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One of the largest ecological transitions in carnivoran evolution was the shift from terrestrial to aquatic lifestyles, which has driven morphological diversity in skulls and other skeletal structures. In this paper, we investigate the association between those lifestyles and whisker morphology. However, comparing whisker morphology over a range of species is challenging since the number of whiskers and their positions on the mystacial pads vary between species. Also, each whisker will be at a different stage of growth and may have incurred damage due to wear and tear. Identifying a way to easily capture whisker morphology in a small number of whisker samples would be beneficial. Here, we describe individual and species variation in whisker morphology from two-dimensional scans in red fox, European otter and grey seal. A comparison of long, caudal whiskers shows inter-species differences most clearly. We go on to describe global whisker shape in 24 species of carnivorans, using linear approximations of curvature and taper, as well as traditional morphometric methods. We also qualitatively examine surface texture, or the presence of scales, using scanning electron micrographs. We show that gross whisker shape is highly conserved, with whisker curvature and taper obeying simple linear relationships with length. However, measures of whisker base radius, length, and maybe even curvature, can vary between species and substrate preferences. Specifically, the aquatic species in our sample have thicker, shorter whiskers that are smoother, with less scales present than those of terrestrial species. We suggest that these thicker whiskers may be stiffer and able to maintain their shape and position during underwater sensing, but being stiffer may also increase wear., (© 2023 The Authors. Journal of Morphology published by Wiley Periodicals LLC.)

Background: Studying natural, complex behaviours over a range of different species provides insights into the evolution of the brain and behaviour. Whisker movements reveal complex behaviours; however, there does not yet exist a protocol that is able to capture whisker movements and behaviours in a range of different species., New Method: We develop a new protocol and make recommendations for measuring comparative whisker movements and behaviours. Using two set-ups - an enclosure camera set-up and a high-speed video set-up - we capture and measure the whisker movements of sixteen different captive mammal species from four different animal collections., Results: We demonstrate the ability to describe whisker movements and behaviours across a wide range of mammalian species. We describe whisker movements in European hedgehog, Cape porcupine, domestic rabbit, domestic ferret, weasel, European otter and red fox for the first time. We observe whisker movements in all the species we tested, although movement, positions and behaviours vary in a species-specific way., Comparison With Existing Method(s): The high-speed video set-up is based on the protocols of previous studies. The addition of an enclosure video set-up is entirely new, and allows us to include more species, especially large and shy species that cannot be moved into a high-speed filming arena., Conclusions: We make recommendations for comparative whisker behaviour studies, particularly incorporating individual and species-specific considerations. We believe that flexible, comparative behavioural protocols have wide-ranging applications, specifically to better understand links between the brain and complex behaviours., Competing Interests: Declarations of interest None., (Copyright © 2022 The Authors. Published by Elsevier B.V. All rights reserved.)

Of all mammalian vibrissae, those of certain species of pinnipeds are exceptional. Researchers believe that their curious undulating form evolved for hydrodynamic detection. Our understanding of how these whiskers work depends on a geometrical model that captures the crucial pertinent features of the natural vibrissae including its tapering and curvature. It should also account for the form of the whisker when it flexes under external loading. We introduce and study a normal skeleton of a two-dimensional projection of a harbour seal whisker. The normal skeleton is a complete shape descriptor that involves reduction to the centreline equipped with a thickness function of the orthogonal cross-section. The contours of the whisker shape are extracted from a 2D greyscale scan. Our analysis reveals correspondence between the undulations of the width and oscillations of the centreline curvature as functions of arc length. We discuss two possible explanations for that remarkable feature: one based on consideration of growth and the other of plastic deformation. For the latter we employ a mechanical model to demonstrate appearance of curvature oscillations caused by extensive deflection of the undulating whisker due to external loading., (© 2022 IOP Publishing Ltd.)

Most cetaceans are born with vibrissae but they can be lost or reduced in adulthood, especially in odontocetes. Despite this, some species of odontocetes have been found to have functioning vibrissal follicles (including the follicle itself and any remaining vibrissal hair shaft) that play a role in mechanoreception, proprioception and electroreception. This reveals a greater diversity of vibrissal function in odontocetes than in any other mammalian group. However, we know very little about vibrissal follicle form and function across the Cetacea. Here, we qualitatively describe the gross vibrissal follicle anatomy of fetuses of three species of cetaceans, including two odontocetes: Atlantic white-sided dolphin (Lagenorhynchus acutus), harbour porpoise (Phocoena phocoena), and one mysticete: minke whale (Balaenoptera acutorostrata), and compared our findings to previous anatomical descriptions. All three species had few, short vibrissae contained within a relatively simple, single-part follicle, lacking in muscles. However, we observed differences in vibrissal number, follicle size and shape, and innervation distribution between the species. While all three species had nerve fibers around the follicles, the vibrissal follicles of Balaenoptera acutorostrata were innervated by a deep vibrissal nerve, and the nerve fibers of the odontocetes studied were looser and more branched. For example, in Lagenorhynchus acutus, branches of nerve fibers travelled parallel to the follicle, and innervated more superficial areas, rather than just the base. Our anatomical descriptions lend support to the observation that vibrissal morphology is diverse in cetaceans, and is worth further investigation to fully explore links between form and function., (© 2021 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

