Multiple origins and modularity in the spatiotemporal emergence of cerebellar astrocyte heterogeneity.

The morphological, molecular, and functional heterogeneity of astrocytes is under intense scrutiny, but how this diversity is ontogenetically achieved remains largely unknown. Here, by quantitative in vivo clonal analyses and proliferation studies, we demonstrate that the major cerebellar astrocyte types emerge according to an unprecedented and remarkably orderly developmental program comprising (i) a time-dependent decline in both clone size and progenitor multipotency, associated with clone allocation first to the hemispheres and then to the vermis(ii) distinctive clonal relationships among astrocyte types, revealing diverse lineage potentials of embryonic and postnatal progenitors; and (iii) stereotyped clone architectures and recurrent modularities that correlate to layer-specific dynamics of postnatal proliferation/differentiation. In silico simulations indicate that the sole presence of a unique multipotent progenitor at the source of the whole astrogliogenic program is unlikely and rather suggest the involvement of additional committed components.

Videos look faster as children grow up: Sense of speed and impulsivity throughout primary school.

Previous research has documented systematic biases when adult observers judge the original speed of real-life video clips. What does the unfolding of events in videos-an increasingly pervasive "substitute reality"-look like as children grow up? The current study investigated the sense of speed during childhood and the relation with a number of performance and personality indexes. A group of 142 children aged 6-10 years adjusted the speed of video clips representing various scenes until reaching the apparent right speed. We found a tendency to underestimate speed; on average, videos played at their original speed were considered to be too slow. This tendency was larger in younger children and decreased with age (-3.16%/year). Uncertainty in judging video speed also decreased over the same age period (-10.79%/year). Children then performed a simple visuomotor task requiring response control, which revealed high accuracy and, in older children, faster responses. Children were also assessed for impulsivity/inattention and visuomotor habits through parents' questionnaires. When all variables were considered together, age and video game playing stood out as the only significant predictors of speed biases, both associated with an increase in apparent video speed. Thus, this study provides evidence of a change in the sense of speed during the primary school period, possibly involving the progressive overriding of a slow motion prior and/or the protracted calibration of perceptual mechanisms for speed constancy. The sense of speed, however, did not seem to be influenced by impulsivity traits in the population considered.

Enhancing Visual Exploration through Augmented Gaze: High Acceptance of Immersive Virtual Biking by Oldest Olds.

The diffusion of virtual reality applications dedicated to aging urges us to appraise its acceptance by target populations, especially the oldest olds. We investigated whether immersive virtual biking, and specifically a visuomotor manipulation aimed at improving visual exploration (augmented gaze), was well accepted by elders living in assisted residences. Twenty participants (mean age 89.8 years, five males) performed three 9 min virtual biking sessions pedalling on a cycle ergometer while wearing a Head-Mounted Display which immersed them inside a 360-degree pre-recorded biking video. In the second and third sessions, the relationship between horizontal head rotation and contingent visual shift was experimentally manipulated (augmented gaze), the visual shift being twice (gain = 2.0) or thrice (gain = 3.0) the amount of head rotation. User experience, motion sickness and visual exploration were measured. We found (i) very high user experience ratings, regardless of the gain; (ii) no effect of gain on motion sickness; and (iii) increased visual exploration (slope = +46%) and decreased head rotation (slope = -18%) with augmented gaze. The improvement in visual exploration capacity, coupled with the lack of intolerance signs, suggests that augmented gaze can be a valuable tool to improve the "visual usability" of certain virtual reality applications for elders, including the oldest olds.

Spotting expertise in the eyes: billiards knowledge as revealed by gaze shifts in a dynamic visual prediction task.

In sports, as in other activities and knowledge domains, expertise is a highly valuable asset. We assessed whether expertise in billiards is associated with specific patterns of eye movements in a visual prediction task. Professional players and novices were presented a number of simplified billiard shots on a computer screen, previously filmed in a real set, with the last part of the ball trajectory occluded. They had to predict whether or not the ball would have hit the central skittle. Experts performed better than novices, in terms of both accuracy and response time. By analyzing eye movements, we found that during occlusion, experts rarely extrapolated with the gaze the occluded part of the ball trajectory-a behavior that was widely diffused in novices-even when the unseen path was long and with two bounces interposed. Rather, they looked selectively at specific diagnostic points on the cushions along the ball's visible trajectory, in accordance with a formal metrical system used by professional players to calculate the shot coordinates. Thus, the eye movements of expert observers contained a clear signature of billiard expertise and documented empirically a strategy upgrade in visual problem solving from dynamic, analog simulation in imagery to more efficient rule-based, conceptual knowledge.

