How do hoverflies use their righting reflex?

When taking off from a sloping surface, flies have to reorient themselves dorsoventrally and stabilize their body by actively controlling their flapping wings. We have observed that righting is achieved solely by performing a rolling manoeuvre. How flies manage to do this has not yet been elucidated. It was observed here for the first time that hoverfly reorientation is entirely achieved within 6 wingbeats (48.8 ms) at angular roll velocities of up to 10×103 deg s-1 and that the onset of their head rotation consistently follows that of their body rotation after a time lag of 16 ms. The insects' body roll was found to be triggered by the asymmetric wing stroke amplitude, as expected. The righting process starts immediately with the first wingbeat and seems unlikely to depend on visual feedback. A dynamic model for the fly's righting reflex is presented, which accounts for the head/body movements and the time lag recorded in these experiments. This model consists of a closed-loop control of the body roll, combined with a feedforward control of the head/body angle. During the righting manoeuvre, a strong coupling seems to exist between the activation of the halteres (which measure the body's angular speed) and the gaze stabilization reflex. These findings again confirm the fundamental role played by the halteres in both body and head stabilization processes.

Individual movement features during prism adaptation correlate with after-effects and interlimb transfer.

The human nervous system displays such plasticity that we can adapt our motor behavior to various changes in environmental or body properties. However, how sensorimotor adaptation generalizes to new situations and new effectors, and which factors influence the underlying mechanisms, remains unclear. Here we tested the general hypothesis that differences across participants can be exploited to uncover what drives interlimb transfer. Twenty healthy adults adapted to prismatic glasses while reaching to visual targets with their dominant arm. Classic adaptation and generalization across movement directions were observed but transfer to the non-dominant arm was not significant and inter-individual differences were substantial. Interlimb transfer resulted for some participants in a directional shift of non-dominant arm movements that was consistent with an encoding of visuomotor adaptation in extrinsic coordinates. For some other participants, transfer was consistent with an intrinsic coordinate system. Simple and multiple regression analyses showed that a few kinematic parameters such as peak acceleration (or peak velocity) and variability of movement direction were correlated with interlimb transfer. Low peak acceleration and low variability were related to extrinsic transfer, while high peak acceleration and high variability were related to intrinsic transfer. Motor variability was also positively correlated with the magnitude of the after-effect systematically observed on the dominant arm. Overall, these findings on unconstrained movements support the idea that individual movement features could be linked to the sensorimotor adaptation and its generalization. The study also suggests that distinct movement characteristics may be related to different coordinate frames of action representations in the nervous system.

Modeling visual-based pitch, lift and speed control strategies in hoverflies.

To avoid crashing onto the floor, a free falling fly needs to trigger its wingbeats quickly and control the orientation of its thrust accurately and swiftly to stabilize its pitch and hence its speed. Behavioural data have suggested that the vertical optic flow produced by the fall and crossing the visual field plays a key role in this anti-crash response. Free fall behavior analyses have also suggested that flying insect may not rely on graviception to stabilize their flight. Based on these two assumptions, we have developed a model which accounts for hoverflies´ position and pitch orientation recorded in 3D with a fast stereo camera during experimental free falls. Our dynamic model shows that optic flow-based control combined with closed-loop control of the pitch suffice to stabilize the flight properly. In addition, our model sheds a new light on the visual-based feedback control of fly´s pitch, lift and thrust. Since graviceptive cues are possibly not used by flying insects, the use of a vertical reference to control the pitch is discussed, based on the results obtained on a complete dynamic model of a virtual fly falling in a textured corridor. This model would provide a useful tool for understanding more clearly how insects may or not estimate their absolute attitude.

Role of the light source position in freely falling hoverflies' stabilization performances.

