Spatial arrangement of the whiskers of harbor seals (Phoca vitulina) compared to whisker arrangements of house mice (Mus musculus) and brown rats (Rattus norvegicus).

Whiskers (vibrissae) are important tactile sensors for most mammals. We introduce a novel approach to quantitatively compare 3D geometry of whisker arrays across species with different whisker numbers and arrangements, focusing on harbor seals (Phoca vitulina), house mice (Mus musculus) and Norway rats (Rattus norvegicus). Whiskers of all three species decrease in arclength and increase in curvature from caudal to rostral. They emerge from the face with elevation angles that vary linearly with dorsoventral position, and with curvature orientations that vary diagonally as linear combinations of dorsoventral and rostrocaudal positions. In seals, this diagonal varies linearly with horizontal emergence angles, and is orthogonal to the diagonal for rats and mice. This work provides the first evidence for common elements of whisker arrangements across species in different mammalian orders. Placing the equation-based whisker array on a CAD model of a seal head enables future mechanical studies of whisker-based sensing, including wake-tracking.

Continuous, multidimensional coding of 3D complex tactile stimuli by primary sensory neurons of the vibrissal system.

Across all sensory modalities, first-stage sensory neurons are an information bottleneck: they must convey all information available for an animal to perceive and act in its environment. Our understanding of coding properties of primary sensory neurons in the auditory and visual systems has been aided by the use of increasingly complex, naturalistic stimulus sets. By comparison, encoding properties of primary somatosensory afferents are poorly understood. Here, we use the rodent whisker system to examine how tactile information is represented in primary sensory neurons of the trigeminal ganglion (Vg). Vg neurons have long been thought to segregate into functional classes associated with separate streams of information processing. However, this view is based on Vg responses to restricted stimulus sets which potentially underreport the coding capabilities of these neurons. In contrast, the current study records Vg responses to complex three-dimensional (3D) stimulation while quantifying the complete 3D whisker shape and mechanics, thereby beginning to reveal their full representational capabilities. The results show that individual Vg neurons simultaneously represent multiple mechanical features of a stimulus, do not preferentially encode principal components of the stimuli, and represent continuous and tiled variations of all available mechanical information. These results directly contrast with proposed codes in which subpopulations of Vg neurons encode select stimulus features. Instead, individual Vg neurons likely overcome the information bottleneck by encoding large regions of a complex sensory space. This proposed tiled and multidimensional representation at the Vg directly constrains the computations performed by more central neurons of the vibrissotrigeminal pathway.

A dynamical model for generating synthetic data to quantify active tactile sensing behavior in the rat.

As it becomes possible to simulate increasingly complex neural networks, it becomes correspondingly important to model the sensory information that animals actively acquire: the biomechanics of sensory acquisition directly determines the sensory input and therefore neural processing. Here, we exploit the tractable mechanics of the well-studied rodent vibrissal ("whisker") system to present a model that can simulate the signals acquired by a full sensor array actively sampling the environment. Rodents actively "whisk" ∼60 vibrissae (whiskers) to obtain tactile information, and this system is therefore ideal to study closed-loop sensorimotor processing. The simulation framework presented here, WHISKiT Physics, incorporates realistic morphology of the rat whisker array to predict the time-varying mechanical signals generated at each whisker base during sensory acquisition. Single-whisker dynamics were optimized based on experimental data and then validated against free tip oscillations and dynamic responses to collisions. The model is then extrapolated to include all whiskers in the array, incorporating each whisker's individual geometry. Simulation examples in laboratory and natural environments demonstrate that WHISKiT Physics can predict input signals during various behaviors, currently impossible in the biological animal. In one exemplary use of the model, the results suggest that active whisking increases in-plane whisker bending compared to passive stimulation and that principal component analysis can reveal the relative contributions of whisker identity and mechanics at each whisker base to the vibrissotactile response. These results highlight how interactions between array morphology and individual whisker geometry and dynamics shape the signals that the brain must process.

Comparative morphology of the whiskers and faces of mice (Mus musculus) and rats (Rattus norvegicus).