If You Are Old, Videos Look Slow. The Paradoxical Effect of Age-Related Motor Decline on the Kinematic Interpretation of Visual Scenes.

Perception and action are tightly coupled. However, there is still little recognition of how individual motor constraints impact perception in everyday life. Here we asked whether and how the motor slowing that accompanies aging influences the sense of visual speed. Ninety-four participants aged between 18 and 90 judged the natural speed of video clips reproducing real human or physical motion (SoS, Sense-of-Speed adjustment task). They also performed a finger tapping task and a visual search task, which estimated their motor speed and visuospatial attention speed, respectively. Remarkably, aged people judged videos to be too slow (speed underestimation), as compared to younger people: the Point of Subjective Equality (PSE), which estimated the speed bias in the SoS task, was +4% in young adults (<40), +12% in old adults (40-70) and +16% in elders. On average, PSE increased with age at a rate of 0.2% per year, with perceptual precision, adjustment rate, and completion time progressively worsening. Crucially, low motor speed, but not low attentional speed, turned out to be the key predictor of video speed underestimation. These findings suggest the existence of a counterintuitive compensatory coupling between action and perception in judging dynamic scenes, an effect that becomes particularly germane during aging.

Decoding overt shifts of attention in depth through pupillary and cortical frequency tagging.

Objective. We have recently developed a prototype of a novel human-computer interface for assistive communication based on voluntary shifts of attention (gaze) from a far target to a near target associated with a decrease of pupil size (Pupillary Accommodative Response, PAR), an automatic vegetative response that can be easily recorded. We report here an extension of that approach based on pupillary and cortical frequency tagging.Approach. In 18 healthy volunteers, we investigated the possibility of decoding attention shifts in depth by exploiting the evoked oscillatory responses of the pupil (Pupillary Oscillatory Response, POR, recorded through a low-cost device) and visual cortex (Steady-State Visual Evoked Potentials, SSVEP, recorded from 4 scalp electrodes). With a simple binary communication protocol (focusing on a far target meaning 'No', focusing on the near target meaning 'Yes'), we aimed at discriminating when observer's overt attention (gaze) shifted from the far to the near target, which were flickering at different frequencies.Main results. By applying a binary linear classifier (Support Vector Machine, SVM, with leave-one-out cross validation) to POR and SSVEP signals, we found that, with only twenty trials and no subjects' behavioural training, the offline median decoding accuracy was 75% and 80% with POR and SSVEP signals, respectively. When the two signals were combined together, accuracy reached 83%. The number of observers for whom accuracy was higher than 70% was 11/18, 12/18 and 14/18 with POR, SVVEP and combined features, respectively. A signal detection analysis confirmed these results.Significance. The present findings suggest that exploiting frequency tagging with pupillary or cortical responses during an attention shift in the depth plane, either separately or combined together, is a promising approach to realize a device for communicating with Complete Locked-In Syndrome (CLIS) patients when oculomotor control is unreliable and traditional assistive communication, even based on PAR, is unsuccessful.

Did i do that? Cognitive flexibility and self-agency in patients with obsessivecompulsive disorder.

Self-agency can be understood as the ability to infer causal relationships between actions and sensory events. Obsessive-Compulsive Disorder (OCD) patients with checking compulsions often report lack of "action-completion" sensations, possibly due to an altered sense of agency in these patients. The present study aimed to investigate whether self-agency was related to cognitive flexibility in OCD checkers. In 18 adult OCD checkers and 18 age- and gender-matched healthy controls, cognitive flexibility was assessed with the Intra-Extra Dimensional Set Shift Task (IED). Self-agency attribution was evaluated in two tasks that targeted the novel construct of "gaze-agency", the capability of an observer to identify his or her own eye movements as the cause of a concurrent event (here, an auditory beep). This technique allows sensitive measurement of agency under subtly varying investigator-controlled conditions. OCD checkers manifested significantly inferior performance correctly ascribing the beeps to their own ocular saccades than controls, even when after a hint was provided. Although cognitive inflexibility (errors on the IED) did not differ significantly between the two groups, within the OCD sample there were positive correlations between errors in self-agency attribution and total and extra-dimensional shift errors. These findings show that cognitive inflexibility is related to self-agency in OCD.

Cortical Resonance to Visible and Invisible Visual Rhythms.