The stabilization of plummeting hoverflies was filmed and analysed in terms of their wingbeat initiation times as well as the crash and stabilization rates. The flies experienced near-weightlessness for a period of time that depended on their ability to counteract the free fall by triggering their wingbeats. In this paradigm, hoverflies' flight stabilization strategies were investigated here for the first time under two different positions of the light source (overhead and bottom lighting). The crash rates were higher in bottom lighting conditions than with top lighting. In addition, adding a texture to the walls reduced the crash rates only in the overhead lighting condition. The position of the lighting also significantly affected both the stabilization rates and the time taken by the flies to stabilize, which decreased and increased under bottom lighting conditions, respectively, whereas textured walls increased the stabilization rates under both lighting conditions. These results support the idea that flies may mainly base their flight control strategy on visual cues and particularly that the light distribution in the visual field may provide reliable, efficient cues for estimating their orientation with respect to an allocentric reference frame. In addition, the finding that the hoverflies' optic flow-based motion detection ability is affected by the position of the light source in their visual field suggests the occurrence of interactions between movement perception and this visual vertical perception process.

To crash or not to crash: how do hoverflies cope with free-fall situations and weightlessness?

Insects' aptitude to perform hovering, automatic landing and tracking tasks involves accurately controlling their head and body roll and pitch movements, but how this attitude control depends on an internal estimation of gravity orientation is still an open question. Gravity perception in flying insects has mainly been studied in terms of grounded animals' tactile orientation responses, but it has not yet been established whether hoverflies use gravity perception cues to detect a nearly weightless state at an early stage. Ground-based microgravity simulators provide biologists with useful tools for studying the effects of changes in gravity. However, in view of the cost and the complexity of these set-ups, an alternative Earth-based free-fall procedure was developed with which flying insects can be briefly exposed to microgravity under various visual conditions. Hoverflies frequently initiated wingbeats in response to an imposed free fall in all the conditions tested, but managed to avoid crashing only in variably structured visual environments, and only episodically in darkness. Our results reveal that the crash-avoidance performance of these insects in various visual environments suggests the existence of a multisensory control system based mainly on vision rather than gravity perception.

To transfer or not to transfer? Kinematics and laterality quotient predict interlimb transfer of motor learning.

Humans can remarkably adapt their motor behavior to novel environmental conditions, yet it remains unclear which factors enable us to transfer what we have learned with one limb to the other. Here we tested the hypothesis that interlimb transfer of sensorimotor adaptation is determined by environmental conditions but also by individual characteristics. We specifically examined the adaptation of unconstrained reaching movements to a novel Coriolis, velocity-dependent force field. Right-handed subjects sat at the center of a rotating platform and performed forward reaching movements with the upper limb toward flashed visual targets in prerotation, per-rotation (i.e., adaptation), and postrotation tests. Here only the dominant arm was used during adaptation and interlimb transfer was assessed by comparing performance of the nondominant arm before and after dominant-arm adaptation. Vision and no-vision conditions did not significantly influence interlimb transfer of trajectory adaptation, which on average was significant but limited. We uncovered a substantial heterogeneity of interlimb transfer across subjects and found that interlimb transfer can be qualitatively and quantitatively predicted for each healthy young individual. A classifier showed that in our study, interlimb transfer could be predicted based on the subject's task performance, most notably motor variability during learning, and his or her laterality quotient. Positive correlations suggested that variability of motor performance and lateralization of arm movement control facilitate interlimb transfer. We further show that these individual characteristics can predict the presence and the magnitude of interlimb transfer of left-handers. Overall, this study suggests that individual characteristics shape the way the nervous system can generalize motor learning.

Behavioural evidence for a visual and proprioceptive control of head roll in hoverflies (Episyrphus balteatus).