Understanding neural function requires quantification of the sensory signals that an animal's brain evolved to interpret. These signals in turn depend on the morphology and mechanics of the animal's sensory structures. Although the house mouse (Mus musculus) is one of the most common model species used in neuroscience, the spatial arrangement of its facial sensors has not yet been quantified. To address this gap, the present study quantifies the facial morphology of the mouse, with a particular focus on the geometry of its vibrissae (whiskers). The study develops equations that establish relationships between the three-dimensional (3D) locations of whisker basepoints, whisker geometry (arclength, curvature) and the 3D angles at which the whiskers emerge from the face. Additionally, the positions of facial sensory organs are quantified relative to bregma-lambda. Comparisons with the Norway rat (Rattus norvegicus) indicate that when normalized for head size, the whiskers of these two species have similar spacing density. The rostral-caudal distances between facial landmarks of the rat are a factor of ∼2.0 greater than the mouse, while the scale of bilateral distances is larger and more variable. We interpret these data to suggest that the larger size of rats compared with mice is a derived (apomorphic) trait. As rodents are increasingly important models in behavioral neuroscience, the morphological model developed here will help researchers generate naturalistic, multimodal patterns of stimulation for neurophysiological experiments and allow the generation of synthetic datasets and simulations to close the loop between brain, body and environment.

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.

Constraints on the deformation of the vibrissa within the follicle.

Nearly all mammals have a vibrissal system specialized for tactile sensation, composed of whiskers growing from sensor-rich follicles in the skin. When a whisker deflects against an object, it deforms within the follicle and exerts forces on the mechanoreceptors inside. In addition, during active whisking behavior, muscle contractions around the follicle and increases in blood pressure in the ring sinus will affect the whisker deformation profile. To date, however, it is not yet possible to experimentally measure how the whisker deforms in an intact follicle or its effects on different groups of mechanoreceptors. The present study develops a novel model to predict vibrissal deformation within the follicle sinus complex. The model is based on experimental results from a previous ex vivo study on whisker deformation within the follicle, and on a new histological analysis of follicle tissue. It is then used to simulate whisker deformation within the follicle during passive touch and active whisking. Results suggest that the most likely whisker deformation profile is "S-shaped," crossing the midline of the follicle right below the ring sinus. Simulations of active whisking indicate that an increase in overall muscle stiffness, an increase in the ratio between deep and superficial intrinsic muscle stiffness, and an increase in sinus blood pressure will all enhance tactile sensitivity. Finally, we discuss how the deformation profiles might map to the responses of primary afferents of each mechanoreceptor type. The mechanical model presented in this study is an important first step in simulating mechanical interactions within whisker follicles.

Whisker Vibrations and the Activity of Trigeminal Primary Afferents in Response to Airflow.

Rodents are the most commonly studied model system in neuroscience, but surprisingly few studies investigate the natural sensory stimuli that rodent nervous systems evolved to interpret. Even fewer studies examine neural responses to these natural stimuli. Decades of research have investigated the rat vibrissal (whisker) system in the context of direct touch and tactile stimulation, but recent work has shown that rats also use their whiskers to help detect and localize airflow. The present study investigates the neural basis for this ability as dictated by the mechanical response of whiskers to airflow. Mechanical experiments show that a whisker's vibration magnitude depends on airspeed and the intrinsic shape of the whisker. Surprisingly, the direction of the whisker's vibration changes as a function of airflow speed: vibrations transition from parallel to perpendicular with respect to the airflow as airspeed increases. Recordings from primary sensory trigeminal ganglion neurons show that these neurons exhibit responses consistent with those that would be predicted from direct touch. Trigeminal neuron firing rate increases with airspeed, is modulated by the orientation of the whisker relative to the airflow, and is influenced by the whisker's resonant frequencies. We develop a simple model to describe how a population of neurons could leverage mechanical relationships to decode both airspeed and direction. These results open new avenues for studying vibrissotactile regions of the brain in the context of evolutionarily important airflow-sensing behaviors and olfactory search. Although this study used only female rats, all results are expected to generalize to male rats.SIGNIFICANCE STATEMENT The rodent vibrissal (whisker) system has been studied for decades in the context of direct tactile sensation, but recent work has indicated that rats also use whiskers to help localize airflow. Neural circuits in somatosensory regions of the rodent brain thus likely evolved in part to process airflow information. This study investigates the whiskers' mechanical response to airflow and the associated neural response. Airspeed affects the magnitude of whisker vibration and the response magnitude of whisker-sensitive primary sensory neurons in the trigeminal ganglion. Surprisingly, the direction of vibration and the associated directionally dependent neural response changes with airspeed. These findings suggest a population code for airflow speed and direction and open new avenues for studying vibrissotactile regions of the brain.

