Whisker Contact Detection of Rodents Based on Slow and Fast Mechanical Inputs.
ABSTRACT: Rodents use their whiskers to locate nearby objects with an extreme precision. To perform such tasks, they need to detect whisker/object contacts with a high temporal accuracy. This contact detection is conveyed by classes of mechanoreceptors whose neural activity is sensitive to either slow or fast time varying mechanical stresses acting at the base of the whiskers. We developed a biomimetic approach to separate and characterize slow quasi-static and fast vibrational stress signals acting on a whisker base in realistic exploratory phases, using experiments on both real and artificial whiskers. Both slow and fast mechanical inputs are successfully captured using a mechanical model of the whisker. We present and discuss consequences of the whisking process in purely mechanical terms and hypothesize that free whisking in air sets a mechanical threshold for contact detection. The time resolution and robustness of the contact detection strategies based on either slow or fast stress signals are determined. Contact detection based on the vibrational signal is faster and more robust to exploratory conditions than the slow quasi-static component, although both slow/fast components allow localizing the object.
Project description:Rats discriminate surface textures using their whiskers (vibrissae), but how whiskers extract texture information, and how this information is encoded by the brain, are not known. In the resonance model, whisker motion across different textures excites mechanical resonance in distinct subsets of whiskers, due to variation across whiskers in resonance frequency, which varies with whisker length. Texture information is therefore encoded by the spatial pattern of activated whiskers. In the competing kinetic signature model, different textures excite resonance equally across whiskers, and instead, texture is encoded by characteristic, nonuniform temporal patterns of whisker motion. We tested these models by measuring whisker motion in awake, behaving rats whisking in air and onto sandpaper surfaces. Resonant motion was prominent during whisking in air, with fundamental frequencies ranging from approximately 35 Hz for the long Delta whisker to approximately 110 Hz for the shorter D3 whisker. Resonant vibrations also occurred while whisking against textures, but the amplitude of resonance within single whiskers was independent of texture, contradicting the resonance model. Rather, whiskers resonated transiently during discrete, high-velocity, and high-acceleration slip-stick events, which occurred prominently during whisking on surfaces. The rate and magnitude of slip-stick events varied systematically with texture. These results suggest that texture is encoded not by differential resonant motion across whiskers, but by the magnitude and temporal pattern of slip-stick motion. These findings predict a temporal code for texture in neural spike trains.
Project description:Whisking mediated touch is an active sense whereby whisker movements are modulated by sensory input and behavioral context. Here we studied the effects of touching an object on whisking in head-fixed rats. Simultaneous movements of whiskers C1, C2, and D1 were tracked bilaterally and their movements compared. During free-air whisking, whisker protractions were typically characterized by a single acceleration-deceleration event, whisking amplitude and velocity were correlated, and whisk duration correlated with neither amplitude nor velocity. Upon contact with an object, a second acceleration-deceleration event occurred in about 25% of whisk cycles, involving both contacting (C2) and non-contacting (C1, D1) whiskers ipsilateral to the object. In these cases, the rostral whisker (C2) remained in contact with the object throughout the double-peak phase, which effectively prolonged the duration of C2 contact. These "touch-induced pumps" (TIPs) were detected, on average, 17.9 ms after contact. On a slower time scale, starting at the cycle following first touch, contralateral amplitude increased while ipsilateral amplitude decreased. Our results demonstrate that sensory-induced motor modulations occur at various timescales, and directly affect object palpation.
Project description:All small mammals have prominent facial whiskers that they employ as tactile sensors to guide navigation and foraging in complex habitats. Nocturnal, arboreal mammals tend to have the longest and most densely packed whiskers, and semi-aquatic mammals have the most sensitive. Here we present evidence to indicate that many small mammals use their whiskers to tactually guide safe foot positioning. Specifically, in 11, small, non-flying mammal species, we demonstrate that forepaw placement always falls within the ground contact zone of the whisker field and that forepaw width is always smaller than whisker span. We also demonstrate commonalities of whisker scanning movements (whisking) and elements of active control, associated with increasing contact with objects of interest, across multiple small mammal species that have previously only been shown in common laboratory animals. Overall, we propose that guiding locomotion, alongside environment exploration, is a common function of whisker touch sensing in small, quadrupedal mammals.