Humans are rather poor in judging the right speed of video scenes. For example, a soccer match may be sped up so as to last only 80 min without observers noticing it. However, both adults and children seem to have a systematic, though often biased, notion of what should be the right speed of a given video scene. We therefore explored cortical responsiveness to video speed manipulations in search of possible differences between explicit and implicit speed processing. We applied sinusoidal speed modulations to a video clip depicting a naturalistic scene as well as a traditional laboratory visual stimulus (random dot kinematogram, RDK), and measured both perceptual sensitivity and cortical responses (steady-state visual evoked potentials, SSVEPs) to speed modulations. In five observers, we found a clear perceptual sensitivity increase and a moderate SSVEP amplitude increase with increasing speed modulation strength. Cortical responses were also found with weak, undetected speed modulations. These preliminary findings suggest that the cortex responds globally to periodic video speed modulations, even when observers do not notice them. This entrainment mechanism may be the basis of automatic resonance to the rhythms of the external world.

Blind saccades: an asynchrony between seeing and looking.

Saccades may not always wait for the completion of the perceptual analysis. By taking advantage of a motion-induced illusion of position and of the spontaneous scatter of saccade latency, we showed that in normal observers, regular saccades (latency, approximately 200 ms) were accurately directed to the target, whereas at higher latencies, saccades were increasingly biased by visual motion until they reflected the perceptual illusion. We reconstructed the time course of saccadic direction coding and identified an early phase in which saccades are mostly predictive (latencies less than approximately 100 ms), followed by a phase in which saccades are guided by the target position signal (latencies approximately 100-250 ms), and a later phase associated with the buildup of mislocalization (approximately 250-450 ms). This transient dissociation between action and perception indicates that seeing and looking are based on asynchronous processes, possibly because of independent thresholds for saccades and perceptual localization. The metrics of a saccade would then reflect the evolution of cortical visual signals from a predictive state to a perceptual state, passing through an intermediate visuomotor state. If saccades occur during the visuomotor state, they escape the tricks of perception.

Motion prediction at low contrast.

Accurate motion prediction is fundamental for survival. How does this reconcile with the well-known speed underestimation of low-contrast stimuli? Here we asked whether this contrast-dependent perceptual bias is retained in motion prediction under two different saccadic planning conditions: making a saccade to an occluded moving target, and real-time gaze interaction with multiple moving targets. In a first experiment, observers made a saccade to the mentally extrapolated position of a moving target (imagery condition). In a second experiment, observers had to prevent collisions among multiple moving targets by glancing at them through a gaze-contingent display or by hitting them with the touchpad cursor (interaction condition). In both experiments, target contrast was manipulated. We found that, whereas saccades to the imagined moving target were systematically biased by contrast, the gaze interaction performance, as measured by missed collisions, was generally unaffected - even though low-contrast targets looked slower. Interceptive actions increased at low contrast, but only when the gaze was used for interaction. Thus, perceptual speed underestimation transfers to saccades made to imagined low-contrast targets, without however necessarily being detrimental to effective performance when real-time interaction with multiple targets is required. This differential effect of stimulus contrast suggests that in complex dynamic conditions saccades are rather tolerant to visual speed biases.

Head movements modulate visual responsiveness in the absence of gaze shifts.

Visuospatial attention is strongly associated with saccades. Given that gaze shifts are often accomplished by combined eye-head movements, attention may also be coupled to head movements. We showed that simply turning the head without shifting the gaze is sufficient to cause a transient unbalance in responding to a visual stimulus. Manual responses to a stimulus flashed shortly before the onset of a horizontal head movement were faster in congruent trials, when the head moved towards the stimulus, than in incongruent trials, when the head moved away from the stimulus. These effects are similar to those observed for saccades. We take this as evidence for a tight link between visuospatial attention and head movements, even when the gaze does not shift.

Speed Biases With Real-Life Video Clips.

We live almost literally immersed in an artificial visual world, especially motion pictures. In this exploratory study, we asked whether the best speed for reproducing a video is its original, shooting speed. By using adjustment and double staircase methods, we examined speed biases in viewing real-life video clips in three experiments, and assessed their robustness by manipulating visual and auditory factors. With the tested stimuli (short clips of human motion, mixed human-physical motion, physical motion and ego-motion), speed underestimation was the rule rather than the exception, although it depended largely on clip content, ranging on average from 2% (ego-motion) to 32% (physical motion). Manipulating display size or adding arbitrary soundtracks did not modify these speed biases. Estimated speed was not correlated with estimated duration of these same video clips. These results indicate that the sense of speed for real-life video clips can be systematically biased, independently of the impression of elapsed time. Measuring subjective visual tempo may integrate traditional methods that assess time perception: speed biases may be exploited to develop a simple, objective test of reality flow, to be used for example in clinical and developmental contexts. From the perspective of video media, measuring speed biases may help to optimize video reproduction speed and validate "natural" video compression techniques based on sub-threshold temporal squeezing.

