Sensitivity of occipito-temporal cortex, premotor and Broca's areas to visible speech gestures in a familiar language.

When looking at a speaking person, the analysis of facial kinematics contributes to language discrimination and to the decoding of the time flow of visual speech. To disentangle these two factors, we investigated behavioural and fMRI responses to familiar and unfamiliar languages when observing speech gestures with natural or reversed kinematics. Twenty Italian volunteers viewed silent video-clips of speech shown as recorded (Forward, biological motion) or reversed in time (Backward, non-biological motion), in Italian (familiar language) or Arabic (non-familiar language). fMRI revealed that language (Italian/Arabic) and time-rendering (Forward/Backward) modulated distinct areas in the ventral occipito-temporal cortex, suggesting that visual speech analysis begins in this region, earlier than previously thought. Left premotor ventral (superior subdivision) and dorsal areas were preferentially activated with the familiar language independently of time-rendering, challenging the view that the role of these regions in speech processing is purely articulatory. The left premotor ventral region in the frontal operculum, thought to include part of the Broca's area, responded to the natural familiar language, consistent with the hypothesis of motor simulation of speech gestures.

Wildlife and Bait Density Monitoring to Describe the Effectiveness of a Rabies Vaccination Program in Foxes.

Fox rabies has been eliminated from vast areas of West and Central Europe, but cases still occur in the Balkans. Oral vaccination is an effective method for reducing the incidence of the disease in wildlife, but it requires monitoring if bait density is adequate for the density of the wildlife reservoir. We developed a methodology to evaluate the effectiveness of aerial vaccination campaigns conducted in Montenegro during autumn 2011 and spring 2012. The effectiveness of the vaccination campaign was assessed by (i) estimating the density of baits, (ii) estimating the distribution of the red fox, (iii) identifying critical areas of insufficient bait density by combining both variables. Although the two vaccination campaigns resulted in 45% and 47% of the country's total area not reaching recommended density of 20 baits/km2, the consecutive delivery of both campaigns reduced these "gaps" to 6%. By combining bait and reservoir density data, we were able to show that bait density was lower than fox density in only 5% of Montenegro's territory. The methodology described can be used for real-time evaluation of aerial vaccine delivery campaigns, to identify areas with insufficient bait densities.

Drawing ellipses in water: evidence for dynamic constraints in the relation between velocity and path curvature.

Several types of continuous human movements comply with the so-called Two-Thirds Power Law (2/3-PL) stating that velocity (V) is a power function of the radius of curvature (R) of the endpoint trajectory. The origin of the 2/3-PL has been the object of much debate. An experiment investigated further this issue by comparing two-dimensional drawing movements performed in air and water. In both conditions, participants traced continuously quasi-elliptic trajectories (period T = 1.5 s). Other experimental factors were the movement plane (horizontal/vertical), and whether the movement was performed free-hand, or by following the edge of a template. In all cases a power function provided a good approximation to the V-R relation. The main result was that the exponent of the power function in water was significantly smaller than in air. This appears incompatible with the idea that the power relationship depends only on kinematic constraints and suggests a significant contribution of dynamic factors. We argue that a satisfactory explanation of the observed behavior must take into account the interplay between the properties of the central motor commands and the visco-elastic nature of the mechanical plant that implements the commands.

Grip forces during fast point-to-point and continuous hand movements.

Three experiments investigated the grip force exerted by the fingers on an object displaced actively in the near-body space. In one condition (unimanual) the object was held by one hand with the tripod grip and was moved briskly back and forth along one of the three coordinate directions (up-down, left-right, near-far). In the second condition (bimanual) the same point-to-point movements were performed while holding the object with the index and middle fingers of both hands. In the third condition (bimanual) the object was held as in the second condition and moved along a circular path lying in one of the three coordinate planes (horizontal, frontal, sagittal). In all conditions participants were asked to exert a baseline level of grip force largely exceeding the safety margin against slippage. Both grip forces and hand displacements were measured with high accuracy. As reported in previous studies, in the two point-to-point conditions we observed an upsurge of the grip force at the onset and at the end the movements. However, the timing of the transient increases of the grip force relative to hand kinematics did not confirm the hypothesis set forth by several previous studies that grip modulation is a pre-planned action based on an internal model of the expected effects of the movement. In the third condition, the systematic modulation of the grip force also for circular movements was again at variance with the internal model hypothesis because it cannot be construed as a pre-planned action aiming at countering large changes in dynamic load. We argue that a parsimonious account of the covariations of load and grip forces can be offered by taking into account the visco-elastic properties of the neuromuscular system.