Defining "active sensing" through an analysis of sensing energetics: homeoactive and alloactive sensing.

The term "active sensing" has been defined in multiple ways. Most strictly, the term refers to sensing that uses self-generated energy to sample the environment (e.g., echolocation). More broadly, the definition includes all sensing that occurs when the sensor is moving (e.g., tactile stimuli obtained by an immobile versus moving fingertip) and, broader still, includes all sensing guided by attention or intent (e.g., purposeful eye movements). The present work offers a framework to help disambiguate aspects of the "active sensing" terminology and reveals properties of tactile sensing unique among all modalities. The framework begins with the well-described "sensorimotor loop," which expresses the perceptual process as a cycle involving four subsystems: environment, sensor, nervous system, and actuator. Using system dynamics, we examine how information flows through the loop. This "sensory-energetic loop" reveals two distinct sensing mechanisms that subdivide active sensing into homeoactive and alloactive sensing. In homeoactive sensing, the animal can change the state of the environment, while in alloactive sensing the animal can alter only the sensor's configurational parameters and thus the mapping between input and output. Given these new definitions, examination of the sensory-energetic loop helps identify two unique characteristics of tactile sensing: 1) in tactile systems, alloactive and homeoactive sensing merge to a mutually controlled sensing mechanism, and 2) tactile sensing may require fundamentally different predictions to anticipate reafferent input. We expect this framework may help resolve ambiguities in the active sensing community and form a basis for future theoretical and experimental work regarding alloactive and homeoactive sensing.

Quantification of vibrissal mechanical properties across the rat mystacial pad.

Recent work has quantified the geometric parameters of individual rat vibrissae (whiskers) and developed equations that describe how these parameters vary as a function of row and column position across the array. This characterization included a detailed quantification of whisker base diameter and arc length as well as the geometry of the whisker medulla. The present study now uses these equations for whisker geometry to quantify several properties of the whisker that govern its mechanical behavior. We first show that the average density of a whisker is lower in its proximal region than in its distal region. This density variation appears to be largely attributable to the presence of the whisker cuticle rather than the medulla. The density variation has very little effect on the center of mass of the whisker. We next show that the presence of the medulla decreases the deflection of the whisker under its own weight and also decreases its mass moment of inertia while sacrificing <1% stiffness at the whisker base compared with a solid whisker. Finally, we quantify two dimensionless parameters across the array. First, the deflection-to-length ratio decreases from caudal to rostral: caudal whiskers are longer but deflect more under their own weight. Second, the nondimensionalized radius of gyration is approximately constant across the array, which may simplify control of whisking by the intrinsic muscles. We anticipate that future work will exploit the mechanical properties computed in the present study to improve simulations of the mechanosensory signals associated with vibrissotactile exploratory behavior. NEW & NOTEWORTHY The mechanical signals transmitted by a whisker depend critically on its geometry. We used measurements of whisker geometry and mass to quantify the center of mass, mass moment of inertia, radius of gyration, and deflection under gravity of the whisker. We describe how variations in these quantities across the array could enhance sensing behaviors while reducing energy costs and simplifying whisking control. Most importantly, we provide derivations for these quantities for use in future simulation work.

Variations in vibrissal geometry across the rat mystacial pad: base diameter, medulla, and taper.

Many rodents tactually sense the world through active motions of their vibrissae (whiskers), which are regularly arranged in rows and columns (arcs) on the face. The present study quantifies several geometric parameters of rat whiskers that determine the tactile information acquired. Findings include the following. 1) A meta-analysis of seven studies shows that whisker base diameter varies with arc length with a surprisingly strong dependence on the whisker's row position within the array. 2) The length of the whisker medulla varies linearly with whisker length, and the medulla's base diameter varies linearly with whisker base diameter. 3) Two parameters are required to characterize whisker "taper": radius ratio (base radius divided by tip radius) and radius slope (the difference between base and tip radius, divided by arc length). A meta-analysis of five studies shows that radius ratio exhibits large variability due to variations in tip radius, while radius slope varies systematically across the array. 4) Within the resolution of the present study, radius slope does not differ between the proximal and distal segments of the whisker, where "proximal" is defined by the presence of the medulla. 5) Radius slope of the medulla is offset by a constant value from radius slope of the proximal portion of the whisker. We conclude with equations for all geometric parameters as functions of row and column position.NEW & NOTEWORTHY Rats tactually explore their world by brushing and tapping their whiskers against objects. Each whisker's geometry will have a large influence on its mechanics and thus on the tactile signals the rat obtains. We performed a meta-analysis of seven studies to generate equations that describe systematic variations in whisker geometry across the rat's face. We also quantified the geometry of the whisker medulla. A database provides access to geometric parameters of over 500 rat whiskers.