The ability of hoverflies to control their head orientation with respect to their body contributes importantly to their agility and their autonomous navigation abilities. Many tasks performed by this insect during flight, especially while hovering, involve a head stabilization reflex. This reflex, which is mediated by multisensory channels, prevents the visual processing from being disturbed by motion blur and maintains a consistent perception of the visual environment. The so-called dorsal light response (DLR) is another head control reflex, which makes insects sensitive to the brightest part of the visual field. In this study, we experimentally validate and quantify the control loop driving the head roll with respect to the horizon in hoverflies. The new approach developed here consisted of using an upside-down horizon in a body roll paradigm. In this unusual configuration, tethered flying hoverflies surprisingly no longer use purely vision-based control for head stabilization. These results shed new light on the role of neck proprioceptor organs in head and body stabilization with respect to the horizon. Based on the responses obtained with male and female hoverflies, an improved model was then developed in which the output signals delivered by the neck proprioceptor organs are combined with the visual error in the estimated position of the body roll. An internal estimation of the body roll angle with respect to the horizon might explain the extremely accurate flight performances achieved by some hovering insects.

Getting back in the loop: Does autonomous driving duration affect driver's takeover performance?

The level 3 autonomous driving function allows the driver to perform non-driving-related tasks such as watching movies or reading while the system manages the driving task. However, when a difficult situation arises, the driver is requested to return to the loop of control. This switching from driver to passenger then back to driver may modify the driving paradigm, potentially causing an out-of-the-loop state. We tested the hypothesis of a linear (progressive) impact of various autonomous driving durations: the longer the level 3 autonomous function is used, the poorer the driver's takeover performance. Fifty-two participants were divided into 4 groups, each group being assigned a specific period of autonomous driving (5, 15, 45, or 60 min), followed by a takeover request with a time budget of 8.3 s. Takeover performance was assessed over two successive drives via reaction times and manual driving metrics (trajectories). The initial hypothesis (linearity) was not confirmed: there was a nonlinear relationship between autonomous driving duration and takeover performance, with one duration (15 min) appearing safer overall and mixed performance within groups. Repetition induced a major change in performance during the second drive, indicating rapid adaptation to the situation. The non-driving-related task appears critical in several respects (dynamics, content, driver interest) to proper use of level 3 automation. All this supports previous research prompting reservations about the prospect of car driving becoming like train travel.

Bayesian networks and information theory for audio-visual perception modeling.

Thanks to their different senses, human observers acquire multiple information coming from their environment. Complex cross-modal interactions occur during this perceptual process. This article proposes a framework to analyze and model these interactions through a rigorous and systematic data-driven process. This requires considering the general relationships between the physical events or factors involved in the process, not only in quantitative terms, but also in term of the influence of one factor on another. We use tools from information theory and probabilistic reasoning to derive relationships between the random variables of interest, where the central notion is that of conditional independence. Using mutual information analysis to guide the model elicitation process, a probabilistic causal model encoded as a Bayesian network is obtained. We exemplify the method by using data collected in an audio-visual localization task for human subjects, and we show that it yields a well-motivated model with good predictive ability. The model elicitation process offers new prospects for the investigation of the cognitive mechanisms of multisensory perception.

Adaptation of reactive and voluntary saccades: different patterns of adaptation revealed in the antisaccade task.

Sensorimotor adaptation restores and maintains the accuracy of goal-directed movements. It remains unclear whether these adaptive mechanisms modify actions by controlling peripheral premotor stages that send commands to the effectors and/or earlier processing stages involved in registration of target location. Here, we studied the effect of adaptation of saccadic eye movements, a well-established model of sensorimotor adaptation, in an antisaccade task. This task introduces a clear spatial dissociation between the actual target direction and the requested saccade direction because the correct movement direction is in the opposite direction from the target location. We used this requirement of a vector inversion to assess the level(s) of saccadic adaptation for two different types of adapted saccades. In two different experiments, we tested the transfer to antisaccades of the adaptation in one direction of reactive saccades to jumping targets and of scanning voluntary saccades within a target array. In the first experiment, we found that adaptation of reactive saccades transferred only to antisaccades in the adapted direction. In contrast, in the second experiment, adaptation of scanning voluntary saccades transferred to antisaccades in both the adapted and non-adapted directions. We conclude that adaptation of reactive saccades acts only downstream of the vector inversion required in the antisaccade task, whereas adaptation of voluntary saccades has a distributed influence, acting both upstream and downstream of vector inversion.