Low perceptual sensitivity to altered video speed in viewing a soccer match.

When watching videos, our sense of reality is continuously challenged. How much can a fundamental dimension of experience such as visual flow be modified before breaking the perception of real time? Here we found a remarkable indifference to speed manipulations applied to a popular video content, a soccer match. In a condition that mimicked real-life TV watching, none of 100 naïve observers spontaneously noticed speed alterations up/down to 12%, even when asked to report motion anomalies, and showed very low sensitivity to video speed changes (Just Noticeable Difference, JND = 18%). When tested with a constant-stimuli speed discrimination task, JND was still high, though much reduced (9%). The presence of the original voice-over with compensation for pitch did not affect perceptual performance. Thus, our results document a rather broad tolerance to speed manipulations in video viewing, even under attentive scrutiny. This finding may have important implications. For example, it can validate video compression strategies based on sub-threshold temporal squeezing. This way, a soccer match can last only 80 min and still be perceived as natural. More generally, knowing the boundaries of natural speed perception may help to optimize the flow of artificial visual stimuli which increasingly surround us.

Who Has Done It? Exploring Gaze Agency in Obsessive-Compulsive Checkers.

The sense of agency (SoA) is a multifaceted construct, which can be defined as the ability to understand the causal relationships between our actions and sensory events. Obsessive-Compulsive Disorder (OCD) patients with checking compulsions often report a "lack of action completion" sensations, which has been conceptualized in the so-called "Not Just Right Experiences" construct. An intriguing explanation of this phenomenon comes from Belayachi and Van der Linden (2009, 2010), who suggest that OCD-checking patients are more prone to specify their action in a relatively molecular and inflexible way. Currently, there are no studies in literature which address this issue in OCD patients, except for the one of Gentsch et al. (2012), who suggested an altered SoA in these patients. Here we exploited a novel construct, gaze agency, to evaluate causal attribution capabilities in a group of 21 OCD patients (checkers) and matched healthy controls (HCs). Basically, two tasks targeted observers' capability to identify their own eye movements as the cause of concurrently presented beeps, which allowed us to measure agency sensitivity as well as subtle agency alterations in an ecological setting. We found a poorer performance in OCD patients as compared to HCs in many parameters of our tasks, suggesting a difficulty with causal attribution possibly due to both a reduced cognitive flexibility and a less functional gaze agency in OCD patients.

Mental extrapolation of motion modulates responsiveness to visual stimuli.

Mental imagery is often considered to be an attentional state. We investigated whether imagining a stimulus in motion involves a corresponding movement of attention. Subjects fixating a central target extrapolated in imagery the motion of a spot that moved along a circular trajectory and then vanished. During imagery, a flash was presented with various backward and forward displacements relative to the direction of the imagined spot. Subjects had to make a saccade to the flash. Saccades were delayed by as much as 50 ms when the flash appeared displaced from the imagined spot, compared to when the flash was presented in its proximity. A similar delay in latency was obtained when subjects responded with a button press. In an "Observation" condition, in which the spot did not disappear, saccade latencies were similarly delayed, although mainly for backward flash displacements. These findings suggest that motion imagery is associated with a movement of visuospatial attention.

Covert enaction at work: Recording the continuous movements of visuospatial attention to visible or imagined targets by means of Steady-State Visual Evoked Potentials (SSVEPs).