Evidence for Functional Groupings of Vibrissae across the Rodent Mystacial Pad.

During natural exploration, rats exhibit two particularly conspicuous vibrissal-mediated behaviors: they follow along walls, and "dab" their snouts on the ground at frequencies related to the whisking cycle. In general, the walls and ground may be located at any distance from, and at any orientation relative to, the rat's head, which raises the question of how the rat might determine the position and orientation of these surfaces. Previous studies have compellingly demonstrated that rats can accurately determine the horizontal angle at which a vibrissa first touches an object, and we therefore asked whether this parameter could provide the rat with information about the pitch, distance, and yaw of a surface relative to its head. We used a three-dimensional model of the whisker array to construct mappings between the horizontal angle of contact of each vibrissa and every possible (pitch, distance, and yaw) configuration of the head relative to a flat surface. The mappings revealed striking differences in the patterns of contact for vibrissae in different regions of the array. The exterior (A, D, E) rows provide information about the relative pitch of the surface regardless of distance. The interior (B, C) rows provide distance cues regardless of head pitch. Yaw is linearly correlated with the difference between the number of right and left whiskers touching the surface. Compared to the long reaches that whiskers can make to the side and below the rat, the reachable distance in front of the rat's nose is relatively small. We confirmed key predictions of these functional groupings in a behavioral experiment that monitored the contact patterns that the vibrissae made with a flat vertical surface. These results suggest that vibrissae in different regions of the array are not interchangeable sensors, but rather functionally grouped to acquire particular types of information about the environment.

Mechanical responses of rat vibrissae to airflow.

The survival of many animals depends in part on their ability to sense the flow of the surrounding fluid medium. To date, however, little is known about how terrestrial mammals sense airflow direction or speed. The present work analyzes the mechanical response of isolated rat macrovibrissae (whiskers) to airflow to assess their viability as flow sensors. Results show that the whisker bends primarily in the direction of airflow and vibrates around a new average position at frequencies related to its resonant modes. The bending direction is not affected by airflow speed or by geometric properties of the whisker. In contrast, the bending magnitude increases strongly with airflow speed and with the ratio of the whisker's arc length to base diameter. To a much smaller degree, the bending magnitude also varies with the orientation of the whisker's intrinsic curvature relative to the direction of airflow. These results are used to predict the mechanical responses of vibrissae to airflow across the entire array, and to show that the rat could actively adjust the airflow data that the vibrissae acquire by changing the orientation of its whiskers. We suggest that, like the whiskers of pinnipeds, the macrovibrissae of terrestrial mammals are multimodal sensors - able to sense both airflow and touch - and that they may play a particularly important role in anemotaxis.

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.

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.

Probability distributions of whisker-surface contact: quantifying elements of the rat vibrissotactile natural scene.

Analysis of natural scene statistics has been a powerful approach for understanding neural coding in the auditory and visual systems. In the field of somatosensation, it has been more challenging to quantify the natural tactile scene, in part because somatosensory signals are so tightly linked to the animal's movements. The present work takes a step towards quantifying the natural tactile scene for the rat vibrissal system by simulating rat whisking motions to systematically investigate the probabilities of whisker-object contact in naturalistic environments. The simulations permit an exhaustive search through the complete space of possible contact patterns, thereby allowing for the characterization of the patterns that would most likely occur during long sequences of natural exploratory behavior. We specifically quantified the probabilities of 'concomitant contact', that is, given that a particular whisker makes contact with a surface during a whisk, what is the probability that each of the other whiskers will also make contact with the surface during that whisk? Probabilities of concomitant contact were quantified in simulations that assumed increasingly naturalistic conditions: first, the space of all possible head poses; second, the space of behaviorally preferred head poses as measured experimentally; and third, common head poses in environments such as cages and burrows. As environments became more naturalistic, the probability distributions shifted from exhibiting a 'row-wise' structure to a more diagonal structure. Results also reveal that the rat appears to use motor strategies (e.g. head pitches) that generate contact patterns that are particularly well suited to extract information in the presence of uncertainty.

