognitive deficits. We examined whether individuals with WS have intact perception of biological motion, which requires global spatial integration of local motion signals into a unitary percept of a human form. Children with WS, normal mental-age-matched children, and normal adults viewed point-light-walker (PLW) displays portraying a human figure walking to the left or right. Children with WS were as good as or better than control children in their ability to judge the walker's direction, even when it was masked with dynamic noise that mimicked the local motion of the PLW lights. These results show that mechanisms underlying the perception of at least some kinds of biological motion are unimpaired in children with WS. They provide the first evidence of selective sparing of a specialized spatial system in individuals with a known genetic impairment.     

Body configuration modulates the usage of local cues to direction in biological-motion perception

The presence of information in a visual display does not guarantee its use by the visual system. Studies of inversion effects in both face recognition and biological-motion perception have shown that the same information may be used by observers when it is presented in an upright display but not used when the display is inverted. In our study, we tested the inversion effect in scrambled biological-motion displays to investigate mechanisms that validate information contained in the local motion of a point-light walker. Using novel biological-motion stimuli that contained no configural cues to the direction in which a walker was facing, we found that manipulating the relative vertical location of the walker's feet significantly affected observers' performance on a direction-discrimination task. Our data demonstrate that, by themselves, local cues can almost unambiguously indicate the facing direction of the agent in biological-motion stimuli. Additionally, we document a noteworthy interaction between local and global information and offer a new explanation for the effect of local inversion in biological-motion perception.     

Temporal properties in masking biological motion

The perception of biological motion using point light animation techniques was investigated in several experiments. Animations simulating walking were presented with additional masking dots. The temporal properties of the walking motion or the temporal relationship between the walking and masking motions were systematically manipulated. Results showed that (1) perception of biological motion was sensitive to even small temporal perturbation within the walker, (2) the effectiveness of a mask depended upon the temporal phase difference between the mask and point light walker, (3) relatively small temporal differences between the mask and point light walker decreased the effectiveness of the mask, and (4) these effects were not due simply to observers detecting the phase offsets in the display. Temporal properties of the motion are important in perceiving the human form in action, just as in other types of figure-ground segregation. This information may be processed by both motion and form pathways for processing biological motion.     

The informational properties of the throwing arm for anticipation of goal-directed action

We examined the informational value of biological motion from the arm in predicting the location of a thrown ball. In three experiments, participants were classified as being skilled and less skilled based on their actual performance on the task (i.e., using a within-task criterion). We then presented participants with a range of stick figure representations and required them to predict throw direction. In Experiment 1, we presented stick figure movies of a full body throwing action, right throwing arm plus left shoulder and throwing arm only. Participants were able to anticipate throw direction above chance under all conditions irrespective of perceptual skill level, with the perceptually skilled participants excelling under full body conditions. In Experiment 2, we neutralized dynamical differences in motion to opposing throw directions from the shoulder, elbow and wrist of the throwing arm. Neutralizing the wrist location negatively affected anticipation performance in all participants reducing accuracy to below chance. In Experiment 3, we presented movies of the motion wrist location alone and the upper section of the throwing arm (shoulder-elbow). Participants were able to successfully anticipate above chance in these latter two conditions. Our findings suggest that motion of the throwing arm contains multiple sources of information that can help facilitate the anticipation of goal-directed action. Perceptually skilled participants were superior in extracting informational value from motion at both the local and global levels when compared to less skilled counterparts.     

Impaired visual perception of hurtful actions in patients with chronic low back pain

Visually presented biological motion stimuli activate regions in the brain that are also related to musculo-skeletal pain. We therefore hypothesized that chronic pain impairs the perception of visually presented actions that involve body parts that hurt. In the first experiment, chronic back pain (CLBP) patients and healthy controls judged the lifted weight from point-light biological motion displays. An actor either lifted an invisible container (5, 10, or 15 kg) from the floor, or lifted and manipulated it from the right to the left. The latter involved twisting of the lower back and would be very painful for CLBP patients. All participants recognized the displayed actions, but CLBP patients were impaired in judging the difference in handled weights, especially for the trunk rotation. The second experiment involved discrimination between forward and backward walking. Here the patients were just as good as the controls, showing that the main result of the first experiment was indeed specific to the sensory aspects of the task, and not to general impairments or attentional deficits. The results thus indicate that the judgment of sensorimotor aspects of a visually displayed movement is specifically affected by chronic low back pain.     