Project description:Haptic perception synthesizes touch with proprioception, the sense of body position. Humans and mice alike experience rich active touch of the face. Because most facial muscles lack proprioceptor endings, the sensory basis of facial proprioception remains unsolved. Facial proprioception may instead rely on mechanoreceptors that encode both touch and self-motion. In rodents, whisker mechanoreceptors provide a signal that informs the brain about whisker position. Whisking involves coordinated orofacial movements, so mechanoreceptors innervating facial regions other than whiskers could also provide information about whisking. To define all sources of sensory information about whisking available to the brain, we recorded spikes from mechanoreceptors innervating diverse parts of the face. Whisker motion was encoded best by whisker mechanoreceptors, but also by those innervating whisker pad hairy skin and supraorbital vibrissae. Redundant self-motion responses may provide the brain with a stable proprioceptive signal despite mechanical perturbations during active touch.
Project description:Rats sweep their facial whiskers back and forth to generate tactile sensory information through contact with environmental structure. The neural processes operating on the signals arising from these whisker contacts are widely studied as a model of sensing in general, even though detailed knowledge of the natural circumstances under which such signals are generated is lacking. We used digital video tracking and wireless recording of mystacial electromyogram signals to assess the effects of whisker-object contact on whisking in freely moving animals exploring simple environments. Our results show that contact leads to reduced protraction (forward whisker motion) on the side of the animal ipsilateral to an obstruction and increased protraction on the contralateral side. Reduced ipsilateral protraction occurs rapidly and in the same whisk cycle as the initial contact. We conclude that whisker movements are actively controlled so as to increase the likelihood of environmental contacts while constraining such interactions to involve a gentle touch. That whisking pattern generation is under strong feedback control has important implications for understanding the nature of the signals reaching upstream neural processes.
Project description:The rat vibrissal system is an important model for the study of somatosensation, but the small size and rapid speed of the vibrissae have precluded measuring precise vibrissal-object contact sequences during behavior. We used a laser light sheet to quantify, with 1 ms resolution, the spatiotemporal structure of whisker-surface contact as five naïve rats freely explored a flat, vertical glass wall. Consistent with previous work, we show that the whisk cycle cannot be uniquely defined because different whiskers often move asynchronously, but that quasi-periodic (~8 Hz) variations in head velocity represent a distinct temporal feature on which to lock analysis. Around times of minimum head velocity, whiskers protract to make contact with the surface, and then sustain contact with the surface for extended durations (~25-60 ms) before detaching. This behavior results in discrete temporal windows in which large numbers of whiskers are in contact with the surface. These "sustained collective contact intervals" (SCCIs) were observed on 100% of whisks for all five rats. The overall spatiotemporal structure of the SCCIs can be qualitatively predicted based on information about head pose and the average whisk cycle. In contrast, precise sequences of whisker-surface contact depend on detailed head and whisker kinematics. Sequences of vibrissal contact were highly variable, equally likely to propagate in all directions across the array. Somewhat more structure was found when sequences of contacts were examined on a row-wise basis. In striking contrast to the high variability associated with contact sequences, a consistent feature of each SCCI was that the contact locations of the whiskers on the glass converged and moved more slowly on the sheet. Together, these findings lead us to propose that the rat uses a strategy of "windowed sampling" to extract an object's spatial features: specifically, the rat spatially integrates quasi-static mechanical signals across whiskers during the period of sustained contact, resembling an "enclosing" haptic procedure.