Unfamiliar Walking Movements Are Detected Early in the Visual Stream: An fMRI Study.

Two experiments investigated the network involved in the visual perception of walking. Video clips of forward and backward walk (real walk direction) were shown either as recorded, or reversed in time (rendering). In Experiment 1 (identification task), participants were asked to indicate whether or not the stimulus was time-reversed. In Experiment 2 (free-viewing), participants viewed the video clips passively. Identification accuracy was good with the more familiar scene, that is, when the visual walk was in the direction of the facing orientation, and at chance level in the opposite case. In both experiments, the temporo-occipital junction (TOJ) was activated more strongly by unfamiliar than familiar scenes. Only in Experiment 1 intraparietal, superior temporal, and inferior temporal regions were also activated. TOJ activation signals the detection in unfamiliar scenes of a mismatch between facing orientation and visual movement direction. We argue that TOJ response to a mismatch prevents the further processing of the visual input required to identify temporal inversions. When no mismatch is detected (familiar stimuli), TOJ would, instead, be involved in the kinematic analysis that makes such identification possible. The study demonstrates that unfamiliar walking movements are detected earlier than so far assumed along the visual movement processing stream.

Implied dynamics biases the visual perception of velocity.

We expand the anecdotic report by Johansson that back-and-forth linear harmonic motions appear uniform. Six experiments explore the role of shape and spatial orientation of the trajectory of a point-light target in the perceptual judgment of uniform motion. In Experiment 1, the target oscillated back-and-forth along a circular arc around an invisible pivot. The imaginary segment from the pivot to the midpoint of the trajectory could be oriented vertically downward (consistent with an upright pendulum), horizontally leftward, or vertically upward (upside-down). In Experiments 2 to 5, the target moved uni-directionally. The effect of suppressing the alternation of movement directions was tested with curvilinear (Experiment 2 and 3) or rectilinear (Experiment 4 and 5) paths. Experiment 6 replicated the upright condition of Experiment 1, but participants were asked to hold the gaze on a fixation point. When some features of the trajectory evoked the motion of either a simple pendulum or a mass-spring system, observers identified as uniform the kinematic profiles close to harmonic motion. The bias towards harmonic motion was most consistent in the upright orientation of Experiment 1 and 6. The bias disappeared when the stimuli were incompatible with both pendulum and mass-spring models (Experiments 3 to 5). The results are compatible with the hypothesis that the perception of dynamic stimuli is biased by the laws of motion obeyed by natural events, so that only natural motions appear uniform.

The identification of unfolding facial expressions.

We asked whether the identification of emotional facial expressions (FEs) involves the simultaneous perception of the facial configuration or the detection of emotion-specific diagnostic cues. We recorded at high speed (500 frames s-1) the unfolding of the FE in five actors, each expressing six emotions (anger, surprise, happiness, disgust, fear, sadness). Recordings were coded every 10 frames (20 ms of real time) with the Facial Action Coding System (FACS, Ekman et al 2002, Salt Lake City, UT: Research Nexus eBook) to identify the facial actions contributing to each expression, and their intensity changes over time. Recordings were shown in slow motion (1/20 of recording speed) to one hundred observers in a forced-choice identification task. Participants were asked to identify the emotion during the presentation as soon as they felt confident to do so. Responses were recorded along with the associated response times (RTs). The RT probability density functions for both correct and incorrect responses were correlated with the facial activity during the presentation. There were systematic correlations between facial activities, response probabilities, and RT peaks, and significant differences in RT distributions for correct and incorrect answers. The results show that a reliable response is possible long before the full FE configuration is reached. This suggests that identification is reached by integrating in time individual diagnostic facial actions, and does not require perceiving the full apex configuration.

The perception of visible speech: estimation of speech rate and detection of time reversals.