Reproducibility of distance and direction errors associated with forward, backward, and sideway walking in the context of blind navigation

The ability to navigate without vision towards a previously seen target has been extensively studied, but its reliability over time has yet to be established. Our aims were to determine distance and direction errors made during blind navigation across four different directions involving three different gait patterns (stepping forward, stepping sideway, and stepping backward), and to establish the test-retest reproducibility of these errors. Twenty young healthy adults participated in two testing sessions separated by 7 days. They were shown targets located, respectively, 8 m ahead, 8 m behind, and 8 m to their right and left. With vision occluded by opaque goggles, they walked forward (target ahead), backward (target behind), and sideway (right and left targets) until they perceived to be on the target. Subjects were not provided with feedback about their performance. Walked distance, angular deviation, and body rotation were measured. The mean estimated distance error was similar across the four walking directions and ranged from 16 to 80 cm with respect to the 8 m target. In contrast, direction errors were significantly larger during sideway navigation (walking in the frontal plane: leftward, 10 degrees +/- 15 degrees deviation; rightward, 18 degrees +/- 13 degrees) than during forward and backward navigation (walking in the sagittal plane). In general, distance and direction errors were only moderately reproducible between the two sessions [intraclass correlation coefficients (ICCs) ranging from 0.682 to 0.705]. Among the four directions, rightward navigation showed the best reproducibility with ICCs ranging from 0.607 to 0.726, and backward navigation had the worst reliability with ICCs ranging from 0.094 to 0.554. These findings indicate that errors associated with blind navigation across different walking directions and involving different gait patterns are only moderately to poorly reproducible on repeated testing, especially for walking backward. The biomechanical constraints and increased cognitive loading imposed by changing the walking pattern to backward stepping may underlie the poor performance in this direction.     

The perception of biological motion across apertures

To understand the visual analysis of biological motion, subjects viewed dynamic, stick figure renditions of a walker, car, or scissors through apertures. As a result of the aperture problem, the motion of each visible edge was ambiguous. Subjects readily identified the human figure but were unable to identify the car or scissors through invisible apertures. Recognition was orientation specific and robust across a range of stimulus durations, and it benefited from limb orientation cues. The results support the theory that the visual system performs spatially global analyses to interpret biological motion displays.     

Perception of biological motion from limited-lifetime stimuli

The visual perception of human movement from sparse point-light walkers is often believed to rely on local motion analysis. We investigated the role of local motion in the perception of human walking, viewed from the side, in different tasks. The motion signal was manipulated by varying point lifetime. We found the task of coherence discrimination, commonly used in biological motion studies, to be inappropriate for testing the role of motion. A task requiring temporal information showed a strong performance drop when fewer points were used or when the image sequence was sampled and displayed at a reduced frame rate. Irrespective of the frame rate, performance did not vary with point lifetime. We concluded that local motion is not required for the perception of tested biological movements, suggesting that the analysis of biological motion does not benefit from examining local motion. The reliance of perception on the number of displayed points and frames supports the idea that biological motion is perceived from a sequence of spatiotemporally sampled forms.     

Local Dynamic Stability of the Locomotion of Lower Extremity Joints and Trunk During Backward Upslope Walking

Backward slope walking was considered as a practical rehabilitation and training skill. However, its gait stability has been hardly studied, resulting in its limited application as a rehabilitation tool. In this study, the effect of walking direction and slope grade were investigated on the local dynamic stability of the motion of lower extremity joints and trunk segment during backward and forward upslope walking (BUW/FUW). The local divergence exponents (λ_S) of 16 adults were calculated during their BUW and FUW at grades of 0%, 5%, 10%, and 15%. Mean standard deviation over strides (MeanSD) was analyzed as their gait variability. Backward walking showed larger λ_S for the abduction-adduction and rotational angles of knee and ankle on inclined surface than forward walking, while λ_S for hip flexion-extension angle at steeper grades was opposite. No grade effect for any joint existed during BUW, while λ_S increased with the increasing grade during FUW. As to the trunk, walking direction did little impact on λ_S. Still, significant larger λ_S for its medial-lateral and vertical motion were found at the steeper grades during both FUW and BUW. Results indicate that during BUW, the backward direction may influence the stability of joint motions, while the trunk stability was challenged by the increasing grades. Therefore, BUW may be a training tool for the stability of both upper and lower body motion during gait.     

A Simon effect with stationary moving stimuli

To clarify whether motion information per se has a separable influence on action control, the authors investigated whether irrelevant direction of motion of stimuli whose overall position was constant over time would affect manual left-right responses (i.e., reveal a motion-based Simon effect). In Experiments 1 and 2, significant Simon effects were obtained for sine-wave gratings moving in a stationary Gaussian window. In Experiment 3, a direction-based Simon effect with random-dot patterns was replicated, except that the perceived direction of motion was based on the displacement of single elements. Experiments 4 and 5 studied motion-based Simon effects to point-light figures that walked in place--displays requiring high-level analysis of global shape and local motion. Motion-based Simon effects occurred when the displays could be interpreted as an upright human walker, showing that a high-level representation of motion direction mediated the effects. Thus, the present study establishes links between high-level motion perception and action.     