Whereas overt visuospatial attention is customarily measured with eye tracking, covert attention is assessed by various methods. Here we exploited Steady-State Visual Evoked Potentials (SSVEPs) - the oscillatory responses of the visual cortex to incoming flickering stimuli - to record the movements of covert visuospatial attention in a way operatively similar to eye tracking (attention tracking), which allowed us to compare motion observation and motion extrapolation with and without eye movements. Observers fixated a central dot and covertly tracked a target oscillating horizontally and sinusoidally. In the background, the left and the right halves of the screen flickered at two different frequencies, generating two SSVEPs in occipital regions whose size varied reciprocally as observers attended to the moving target. The two signals were combined into a single quantity that was modulated at the target frequency in a quasi-sinusoidal way, often clearly visible in single trials. The modulation continued almost unchanged when the target was switched off and observers mentally extrapolated its motion in imagery, and also when observers pointed their finger at the moving target during covert tracking, or imagined doing so. The amplitude of modulation during covert tracking was ∼25-30% of that measured when observers followed the target with their eyes. We used 4 electrodes in parieto-occipital areas, but similar results were achieved with a single electrode in Oz. In a second experiment we tested ramp and step motion. During overt tracking, SSVEPs were remarkably accurate, showing both saccadic-like and smooth pursuit-like modulations of cortical responsiveness, although during covert tracking the modulation deteriorated. Covert tracking was better with sinusoidal motion than ramp motion, and better with moving targets than stationary ones. The clear modulation of cortical responsiveness recorded during both overt and covert tracking, identical for motion observation and motion extrapolation, suggests to include covert attention movements in enactive theories of mental imagery.

Assessing Self-Awareness through Gaze Agency.

We define gaze agency as the awareness of the causal effect of one's own eye movements in gaze-contingent environments, which might soon become a widespread reality with the diffusion of gaze-operated devices. Here we propose a method for measuring gaze agency based on self-monitoring propensity and sensitivity. In one task, naïf observers watched bouncing balls on a computer monitor with the goal of discovering the cause of concurrently presented beeps, which were generated in real-time by their saccades or by other events (Discovery Task). We manipulated observers' self-awareness by pre-exposing them to a condition in which beeps depended on gaze direction or by focusing their attention to their own eyes. These manipulations increased propensity to agency discovery. In a second task, which served to monitor agency sensitivity at the sensori-motor level, observers were explicitly asked to detect gaze agency (Detection Task). Both tasks turned out to be well suited to measure both increases and decreases of gaze agency. We did not find evident oculomotor correlates of agency discovery or detection. A strength of our approach is that it probes self-monitoring propensity-difficult to evaluate with traditional tasks based on bodily agency. In addition to putting a lens on this novel cognitive function, measuring gaze agency could reveal subtle self-awareness deficits in pathological conditions and during development.

Both vision-for-perception and vision-for-action follow Weber's law at small object sizes, but violate it at larger sizes.

According to a previous report, the visual coding of size does not obey Weber's law when aimed at guiding a grasp (Ganel et al., 2008a). This result has been interpreted as evidence for a fundamental difference between sensory processing in vision-for-perception, which needs to compress a wide range of physical objects to a restricted range of percepts, and vision-for-action when applied to the much narrower range of graspable and reachable objects. We compared finger aperture in a motor task (precision grip) and perceptual task (cross modal matching or "manual estimation" of the object's size). Crucially, we tested the whole range of graspable objects. We report that both grips and estimations clearly violate Weber's law with medium-to-large objects, but are essentially consistent with Weber's law with smaller objects. These results differ from previous characterizations of perception-action dissociations in the precision of representations of object size. Implications for current functional interpretations of the dorsal and ventral processing streams in the human visual system are discussed.

Smooth pursuit-like eye movements during mental extrapolation of motion: the facilitatory effect of drowsiness.

We studied eye movements during mental extrapolation of motion. Subjects tracked with the eyes a target oscillating sinusoidally by +/- 5 degrees on the horizontal plane at frequencies between 0.15 and 0.5 Hz. After 4 cycles the target disappeared and subjects had to mentally extrapolate its motion in imagery for 5 further cycles. Subjects were invited to relax. We recorded eye movements with infrared oculography and the lid aperture with a webcam. Typically, in the imagery phase, sequences of saccades "reproduced" the macroscopic characteristics of the to-be-imagined motion. However, slow eye movements substituting the sequence of saccades emerged repeatedly in the course of the experiment, to the extent that sometimes it was almost impossible to distinguish the pattern of eye movements during tracking from that during imagery. These smooth pursuit-like eye movements lasted up to 17 s, although they were more commonly intermingled with saccades. It turned out that this phenomenon occurred mostly, although not exclusively, when the lids were partially lowered suggesting an association with drowsiness. The average gain of smooth pursuit-like eye movements in imagery passed from 0.09 when the lids were open, to 0.27 when they were partially lowered. A control experiment excluded that the lowering of the lids per se had played any important role. This finding has implications for both the physiology of the oculomotor system and for the debate on the relation between imagery and perception.