Four experiments investigated the perception of visible speech. Experiment 1 addressed the perception of speech rate. Observers were shown video-clips of the lower face of actors speaking at their spontaneous rate. Then, they were shown muted versions of the video-clips, which were either accelerated or decelerated. The task (scaling) was to compare visually the speech rate of the stimulus to the spontaneous rate of the actor being shown. Rate estimates were accurate when the video-clips were shown in the normal direction (forward mode). In contrast, speech rate was underestimated when the video-clips were shown in reverse (backward mode). Experiments 2-4 (2AFC) investigated how accurately one discriminates forward and backward speech movements. Unlike in Experiment 1, observers were never exposed to the sound track of the video-clips. Performance was well above chance when playback mode was crossed with rate modulation, and the number of repetitions of the stimuli allowed some amount of speechreading to take place in forward mode (Experiment 2). In Experiment 3, speechreading was made much more difficult by using a different and larger set of muted video-clips. Yet, accuracy decreased only slightly with respect to Experiment 2. Thus, kinematic rather then speechreading cues are most important for discriminating movement direction. Performance worsened, but remained above chance level when the same stimuli of Experiment 3 were rotated upside down (Experiment 4). We argue that the results are in keeping with the hypothesis that visual perception taps into implicit motor competence. Thus, lawful instances of biological movements (forward stimuli) are processed differently from backward stimuli representing movements that the observer cannot perform.

Detecting temporal reversals in human locomotion.

An experiment investigated the ability by human observers to detect temporal reversals in dynamic displays of human locomotion. We video-taped the lower portion of the body of actors walking at their preferred speed either in the normal, forward direction (FW) or in the backward direction (BW). The videos were presented in a random order either as recorded (N) or in reverse (R). In one session, we presented both normal and time-reversed stimuli in the original upright orientation. In a second session, the stimuli were rotated by 180° around the horizontal axis. Observers were informed that the real recorded movement was either forward or backward and were asked to decide whether or not the movement had been time-reversed prior to the presentation. Although the kinematics of forward and backward human locomotion is quite similar, the detection of temporal reversals followed a consistent pattern showing a good accuracy in condition FW-N and a reduced but still above-chance performance in condition BW-R (by design, in both conditions actors appeared to walk forward). Performance was instead at chance level in the other two conditions where the apparent direction of the movement was backward. Inverting the spatial orientation of the stimuli reduced but did not suppress the ability to detect temporal reversals in the two conditions with apparent forward direction of movement. It is argued that implicit motor competence is at least in part instrumental for extracting the subtle discriminal information from the stimuli.

Is there a dynamic advantage for facial expressions?

Some evidence suggests that it is easier to identify facial expressions (FEs) shown as dynamic displays than as photographs (dynamic advantage hypothesis). Previously, this has been tested by using dynamic FEs simulated either by morphing a neutral face into an emotional one or by computer animations. For the first time, we tested the dynamic advantage hypothesis by using high-speed recordings of actors' FEs. In the dynamic condition, stimuli were graded blends of two recordings (duration: 4.18 s), each describing the unfolding of an expression from neutral to apex. In the static condition, stimuli (duration: 3 s) were blends of just the apex of the same recordings. Stimuli for both conditions were generated by linearly morphing one expression into the other. Performance was estimated by a forced-choice task asking participants to identify which prototype the morphed stimulus was more similar to. Identification accuracy was not different between conditions. Response times (RTs) measured from stimulus onset were shorter for static than for dynamic stimuli. Yet, most responses to dynamic stimuli were given before expressions reached their apex. Thus, with a threshold model, we tested whether discriminative information is integrated more effectively in dynamic than in static conditions. We did not find any systematic difference. In short, neither identification accuracy nor RTs supported the dynamic advantage hypothesis.

Effect of selective and distributed training on visual identification of orientation.

An experiment contrasted the effect of four training schedules in a visual orientation reproduction task. Two selective schedules involved repeated presentation of a single target orientation. Two non-selective schedules involved targets covering the first quadrant either at fixed, equispaced orientations, or distributed randomly. In pre-training sessions, we observed the classical oblique effect (precision for vertical and horizontal stimuli higher than for oblique ones). Practice improved precision with both distributed schedules, but was ineffectual for non-selective schedules. However, a significant oblique effect persisted under all conditions. We argue that the pattern of results is compatible with the hypothesis that the oblique effect reflects both the intrinsic neuronal properties of the primary visual system, and the structure of the visual space imposed by higher, more cognitive processes. The results challenge the thesis that only attentional and post-perceptual factors are able to affect the working of the early visual system.

Velocity control in Parkinson's disease: a quantitative analysis of isochrony in scribbling movements.

