Statistical and information properties of head direction cells

The human channel capacity for identifying sensory stimuli is compared with channel capacities based on neurophysiological findings. Studies have shown that cells in the postsubiculum (PoS) and the anterior dorsal thalamus (ADN) of the rat discharge as a function of the animal’s head direction in the horizontal plane. We compute the statistical properties of the firing rates of head direction (HD) cells and the potential amount of information transmitted by these cells according to two theoretical models. Thecell response model for single cells indicates that information transmitted is much less than 0.5 bits. Thepopulation response model developed for cell ensembles generates values in the range of 1–3.2 bits, suggesting that a cell population can distinguish between two and nine head directions, depending on the value used for the standard deviation of directions over which a cell fires. These values are similar to those found in human psychophysical studies of the channel capacity for unidimensional sensory attributes.     

Illusory visual direction during and after backward head tilts

When the head is returned to upright after prolonged backward tilt, people who are asked to look straight ahead look higher than they did before the backward tilting. One explanation attributes the illusory visual direction to a change in muscle responsiveness which develops after the head is returned to upright. According to this explanation, the illusory shift should be absent, or at least reduced, if the subjects are not returned to upright before testing. In the present study, the illusion was the same whether subjects remained tilted for testing or were returned to upright.     

Central visual learning and illusory visual direction after backward head tilts

When the head is returned to upright after prolonged backward tilt, people who are asked to look straight ahead look higher than they did before the backward tilting. This has been interpreted in terms of hypotheses about central visual learning or by hypotheses about peripheral muscle physiology. According to the learning hypotheses, the illusion of visual direction that occurs after head tilts depends upon the presence of discordant cues about direction. In the present study, the illusion was the same with or without discordant information.     

Discriminating direction of motion trajectories from angular speed and background information

The effects of a background scene on the perception of the trajectory of an approaching object and its relation to changes in angular speed and angular size were examined in five experiments. Observers judged the direction (upward or downward) of two sequentially presented motion trajectories simulating a sphere traveling toward the observer at a constant 3-D speed from a fixed distance. In Experiments 1–4, we examined the effects of changes in angular speed and the presence of a scene background, with changes in angular size based either on the trajectories being discriminated or on an intermediate trajectory. In Experiment 5, we examined the effects of changes in angular speed and scene background, with angular size either constant or consistent with an intermediate 3-D trajectory. Overall, we found that (1) observers were able to judge the direction of object motion trajectories from angular speed changes; (2) observers were more accurate with a 3-D scene background, as compared with a uniform background, suggesting that scene information is important for recovering object motion trajectories; and (3) observers were more accurate in judging motion trajectories based on angular speed when the angular size function was consistent with motion in depth than when the angular size was constant.     

Statistical properties of staircase estimates

The bias and variability of staircase estimators were studied by means of repeated computer simulations of staircase runs. Both forced-choice and yes-no staircases were simulated. The influence of the shape of the psychometric function, the location and spacing of the stimuli, the number of trials in a run, and the method of deriving the estimate from the data are discussed. The forced-choice staircase is compared to the yes-no staircase, and the limitations of the simulation procedure are outlined.     

Eye-position aftereffects of backward head tilt manifested by illusory visual direction

Three experiments showed posttest-minus-pretest shifts in subjective straight-ahead eye position when subjects read for 3, 6, or 9 min with their heads tilted back 20° from upright. These shifts were significant relative to control conditions in which subjects read with their heads upright. All subjects read with the same straight-ahead eye-in-head position. Variability-reducing procedures were developed to provide better measures over Experiments 1, 2. and 3. Explanations in terms of deliberate compensation, head-position asymmetries, eye-position asymmetries, and progressive error were ruled out. It was hypothesized that the shifts were caused by negative aftereffects of compensation for the doll reflex. The doll reflex rotates the eyes down without central registration. causing an upward illusory shift of visual direction similar to what is caused by wedge prisms. Perceptual-motor adaptation to this shift, i.e., doll adaptation, causes an illusory shift in the opposite direction when the head is returned to upright.     

The influence of head contour and nose angle on the perception of eye-gaze direction

We report seven experiments that investigate the influence that head orientation exerts on the perception of eye-gaze direction. In each of these experiments, participants were asked to decide whether the eyes in a brief and masked presentation were looking directly at them or were averted. In each case, the eyes could be presented alone, or in the context of congruent or incongruent stimuli In Experiment 1A, the congruent and incongruent stimuli were provided by the orientation of face features and head outline. Discrimination of gaze direction was found to be better when face and gaze were congruent than in both of the other conditions, an effect that was not eliminated by inversion of the stimuli (Experiment 1B). In Experiment 2A, the internal face features were removed, but the outline of the head profile was found to produce an identical pattern of effects on gaze discrimination, effects that were again insensitive to inversion (Experiment 2B) and which persisted when lateral displacement of the eyes was controlled (Experiment 2C). Finally, in Experiment 3A, nose angle was also found to influence participants' ability to discriminate direct gaze from averted gaze, but here the effect was eliminated by inversion of the stimuli (Experiment 3B). We concluded that an image-based mechanism is responsible for the influence of head profile on gaze perception, whereas the analysis of nose angle involves the configural processing of face features.     