Influence of gaze elevation on estimating the possibility of passing under high obstacles during body tilt.

We investigated the influence of gaze elevation on judging the possibility of passing under high obstacles during pitch body tilts, while stationary, in absence of allocentric cues. Specifically, we aimed at studying the influence of egocentric references upon geocentric judgements. Seated subjects, orientated at various body orientations, were asked to perceptually estimate the possibility of passing under a projected horizontal line while keeping their gaze on a fixation target and imagining a horizontal body displacement. The results showed a global overestimation of the possibility of passing under the line, and confirmed the influence of body orientation reported by Bringoux et al. (Exp Brain Res 185(4):673-680, 2008). More strikingly, a linear influence of gaze elevation was found on perceptual estimates. Precisely, downward eye elevation yielded increased overestimations, and conversely upward gaze elevation yielded decreased overestimations. Furthermore, body and gaze orientation effects were independent and combined additively to yield a global egocentric influence with a weight of 45 and 54%, respectively. Overall, our data suggest that multiple egocentric references can jointly affect the estimated possibility of passing under high obstacles. These results are discussed in terms of "interpenetrability" between geocentric and egocentric reference frames and clearly demonstrate that gaze elevation is involved, as body orientation, in geocentric spatial localization.

Eye-hand coordination in rhythmical pointing.

The authors investigated the relation between hand kinematics and eye movements in 2 variants of a rhythmical Fitts's task in which eye movements were necessary or not necessary. P. M. Fitts's (1954) law held in both conditions with similar slope and marginal differences in hand-kinematic patterns and movement continuity. Movement continuity and eye-hand synchronization were more directly related to movement time than to task index of difficulty. When movement time was decreased to fewer than 350 ms, eye-hand synchronization switched from continuous monitoring to intermittent control. The 1:1 frequency ratio with stable pi/6 relative phase changed for 1:3 and 1:5 frequency ratios with less stable phase relations. The authors conclude that eye and hand movements in a rhythmical Fitts's task are dynamically synchronized to produce the best behavioral performance.

Detection and prediction of driver drowsiness using artificial neural network models.

Not just detecting but also predicting impairment of a car driver's operational state is a challenge. This study aims to determine whether the standard sources of information used to detect drowsiness can also be used to predict when a given drowsiness level will be reached. Moreover, we explore whether adding data such as driving time and participant information improves the accuracy of detection and prediction of drowsiness. Twenty-one participants drove a car simulator for 110min under conditions optimized to induce drowsiness. We measured physiological and behavioral indicators such as heart rate and variability, respiration rate, head and eyelid movements (blink duration, frequency and PERCLOS) and recorded driving behavior such as time-to-lane-crossing, speed, steering wheel angle, position on the lane. Different combinations of this information were tested against the real state of the driver, namely the ground truth, as defined from video recordings via the Trained Observer Rating. Two models using artificial neural networks were developed, one to detect the degree of drowsiness every minute, and the other to predict every minute the time required to reach a particular drowsiness level (moderately drowsy). The best performance in both detection and prediction is obtained with behavioral indicators and additional information. The model can detect the drowsiness level with a mean square error of 0.22 and can predict when a given drowsiness level will be reached with a mean square error of 4.18min. This study shows that, on a controlled and very monotonous environment conducive to drowsiness in a driving simulator, the dynamics of driver impairment can be predicted.

Coupling kinematics of memory and kinematics of movement: the conditions for a psychological relativity.