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.

Correction: Quantifying the three-dimensional facial morphology of the laboratory rat with a focus on the vibrissae.

[This corrects the article DOI: 10.1371/journal.pone.0194981.].

Spatial arrangement of the whiskers of harbor seals ( Phoca vitulina ) compared to whisker arrangements of house mice ( Mus musculus ) and brown rats ( Rattus norvegicus ).

Whiskers (vibrissae) are important tactile sensors for most mammals. We introduce a novel approach to quantitatively compare 3D geometry of whisker arrays across species with different whisker numbers and arrangements, focusing on harbor seals ( Phoca vitulina ), house mice ( Mus musculus ) and Norway rats ( Rattus norvegicus ). Whiskers of all three species decrease in arclength and increase in curvature from caudal to rostral. They emerge from the face with elevation angles that vary linearly with dorsoventral position, and with curvature orientations that vary diagonally as linear combinations of dorsoventral and rostrocaudal positions. In seals, this diagonal varies linearly with horizontal emergence angles, and is orthogonal to the diagonal for rats and mice. This work provides the first evidence for common elements of whisker arrangements across species in different mammalian orders. Placing the equation-based whisker array on a CAD model of a seal head enables future mechanical studies of whisker-based sensing, including wake-tracking. SUMMARY STATEMENT: We quantify the three-dimensional positions and orientations of the whiskers across the face of the harbor seal, and compare this geometry with the whisker arrays of rats and mice.

On the intrinsic curvature of animal whiskers.

Facial vibrissae (whiskers) are thin, tapered, flexible, hair-like structures that are an important source of tactile sensory information for many species of mammals. In contrast to insect antennae, whiskers have no sensors along their lengths. Instead, when a whisker touches an object, the resulting deformation is transmitted to mechanoreceptors in a follicle at the whisker base. Previous work has shown that the mechanical signals transmitted along the whisker will depend strongly on the whisker's geometric parameters, specifically on its taper (how diameter varies with arc length) and on the way in which the whisker curves, often called "intrinsic curvature." Although previous studies have largely agreed on how to define taper, multiple methods have been used to quantify intrinsic curvature. The present work compares and contrasts different mathematical approaches towards quantifying this important parameter. We begin by reviewing and clarifying the definition of "intrinsic curvature," and then show results of fitting whisker shapes with several different functions, including polynomial, fractional exponent, elliptical, and Cesàro. Comparisons are performed across ten species of whiskered animals, ranging from rodents to pinnipeds. We conclude with a discussion of the advantages and disadvantages of using the various models for different modeling situations. The fractional exponent model offers an approach towards developing a species-specific parameter to characterize whisker shapes within a species. Constructing models of how the whisker curves is important for the creation of mechanical models of tactile sensory acquisition behaviors, for studies of comparative evolution, morphology, and anatomy, and for designing artificial systems that can begin to emulate the whisker-based tactile sensing of animals.

Mechanical signals at the base of a rat vibrissa: the effect of intrinsic vibrissa curvature and implications for tactile exploration.

Rats actively tap and sweep their large mystacial vibrissae (whiskers) against objects to tactually explore their surroundings. When a vibrissa makes contact with an object, it bends, and this bending generates forces and bending moments at the vibrissa base. Researchers have only recently begun to quantify these mechanical variables. The present study quantifies the forces and bending moments at the vibrissa base with a quasi-static model of vibrissa deflection. The model was validated with experiments on real vibrissae. Initial simulations demonstrated that almost all vibrissa-object collisions during natural behavior will occur with the concave side of the vibrissa facing the object, and we therefore paid particular attention to the role of the vibrissa's intrinsic curvature in shaping the forces at the base. Both simulations and experiments showed that vibrissae with larger intrinsic curvatures will generate larger axial forces. Simulations also demonstrated that the range of forces and moments at the vibrissal base vary over approximately three orders of magnitude, depending on the location along the vibrissa at which object contact is made. Both simulations and experiments demonstrated that collisions in which the concave side of the vibrissa faces the object generate longer-duration contacts and larger net forces than collisions with the convex side. These results suggest that the orientation of the vibrissa's intrinsic curvature on the mystacial pad may increase forces during object contact and provide increased sensitivity to detailed surface features.