Spatial attentional distribution is modulated by head direction during eccentric gaze

Visual perception during eccentric gaze can be facilitated when a visual stimulus appears in front of the head direction. This study investigated the relative effects of gaze location and head direction on visual perception in central and peripheral vision. Participants identified the orientation of a T-shaped figure presented in the centre of a monitor and simultaneously localised a dot appearing in the periphery, while head direction relative to gaze location was to the left, right or centre. Effects of head direction were found only when the dot appeared far from the gaze fixation point, such that dot detection was superior when it appeared to the left (right) of fixation in the left (right) head direction. Experimental results indicated this was not due to a small shift of gaze location. Thus this study suggests that head direction influences visual perception particularly in peripheral vision where visual acuity decreases.     

Wrinkling and head shape as coordinated sources of age-level information

Changes in the shape of a human head and the development of facial wrinkles were examined as potential sources of information about age level. In Experiment 1, subjects estimated the ages of faces that had been produced by systematically manipulating characteristic head shapes and levels of wrinkles associated with ages 15, 30, 50, and 70 years. The results indicated that observers used both sources of craniofacial change in making age estimates; but the effect of either source of change on perceived age depended upon the level of the other source of change. In Experiment 2, subjects’ ratings of the apparent conflict between levels of head shape and wrinkles further substantiated the conclusion that observers are sensitive to the coordination of products of the two sources of change. These findings suggest that the information specifying perceived age level is a complexrelationship among different types of craniofacial change.     

The mutual influence of gaze and head orientation in the analysis of social attention direction

Three experiments are reported that investigate the hypothesis that head orientation and gaze direction interact in the processing of another individual's direction of social attention. A Stroop-type interference paradigm was adopted, in which gaze and head cues were placed into conflict. In separate blocks of trials, participants were asked to make speeded keypress responses contingent on either the direction of gaze, or the orientation of the head displayed in a digitized photograph of a male face. In Experiments 1 and 2, head and gaze cues showed symmetrical interference effects. Compared with congruent arrangements, incongruent head cues slowed responses to gaze cues, and incongruent gaze cues slowed responses to head cues, suggesting that head and gaze are mutually influential in the analysis of social attention direction. This mutuality was also evident in a cross-modal version of the task (Experiment 3) where participants responded to spoken directional words whilst ignoring the head/gaze images. It is argued that these interference effects arise from the independent influences of gaze and head orientation on decisions concerning social attention direction.     

Motion parallax judgements of depth as a function of the direction and type of head movement

We compared the relative effectiveness of rotating or translating the head, either horizontally or vertically, on the perception of depth resulting from motion parallax. Using Rogers and Graham's (1979) paradigm, we yoked the movement of random dots on a screen to movements of the head, simulating a corrugated surface. In two experiments, subjects nulled the apparent depth or motion seen in the display. Horizontal head movements yielded the most precise depth judgements, irrespective of whether the head translated or rotated. Motion thresholds were higher than those for depth and were independent of direction of head movement. In a third experiment, suprathreshold stimuli that simulated differing amounts of depth were used, and the subjects' perception of depth was virtually the same for all types and directions of head movement. In our stimulus situation, rotating or translating the head either vertically or horizontally produced motion parallax cues for depth that were equally effective. Our results also showed that, within a range, retinal image motion from head movement is converted into a depth signal and that above that range location constancy breaks down and motion is seen.     

The role of direction information in the perception of geometric optic flow components

Theoretically, optic flow, an important source of information for the perception of locomotion and three-dimensional structure of the environment, is described in terms of divergence, curl, and shear components. We measured how the detection of the type of flow field depends on directional information. We manipulated the local directions by rotating them through an angle x relative to the original direction (i.e., the direction of motion at that locus in an unaltered flow field). The results of the first experiments showed that divergence, curl, and shear can be detected even if the directional range of the individual motion vectors is as broad as 180 degrees. Subsequent experiments revealed that the detection of the geometric components of the optic flow field is merely based on the integration of a few (10% of vectors) local directions correctly (within 10 degrees of original direction) specifying the type of flow field. Other directions are irrelevant to this process. This is actually what one would expect if the optic flow is analyzed by special purpose mechanisms that detect and process the geometric components on the basis of the integration of motion information. The results indicate that as far as they integrate motion information, detectors for divergence, curl, and shear operate in a similar manner. Implications of the results for modeling such mechanisms are discussed.     