An experiment was conducted to contrast the motor performance of three groups (N = 20) of participants: (1) patients with confirmed Parkinson Disease (PD) diagnose; (2) age-matched controls; (3) young adults. The task consisted of scribbling freely for 10 s within circular frames of different sizes. Comparison among groups focused on the relation between the figural elements of the trace (overall size and trace length) and the velocity of the drawing movements. Results were analysed within the framework of previous work on normal individuals showing that instantaneous velocity of drawing movements depends jointly on trace curvature (Two-thirds Power Law) and trace extent (Isochrony principle). The motor behaviour of PD patients exhibited all classical symptoms of the disease (reduced average velocity, reduced fluency, micrographia). At a coarse level of analysis both isochrony and the dependence of velocity on curvature, which are supposed to reflect cortical mechanisms, were spared in PD patients. Instead, significant differences with respects to the control groups emerged from an in-depth analysis of the velocity control suggesting that patients did not scale average velocity as effectively as controls. We factored out velocity control by distinguishing the influence of the broad context in which movement is planned--i.e. the size of the limiting frames--from the influence of the local context--i.e. the linear extent of the unit of motor action being executed. The balance between the two factors was found to be distinctively different in PD patients and controls. This difference is discussed in the light of current theorizing on the role of cortical and sub-cortical mechanisms in the aetiology of PD. We argue that the results are congruent with the notion that cortical mechanisms are responsible for generating a parametric template of the desired movement and the BG specify the actual spatio-temporal parameters through a multiplicative gain factor acting on both size and velocity.

Factors affecting the size of the detour effect in the kinaesthetic perception of Euclidean distance.

Three experiments investigated the mechanisms by which we estimate Euclidean distances on the basis of kinaesthetic cues. In all experiments, blindfolded participants followed straight and curvilinear paths with a stylus. Then, with a straight response movement, they estimated the distance between the end-points of the previously explored path. Experiment 1 was designed to validate the hypothesis-made on the basis of results from a previous study-that errors in the kinaesthetic estimations of distances (detour effect) originate from the difficulty to decompose the displacement vector into relevant and irrelevant components, which would become more severe at points of inflection. Using elliptic paths (no inflections), we demonstrated that errors are indeed reduced considerably. The role of the orientation of the work plane was investigated in Experiment 2 in which the same paths used in our previous study were oriented in the frontal rather than the horizontal plane. The results indicate that the detour effect is independent of the orientation. Moreover, despite the asymmetry that gravity introduces between upward and downward movements, errors in the two directions are almost identical. Experiment 3 addressed two issues. First, we demonstrated that introducing a delay between the exploration of the path and the response did not alter significantly the pattern of errors. By contrast, we demonstrated that errors are severely reduced when the number of paths to be explored is reduced by half. The results of the three experiments are discussed within the context of current theories of sensori-motor coding.

Error parsing in visuomotor pointing reveals independent processing of amplitude and direction.

An experiment investigated systematic pointing errors in horizontal movements performed without visual feedback toward 48 targets placed symmetrically around two initial hand positions. Our main goal was to provide evidence in favor of the hypothesis that amplitude and direction of the movements are planned independently on the basis of the hand-target vector (vectorial parametric hypothesis, VP). The analysis was carried out mainly at the individual level. By screening a number of formal models of the potential error components, we found that only models compatible with the VP hypothesis provide an accurate description of the error pattern. A quantitative analysis showed that errors are explained mostly by a bias in the represented initial hand position (46% of the sum of squared errors) and a visuomotor gain bias (26%). Range effect (3%), directional biases (3%), and inertia-dependent amplitude modulations (1%) also provided significant contributions. The error pattern was incompatible with the view that movements are planned by specifying either a final posture or a final position. Instead, the results fully supported the view that, at least in the horizontal plane, amplitude, and direction of pointing movements are planned independently in a hand- or target-centered frame of reference.

Planning short pointing sequences.

An experiment tested the hypothesis that fast, short sequences of movements are planned as a whole, before movement inception. The experimental task consisted of pointing to either one (one-step condition), or two (two-step condition) visual targets aligned along the mid-sagittal axis in a horizontal plane. There were nine possible arrangements of the targets resulting from all combinations of three distances (5, 10, 15 cm), and two trial orders (blocked or random). Performances were characterised by reaction time (RT), movement kinematics, and spatial accuracy. Compared with one-step trials, the first movements of two-step trials had longer RTs (length effect), particularly in random sessions, and when the sequences included short-distance targets. There were also differences in duration (one-target advantage), velocity profile and spatial accuracy that did not depend on the characteristics of the second movement. The results are inconsistent with the assumption that two-step sequences are planned as a whole. Instead, they are in keeping with the alternative hypothesis that part of the preparation of the second step takes place during the execution of the first step.