The organization of spatial and temporal information relative to memory-movement interactions was studied in a recall task. Stimuli consisted of four different configurations of eight dots, presented sequentially. In two configurations, the stimuli were spatiotemporal congruent, with (constant or variable) inter-stimulus distances corresponding to (constant or variable) inter-stimulus time intervals. In the other two configurations, the stimuli were spatiotemporal incongruent, with (constant or variable) inter-stimulus distances not corresponding to (variable or constant) inter-stimulus time intervals. After a learning phase consisting of 20 presentations of the target configuration, participants performed a series of pointing movements to reproduce both spatial and temporal characteristics of the stimulus 60 times in succession without ever re-examining the target configuration. Classically, in incongruent spatiotemporal conditions, the reproduction of, respectively, constant distances or constant time intervals are strongly disturbed by the simultaneous perception of variable inter-stimulus time intervals (Tau effect) or variable distances (Kappa effect). We assume that these spatiotemporal dependencies, which occur when the response relates to one dimension only, crucially depend on the non-integration of motoric aspects in memory.

Modelling gait processes as a combination of sensory-motor and cognitive controls in an attempt to describe accidents on the level in occupational situations.

In occupational situations, accidents referred to as accidents on the level (AoLs) occur most of the time when locomotion control fails. This control is determined by the interactions between the operator and the environment, the task and the used tools. Hence, AoLs prevention requires developing ways to optimise these interactions. More fundamentally, AoLs prevention requires understanding locomotion control in situations where this control is at sake, that is in situations involving one or more AoLs factors. The purpose of this article is to propose a comprehensive model for the control of locomotion in occupational environments. This model featuring the operator, the task and the working space should be an appropriate tool to understand AoLs in the scope of their prevention. Firstly, we describe what occupational AoLs are. In a second part, we present a review of the theoretical and experimental knowledge related to the locomotion system through the various means developed by the Central Nervous System to cope with perturbations of the environment and/or particular constraints from the task. Finally, we propose a simplified systemic model presenting the various levels of control (sensory-motor to cognitive levels) describing locomotion in occupational situations, and we suggest experiments likely to produce the appropriate data to construct the final comprehensive model.

Adapting artificial neural networks to a specific driver enhances detection and prediction of drowsiness.

Monitoring car drivers for drowsiness is crucial but challenging. The high inter-individual variability observed in measurements raises questions about the accuracy of the drowsiness detection process. In this study, we sought to enhance the performance of machine learning models (Artificial Neural Networks: ANNs) by training a model with a group of drivers and then adapting it to a new individual. Twenty-one participants drove a car simulator for 110 min in a monotonous environment. We measured physiological and behavioral indicators and recorded driving behavior. These measurements, in addition to driving time and personal information, served as the ANN inputs. Two ANN-based models were used, one to detect the level of drowsiness every minute, and the other to predict, every minute, how long it would take the driver to reach a specific drowsiness level (moderately drowsy). The ANNs were trained with 20 participants and subsequently adapted using the earliest part of the data recorded from a 21st participant. Then the adapted ANNs were tested with the remaining data from this 21st participant. The same procedure was run for all 21 participants. Varying amounts of data were used to adapt the ANNs, from 1 to 30 min, Model performance was enhanced for each participant. The overall drowsiness monitoring performance of the models was enhanced by roughly 40% for prediction and 80% for detection.

Oculomotor plasticity: are mechanisms of adaptation for reactive and voluntary saccades separate?

Saccadic eye movements are permanently controlled and their accuracy maintained by adaptive mechanisms that compensate for physiological or pathological perturbations. In contrast to the adaptation of reactive saccades (RS) which are automatically triggered by the sudden appearance of a single target, little is known about the adaptation of voluntary saccades which allow us to intentionally scan our environment in nearly all our daily activities. In this study, we addressed this issue in human subjects by determining the properties of adaptation of scanning voluntary saccades (SVS) and comparing these features to those of RS. We also tested the reciprocal transfers of adaptation between the two saccade types. Our results revealed that SVS and RS adaptations disclosed similar adaptation fields, time course and recovery levels, with only a slightly lower after-effect for SVS. Moreover, RS and SVS main sequences both remained unaffected after adaptation. Finally and quite unexpectedly, the pattern of adaptation transfers was asymmetrical, with a much stronger transfer from SVS to RS (79%) than in the reverse direction (22%). These data demonstrate that adaptations of RS and SVS share several behavioural properties but at the same time rely on partially distinct processes. Based on these findings, it is proposed that adaptations of RS and SVS may involve a neural network including both a common site and two separate sites specifically recruited for each saccade type.