Dual-task costs of texting while walking forward and backward are greater for older adults than younger adults

Cognitive-motor dual-tasking involves concurrent performance of two tasks with distinct cognitive and motor demands and is associated with increased fall risk. In this hypothesis-driven study, younger (18–30 years, n = 24) and older (60–75 years, n = 26) adults completed six walking tasks in triplicate. Participants walked forward and backward along a GAITRite mat, in isolation or while performing a verbal fluency task. Verbal fluency tasks involved verbally listing or typing on a smartphone as many words as possible within a given category (e.g., clothes). Using repeated measures MANOVA models, we examined how age, method of fluency task (verbal or texting), and direction of walking altered dual-task performance. Given that tasks like texting and backward walking require greater cognitive resources than verbal and forward walking tasks, respectively, we hypothesized older adults would show higher dual-task costs (DTCs) than younger adults across different task types and walking directions, with degree of impairment more apparent in texting dual-task trials compared to verbal dual-task trials. We also hypothesized that both age groups would have greater DTCs while walking backward than while walking forward, regardless of task.Independent of age group, velocity and stride length were reduced for texting compared to the verbal task during both forward and backward walking; cadence and velocity were reduced while walking forward compared to walking backward for the texting task; and stride length was reduced for forward walking compared to backward walking during the verbal task. Younger adults performed better than older adults on all tasks with the most pronounced differences seen in velocity and stride length during forward-texting and backward-texting. Interaction effects for velocity and stride length while walking forward indicated younger adults performed better than older adults for the texting task but similarly during the verbal task. An interaction for cadence during the verbal task indicated younger adults performed better than older adults while walking backward but similarly while walking forward.In summary, older adults experienced greater gait decrement for all dual-task conditions. The greater declines in velocity and stride length in combination with cadence being stable suggest reductions in velocity during texting were due to shorter strides rather than a reduced rate of stepping. Contrary to our hypotheses, we found greater DTCs while walking forward rather than backward, which may be due to reduced gait performance during single-task backward walking; thus, further decrements with dual-tasking are unlikely. These findings underscore the need for further research investigating fall risk potential associated with texting and walking among aging populations and how interventions targeting stride length during dual-task circumstances may improve performance.     

Contribution of ankle, knee, and hip joints to the perception threshold for support surface rotation.

The purpose of the present experiment was to investigate the extent to which subjects can perceive, at very slow velocities, an angular rotation of the support surface about the medio-lateral axis of the ankle, knee, hip, or neck joint when visual cues are not available. Subjects were passively displaced on a slowly rotating platform at .01, .03, and .05 deg/sec. The subjects' task was to detect movements of the platform in four different postural conditions allowing body oscillations about the ankle, knee, hip, or neck joint. In Experiment 1, subjects had to detect backward and forward rotation (pitching). In Experiment 2, they had to detect left and right rotations of the platform (rolling). In Experiment 3, subjects had to detect both backward/forward and left/right rotations of the platform, with the body fixed and the head either fixed or free to move. Overall, when the body was free to oscillate about the ankle, knee, or hip joints, a similar threshold for movement perception was observed. This threshold was lower for rolling than for pitching. Interestingly, in these postural conditions, an unconscious compensation in the direction opposite to the platform rotation was observed on most trials. The threshold for movement perception was much higher when the head was the only segment free to oscillate about the neck joint. These results suggest that, in static conditions, the otoliths are poor detectors of the direction of gravity forces. They also suggest that accurate perception of body orientation is improved when proprioceptive information can be dynamically integrated.     

Contribution of ankle, knee, and hip joints to the perception threshold for support surface rotation

The purpose of the present experiment was to investigate the extent to which subjects can perceive, at very slow velocities, an angular rotation of the support surface about the medio-lateral axis of the ankle, knee, hip, or neck joint when visual cues are not available. Subjects were passively displaced on a slowly rotating platform at .01, .03, and .05 deg/sec. The subjects’ task was to detect movements of the platform in four different postural conditions allowing body oscillations about the ankle, knee, hip, or neck joint. In Experiment 1, subjects had to detect backward and forward rotation (pitching). In Experiment 2, they had to detect left and right rotations of the platform (rolling). In Experiment 3, subjects had to detect both backward/forward and left/right rotations of the platform, with the body fixed and the head either fixed or free to move. Overall, when the body was free to oscillate about the ankle, knee, or hip joints, a similar threshold for movement perception was observed. This threshold was lower for rolling than for pitching. Interestingly, in these postural conditions, an unconscious compensation in the direction opposite to the platform rotation was observed on most trials. The threshold for movement perception was much higher when the head was the only segment free to oscillate about the neck joint. These results suggest that, in static conditions, the otoliths are poor detectors of the direction of gravity forces. They also suggest that accurate perception of body orientation is improved when proprioceptive information can be dynamically integrated.