Perceptual asynchronies for biological and non-biological visual events.

Four experiments investigated the hypothesis that different attributes of a visual scene are processed by independent channels working asynchronously. Experiment 1 considered the attributes of colour, form, and movement of simple geometrical configurations. In each of three conditions, two of these attributes switched simultaneously between two fixed values (Green/Red, Circle/Square, Fixed/Moving). Participants indicated which of the two attributes changes was perceptually closer in time to a sound signal. Response probabilities varied as a function of the time of occurrence of the sound, showing that the processing of the movement channel is delayed with respect to the other two. A smaller but significant difference was also detected between the processing times for colour and form. Comparing Experiments 1 and 2 showed that movement velocity does not affect the delay with which movement onset is perceived with respect to colour. Experiment 3 contrasted colour and movement in the perception of a biological movement. The stimuli were video clips of a coloured ball being lifted by a hand. The colour of the ball changed a variable amount of time before or after the ball started moving. Participants indicated which of the two changes had occurred first. We found that, unlike in Experiments 1 and 2, movement perception no longer lagged colour perception. Experiment 4 tested the hypothesis that the disappearance of the asynchrony is due to perceptual anticipation. We discuss the implications of the results vis-à-vis current theories on perceptual binding and on the coding of dynamic events.

Amplitude and direction errors in kinesthetic pointing.

We investigated the accuracy with which, in the absence of vision, one can reach again a 2D target location that had been previously identified by a guided movement. A robotic arm guided the participant's hand to a target (locating motion) and away from it (homing motion). Then, the participant pointed freely toward the remembered target position. Two experiments manipulated separately the kinematics of the locating and homing motions. Some robot motions followed a straight path with the bell-shaped velocity profile that is typical of natural movements. Other motions followed curved paths, or had strong acceleration and deceleration peaks. Current motor theories of perception suggest that pointing should be more accurate when the homing and locating motion mimics natural movements. This expectation was not borne out by the results, because amplitude and direction errors were almost independent of the kinematics of the locating and homing phases. In both experiments, participants tended to overshoot the target positions along the lateral directions. In addition, pointing movements towards oblique targets were attracted by the closest diagonal (oblique effect). This error pattern was robust not only with respect to the manner in which participants located the target position (perceptual equivalence), but also with respect to the manner in which they executed the pointing movements (motor equivalence). Because of the similarity of the results with those of previous studies on visual pointing, it is argued that the observed error pattern is basically determined by the idiosyncratic properties of the mechanisms whereby space is represented internally.

The kinaesthetic perception of Euclidean distance: a study of the detour effect.

An experiment investigated the mechanisms by which humans estimate Euclidean distances on the basis of kinaesthetic cues. Blindfolded participants followed straight and curvilinear paths with a hand-held stylus (encoding phase). Then, with a straight movement, they estimated the Euclidean distance between the start- and end-points of the path (response phase). The experiment contrasted an On-axis condition, in which encoding and response movements were spatially aligned, and an Off-axis condition, in which they were displaced laterally. Performances were slightly more accurate in the On-axis condition than in the Off-axis condition. In both conditions, however, errors were consistently smaller when the path covered a larger surface. The results showed that small paths yielded an overestimation of the Euclidean distance, the relative errors increasing with the length of curvilinear paths. The findings are compared with results of other studies in which distances were estimated on the basis of haptic cues.

Altering the visuomotor gain. Evidence that motor plans deal with vector quantities.

Two experiments investigated the effects of providing nonveridical knowledge of the results (KR) in a visuomanual task in which participants pointed to briefly (200 ms) presented targets without seeing their hand. By showing after each trial the movement endpoint displaced radially with respect to its true position, we were able to alter progressively the gain of the visuomanual loop. In experiment 1, the KR was provided only for transversal movements and for one target distance, but the effect generalized to all directions and all distances. Moreover, it also generalized to the other hand that had never been biased. In experiment 2, nonveridical KR was supplied for movements along the two major diagonals which require sharply different muscle synergies. The transfer to other directions and to the other hand was equally substantial. It is argued that the results support the vector coding hypothesis, which holds that the input to the motor execution stage is supplied by specifying independently the amplitude and the direction of the vector from the initial to the final position in an extrinsic frame of reference. We also discuss the possible brain structures involved in the biasing action of the KR.