Adaptive control: a review of the ability to acquire and maintain high sensorimotor performance.

When the relationship which relates us to the environment through vision, often named visual mapping, is durably modified, our behaviour is altered at sensory, motor and cognitive levels. The brain has the ability through the so-called adaptive control to progressively decrease the motor errors despite visual image alteration. Adaptive control in both the visuo-manual and the vestibulo-ocular systems is reviewed. A parallel is tentatively made between adaptation to ordinary optical alterations (resulting from the use of refraction-corrective glasses) in the visual mapping of these two systems and the adaptation to more complex visuo-motor relationships experienced by people working in virtual environments.

Coordination of upper and lower body during balance recovery following a support translation.

Strategies for recovery of posture were studied after lateral mechanical perturbations. 11 participants standing in tandem stance were unexpectedly submitted to lateral support translations with the eyes open or closed at two translation amplitudes. The trajectories of the center of mass of the upper and lower body and muscle activities allowed identification of three strategies, involving either the ankle or the hip only, or both. Hip use increased with vision and with amplitude of perturbation. Short-to-medium latency electromyographic activities were observed in leg and trunk muscles, and long-latency responses in the back leg muscles. Vision increased the activity of both leg and trunk muscles but did not influence the onset of the muscular responses. These data suggest a hierarchy in the selection of these different strategies: the hip is mobilized when the perturbation is more destabilizing but this strategy has a cost and needs specific sensory information supplied by vision.

Sensorimotor Reorganizations of Arm Kinematics and Postural Strategy for Functional Whole-Body Reaching Movements in Microgravity.

Understanding the impact of weightlessness on human behavior during the forthcoming long-term space missions is of critical importance, especially when considering the efficiency of goal-directed movements in these unusual environments. Several studies provided a large set of evidence that gravity is taken into account during the planning stage of arm reaching movements to optimally anticipate its consequence upon the moving limbs. However, less is known about sensorimotor changes required to face weightless environments when individuals have to perform fast and accurate goal-directed actions with whole-body displacement. We thus aimed at characterizing kinematic features of whole-body reaching movements in microgravity, involving high spatiotemporal constraints of execution, to question whether and how humans are able to maintain the performance of a functional behavior in the standards of normogravity execution. Seven participants were asked to reach as fast and as accurately as possible visual targets while standing during microgravity episodes in parabolic flight. Small and large targets were presented either close or far from the participants (requiring, in the latter case, additional whole-body displacement). Results reported that participants successfully performed the reaching task with general temporal features of movement (e.g., movement speed) close to land observations. However, our analyses also demonstrated substantial kinematic changes related to the temporal structure of focal movement and the postural strategy to successfully perform -constrained- whole-body reaching movements in microgravity. These immediate reorganizations are likely achieved by rapidly taking into account the absence of gravity in motor preparation and execution (presumably from cues about body limbs unweighting). Specifically, when compared to normogravity, the arm deceleration phase substantially increased. Furthermore, greater whole-body forward displacements due to smaller trunk flexions occurred when reaching far targets in microgravity. Remarkably, these changes of focal kinematics and postural strategy appear close to those previously reported when participants performed the same task underwater with neutral buoyancy applied to body limbs. Overall, these novel findings reveal that humans are able to maintain the performance of functional goal-directed whole-body actions in weightlessness by successfully managing spatiotemporal constraints of execution in this unusual environment.