Adaptation to field displacement during head movement unrelated to the constancy of visual direction

Adaptation to vertical field displacements dependent on head turning about a vertical axis was demonstrated under two conditions, rapid training with 100 head movements and 1-h-long training with continuous head turning. The effect of rapid training was measured with the slant estimation method. Adaptation after the longer training was ascertained by comparing the uncertainty ranges for apparent target immobility before and after the adaptation period. Adaptation to field displacements in directions parallel to the plane of the head rotation obtained under corresponding conditions was also measured and found to be somewhat greater than adaptation to vertical field displacements. The result of work by Wallach and Frey that adaptation to field displacement in the direction with the head rotation is greater than to displacement against it was corroborated. While the previous result had, been obtained with rapid adaptation and with the slant estimation method, we confirmed it with 1-h training and by measuring the uncertainty ranges before and after the adaptation period.     

Social attention orienting integrates visual information from head and body orientation

Subjects were asked to detect visual, laterally presented reaction signals preceded by head-body cue stimuli in a spatial cueing task. A head rotated towards the reaction signal combined with a front view of a body resulted in shorter reaction times in comparison to the front view of a head and body. In contrast, a cue showing the head and body rotated towards the reaction signal did not result in such a facilitation in reaction times. The results suggest that the brain mechanisms involved in social attention orienting integrate ventrally processed visual information from the head and body orientation. A cue signaling that the other person, in his or her frame of reference, has an averted attention direction shifts the observer's own attention in the same direction.     

Vection during conflicting multisensory information about the axis, magnitude, and direction of self-motion

We examined the vection induced by consistent and conflicting multisensory information about self-motion. Observers viewed displays simulating constant-velocity self-motion in depth while physically oscillating their heads left-right or back-forth in time with a metronome. Their tracked head movements were either ignored or incorporated directly into the self-motion display (as an added simulated self-acceleration). When this head oscillation was updated into displays, sensory conflict was generated by simulating oscillation along: (i) an orthogonal axis to the head movement; or (ii) the same axis, but in a non-ecological direction. Simulated head oscillation always produced stronger vection than 'no display oscillation'--even when the axis/direction of this display motion was inconsistent with the physical head motion. When head-and-display oscillation occurred along the same axis: (i) consistent (in-phase) horizontal display oscillation produced stronger vection than conflicting (out-of-phase) horizontal display oscillation; however, (ii) consistent and conflicting depth oscillation conditions did not induce significantly different vection. Overall, orthogonal-axis oscillation was found to produce very similar vection to same-axis oscillation. Thus, we conclude that while vection appears to be very robust to sensory conflict, there are situations where sensory consistency improves vection.     

Controlling steering and judging heading: retinal flow,visual direction,and extraretinal information

The contribution of retinal flow (RF), extraretinal (ER), and egocentric visual direction (VD) information in locomotor control was explored. First, the recovery of heading from RF was examined when ER information was manipulated; results confirmed that ER signals affect heading judgments. Then the task was translated to steering curved paths, and the availability and veracity of VD were manipulated with either degraded or systematically biased RF. Large steering errors resulted from selective manipulation of RF and VD, providing strong evidence for the combination of RF, ER, and VD. The relative weighting applied to RF and VD was estimated. A point-attractor model is proposed that combines redundant sources of information for robust locomotor control with flexible trajectory planning through active gaze.     

Statistical properties of four effect-size measures for mediation models

This project examined the performance of classical and Bayesian estimators of four effect size measures for the indirect effect in a single-mediator model and a two-mediator model. Compared to the proportion and ratio mediation effect sizes, standardized mediation effect-size measures were relatively unbiased and efficient in the single-mediator model and the two-mediator model. Percentile and bias-corrected bootstrap interval estimates of ab/s _Y , and ab(s _X )/s _Y in the single-mediator model outperformed interval estimates of the proportion and ratio effect sizes in terms of power, Type I error rate, coverage, imbalance, and interval width. For the two-mediator model, standardized effect-size measures were superior to the proportion and ratio effect-size measures. Furthermore, it was found that Bayesian point and interval summaries of posterior distributions of standardized effect-size measures reduced excessive relative bias for certain parameter combinations. The standardized effect-size measures are the best effect-size measures for quantifying mediated effects.     

Psychophysics of perceiving eye-gaze and head direction with peripheral vision: implications for the dynamics of eye-gaze behavior

Two psychophysical experiments are reported, one dealing with the visual perception of the head orientation of another person (the 'looker') and the other dealing with the perception of the looker's direction of eye gaze. The participant viewed the looker with different retinal eccentricities, ranging from foveal to far-peripheral viewing. On average, judgments of head orientation were reliable even out to the extremes of peripheral vision (90 degrees eccentricity), with better performance at the extremes when the participant was able to view the looker changing head orientation from one trial to the next. In sharp contrast, judgments of eye-gaze direction were reliable only out to 4 degrees eccentricity, signifying that the eye-gaze social signal is available to people only when they fixate near the looker's eyes. While not unexpected, this vast difference in availability of information about head direction and eye direction, both of which can serve as indicators of the looker's focus of attention, is important for understanding the dynamics of eye-gaze behavior.