Demonstration of three-dimensional contact point determination and contour reconstruction during active whisking behavior of an awake rat.

The rodent vibrissal (whisker) system has been studied for decades as a model of active touch sensing. There are no sensors along the length of a whisker; all sensing occurs at the whisker base. Therefore, a large open question in many neuroscience studies is how an animal could estimate the three-dimensional (3D) location at which a whisker makes contact with an object. In the present work we simulated the shape of a real rat whisker to demonstrate the existence of several unique mappings from triplets of mechanical signals at the whisker base to the three-dimensional whisker-object contact point. We then used high speed video to record whisker deflections as an awake rat whisked against a peg, and used the mechanics resulting from those deflections to extract the contact points along the peg surface. These results demonstrate that measurement of specific mechanical triplets at the base of a biological whisker can enable 3D contact point determination during natural whisking behavior. The approach is viable even though the biological whisker has non-ideal, non-planar curvature, and even given the rat's real-world choices of whisking parameters. Visual intuition for the quality of the approach is provided in a video that shows the contour of the peg gradually emerging during active whisking behavior.

Tactile Sensing with Whiskers of Various Shapes: Determining the Three-Dimensional Location of Object Contact Based on Mechanical Signals at the Whisker Base.

Almost all mammals use their mystacial vibrissae (whiskers) as important tactile sensors. There are no sensors along the length of a whisker: all sensing is performed by mechanoreceptors at the whisker base. To use artificial whiskers as a sensing tool in robotics, it is essential to be able to determine the three-dimensional (3D) location at which a whisker has made contact with an object. With the assumption of quasistatic, frictionless, single-point contact, previous work demonstrated that the 3D contact point can be uniquely determined if all six components of force and moment are measured at the whisker base, but these measurements require a six-axis load cell. Here, we perform simulations to investigate the extent to which each of the 20 possible "triplet" combinations of the six mechanical signals at the whisker base uniquely determine 3D contact point location. We perform this analysis for four different whisker profiles (shapes): tapered with and without intrinsic curvature, and cylindrical with and without intrinsic curvature. We show that whisker profile has a strong influence on the particular triplet(s) of signals that uniquely map to the 3D contact point. The triplet of bending moment, bending moment direction, and axial force produces unique mappings for tapered whiskers. Four different mappings are unique for a cylindrical whisker without intrinsic curvature, but only when large deflections are excluded. These results inform the neuroscience of vibrissotactile sensing and represent an important step toward the development of artificial whiskers for robotic applications.

Decoupling kinematics and mechanics reveals coding properties of trigeminal ganglion neurons in the rat vibrissal system.

Tactile information available to the rat vibrissal system begins as external forces that cause whisker deformations, which in turn excite mechanoreceptors in the follicle. Despite the fundamental mechanical origin of tactile information, primary sensory neurons in the trigeminal ganglion (Vg) have often been described as encoding the kinematics (geometry) of object contact. Here we aimed to determine the extent to which Vg neurons encode the kinematics vs. mechanics of contact. We used models of whisker bending to quantify mechanical signals (forces and moments) at the whisker base while simultaneously monitoring whisker kinematics and recording single Vg units in both anesthetized rats and awake, body restrained rats. We employed a novel manual stimulation technique to deflect whiskers in a way that decouples kinematics from mechanics, and used Generalized Linear Models (GLMs) to show that Vg neurons more directly encode mechanical signals when the whisker is deflected in this decoupled stimulus space.

Simulations of a Vibrissa Slipping along a Straight Edge and an Analysis of Frictional Effects during Whisking.

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.

Tactile signals transmitted by the vibrissa during active whisking behavior.

The rodent vibrissal-trigeminal system is one of the most widely used models for the study of somatosensation and tactile perception, but to date the field has been unable to quantify the complete set of mechanical input signals generated during natural whisking behavior. In this report we show that during whisking behavior of awake rats (Rattus norvegicus), the whisker will often bend out of its plane of rotation, generating sizeable mechanical (tactile) signals out of the plane. We then develop a model of whisker bending that allows us to compute the three-dimensional tactile signals at the vibrissal base during active whisking behavior. Considerable information can be lost if whisking motions are considered only in two dimensions, and we offer some suggestions for experimentalists concerned with monitoring the direction of bending. These data represent the first quantification of the physical signals transmitted to the mechanoreceptors in the follicle during active whisking behavior.

The search space of the rat during whisking behavior.

Rodents move their vibrissae rhythmically to tactually explore their surroundings. We used a three-dimensional model of the vibrissal array to quantify the rat's 'search space' during whisking. Search space was quantified either as the volume encompassed by the array or as the surface formed by the vibrissal tips. At rest, the average position of the vibrissal tips lies near the rat's mouth, and the tips are all approximately equidistant from the midpoint between the rat's eyes, suggesting spatial registration with the visual system. The intrinsic curvature of the vibrissae greatly increases the volume encompassed by the array, and during a protraction, roll and elevation changes have strong effects on the trajectories of the vibrissal tips. The size of the rat's search space--as measured either by the volume of the array or by the surface area formed by the vibrissal tips--was surprisingly unaffected by protraction angle. In contrast, search space was strongly correlated with the 'spread' of the array, defined as the angle between rostral and caudal-most whiskers. We draw two conclusions: first, that with some caveats, spread can be used as a proxy for changes in search space, and second, in order to change its sensing resolution, the rat must differentially control rostral and caudal vibrissae. Finally, we show that behavioral data can be incorporated into the three-dimensional model to visualize changes in vibrissal search space and sensing resolution during natural exploratory whisking.

Modeling forces and moments at the base of a rat vibrissa during noncontact whisking and whisking against an object.

During exploratory behavior, rats brush and tap their whiskers against objects, and the mechanical signals so generated constitute the primary sensory variables upon which these animals base their vibrissotactile perception of the world. To date, however, we lack a general dynamic model of the vibrissa that includes the effects of inertia, damping, and collisions. We simulated vibrissal dynamics to compute the time-varying forces and bending moment at the vibrissa base during both noncontact (free-air) whisking and whisking against an object (collision). Results show the following: (1) during noncontact whisking, mechanical signals contain components at both the whisking frequency and also twice the whisking frequency (the latter could code whisking speed); (2) when rats whisk rhythmically against an object, the intrinsic dynamics of the vibrissa can be as large as many of the mechanical effects of the collision, however, the axial force could still generate responses that reliably indicate collision based on thresholding; and (3) whisking velocity will have only a small effect on the transient response generated during a whisker-object collision. Instead, the transient response will depend in large part on how the rat chooses to decelerate its vibrissae after the collision. The model allows experimentalists to estimate error bounds on quasi-static descriptions of vibrissal shape, and its predictions can be used to bound realistic expectations from neurons that code vibrissal sensing. We discuss the implications of these results under the assumption that primary sensory neurons of the trigeminal ganglion are sensitive to various combinations of mechanical signals.