Project description:Animals actively regulate the position and movement of their sensory systems to boost the quality and quantity of the sensory information they obtain. The rat vibrissal system is recognized to be an important model system in which to investigate such "active sensing" capabilities. The current study used high-speed video analysis to investigate whisker movements in untrained, freely moving rats encountering unexpected, vertical surfaces. A prominent feature of rat vibrissal movement is the repeated posterior-anterior sweep of the whiskers in which the macrovibrissae are seen to move largely in synchrony. Here we show that a second significant component of whisking behavior is the size of the arc, or "spread," between the whiskers. Observed spread is shown to vary over the whisk cycle and to substantially decrease during exploration of an unexpected surface. We further show that the timing of whisker movements is affected by surface contact such that 1) the whiskers rapidly cease forward protraction following an initial, unexpected contact, and may do so even more rapidly following contact with the same surface in the subsequent whisk cycle, and 2) retraction velocity is reduced following this latter contact, leading to longer second-contact durations. This evidence is taken to support two hypotheses: 1) that the relative velocities of different whiskers may be actively controlled by the rat and 2) that control of whisker velocity and timing may serve to increase the number and duration of whisker-surface contacts while ensuring that such contacts are made with a light touch.
Project description:During tactile exploration, rats sweep their whiskers against objects in a motion called whisking. Here, we investigate how a whisker slips along an object's edge and how friction affects the resulting tactile signals. First, a frictionless model is developed to simulate whisker slip along a straight edge and compared with a previous model that incorporates friction but cannot simulate slip. Results of both models are compared to behavioral data obtained as a rat whisked against a smooth, stainless steel peg. As expected, the frictionless model predicts larger magnitudes of vertical slip than observed experimentally. The frictionless model also predicts forces and moments at the whisker base that are smaller and have a different direction than those predicted by the model with friction. Estimates for the friction coefficient yielded values near 0.48 (whisker/stainless steel). The present work provides the first assessments of the effects of friction on the mechanical signals received by the follicle during active whisking. It also demonstrates a proof-of-principle approach for reducing whisker tracking requirements during experiments and demonstrates the feasibility of simulating a full array of vibrissae whisking against a peg.
Project description:Rats use their whiskers as tactile sensors to sense their environment. Active whisking, moving whiskers back and forth continuously, is one of prominent features observed in rodents. They can discriminate different textures or extract features of a nearby object such as size, shape and distance through active whisking. There have been studies to localize objects with artificial whiskers inspired by rat whiskers. The linear whisker model based on beam theory has been used to estimate the radial distance, that is, the distance between the base of the whisker and a target object. In this paper, we investigate deflection angle measurements instead of forces or moments, based on a linear tapered whisker model to see the role of tapered whiskers found in real animals. We analyze how accurately this model estimates the radial distance, and quantify the estimation errors and noise sensitivity. We also compare the linear model simulation and nonlinear numerical solutions. It is shown that the radial distance can be estimated using deflection angles at two different positions on the tapered whisker. We argue that the tapered whisker has an advantage of estimating the radial distance better, as compared to an untapered whisker, and active sensing allows that estimation without the whisker's material property and thickness or the moment at base. In addition, we investigate the potential of passive sensing for tactile localization.
Project description:We have developed software for fully automated tracking of vibrissae (whiskers) in high-speed videos (>500 Hz) of head-fixed, behaving rodents trimmed to a single row of whiskers. Performance was assessed against a manually curated dataset consisting of 1.32 million video frames comprising 4.5 million whisker traces. The current implementation detects whiskers with a recall of 99.998% and identifies individual whiskers with 99.997% accuracy. The average processing rate for these images was 8 Mpx/s/cpu (2.6 GHz Intel Core2, 2 GB RAM). This translates to 35 processed frames per second for a 640 px×352 px video of 4 whiskers. The speed and accuracy achieved enables quantitative behavioral studies where the analysis of millions of video frames is required. We used the software to analyze the evolving whisking strategies as mice learned a whisker-based detection task over the course of 6 days (8148 trials, 25 million frames) and measure the forces at the sensory follicle that most underlie haptic perception.