Whiskers are present in many species of mammals. They are specialised vibrotactile sensors that sit within strongly innervated follicles. Whisker size and shape will affect the mechanical signals that reach the follicle, and hence the information that reaches the brain. However, whisker size and shape have not been quantified across mammals before. Using a novel method for describing whisker curvature, this study quantifies whisker size and shape across 19 mammalian species. We find that gross two-dimensional whisker shape is relatively conserved across mammals. Indeed, whiskers are all curved, tapered rods that can be summarised by Euler spiral models of curvature and linear models of taper, which has implications for whisker growth and function. We also observe that aquatic and semi-aquatic mammals have relatively thicker, stiffer, and more highly tapered whiskers than arboreal and terrestrial species. In addition, smaller mammals tend to have relatively long, slender, flexible whiskers compared to larger species. Therefore, we propose that whisker morphology varies between larger aquatic species, and smaller scansorial species. These two whisker morphotypes are likely to induce quite different mechanical signals in the follicle, which has implications for follicle anatomy as well as whisker function., (© 2020 The Authors. Journal of Morphology published by Wiley Periodicals LLC.)

Whisker touch is an active sensory system. Previous studies in Pinnipeds have adopted relatively stationary tasks to judge tactile sensitivity, which may not accurately promote natural whisker movements and behaviours. This study developed a novel feeding task, termed fish sweeping to encourage whisker movements. Head and whisker movements were tracked from video footage in Harbor seal (Phoca vitulina), California sea lion (Zalophus californianus) and Pacific walrus (Odobenus rosmarus divergens). All species oriented their head towards the moving fish target and moved their whiskers during the task. Some species also engaged in whisker control behaviours, including head-turning asymmetry in the Pacific walrus, and contact-induced asymmetry in the Pacific walrus and California sea lion: behaviours that have only previously been observed in terrestrial mammals. This study confirms that Pinnipeds should be thought of as whisker specialists, and that whisker control (movement and positioning) is an important aspect of touch sensing in these animals, especially in sea lions and walruses. That the California sea lion controls whisker movement in relation to an object, and also had large values of whisker amplitude, spread and asymmetry, suggests that California sea lions are a promising model with which to further explore active touch sensing.

Active whisking in mice and rats is one of the fastest behaviours known in mammals and is used to guide complex behaviours such as exploration and navigation. During object contact, whisker movements are actively controlled and undergo robust changes in timing, speed and position. This study quantifies whisker movements in 6- to 7-month old male and female 5xFAD mice, and their C57/SJL F1 wild-type (WT) controls. As well as genotype, we examined sex differences and the effects of retinal degeneration (rd). Mice were filmed using a high-speed video camera at 500 frames per second (fps), under infrared light while behaving freely in three tasks: object exploration, sequential object exploration and tunnel running. Measures of whisker position, amplitude, speed and asymmetry were extracted and analysed for each task. The 5xFAD mice had significantly altered whisker angular positions, amplitude and asymmetry during object contacts and female 5xFAD mice with rd had lower mean angular positions during object contact. There were no significant effects of genotype on sequential object exploration or on tunnel running but differences due to sex and rd were found in both tasks, with female mice making larger and faster whisker movements overall, and mice with rd making larger and faster whisker movements during object contact. There were sex differences in whisker movements during sequential object exploration and females with rd had higher whisker retraction speeds in tunnel running. These data show that measuring whisker movements can quantify genotype and sex differences and the effects of rd during exploratory behaviour in these mice., (© 2018 John Wiley & Sons Ltd and International Behavioural and Neural Genetics Society.)

Background: Previous studies have measured whisker movements and locomotion to characterise mouse models of neurodegenerative disease. However, these studies have always been completed in isolation, and do not involve standardized procedures for comparisons across multiple mouse models and background strains., New Method: We present a standard method for conducting whisker movement and locomotion studies, by carrying out qualitative scoring and quantitative measurement of whisker movements from high-speed video footage of mouse models of Amyotrophic Lateral Sclerosis, Huntington's disease, Parkinson's disease, Alzheimer's disease, Cerebellar Ataxia, Somatosensory Cortex Development and Ischemic stroke., Results: Sex, background strain, source breeder and genotype all affected whisker movements. All mouse models, apart from Parkinson's disease, revealed differences in whisker movements during locomotion. R6/2 CAG250 Huntington's disease mice had the strongest behavioural phenotype. Robo3R 3-5 -CKO and RIM-DKOSert mouse models have abnormal somatosensory cortex development and revealed significant changes in whisker movements during object exploration., Comparison With Existing Method(s): Our results have good agreement with past studies, which indicates the robustness and reliability of measuring whisking. We recommend that differences in whisker movements of mice with motor deficits can be captured in open field arenas, but that mice with impairments to sensory or cognitive functioning should also be filmed investigating objects. Scoring clips qualitatively before tracking will help to structure later analyses., Conclusions: Studying whisker movements provides a quantitative measure of sensing, motor control and exploration. However, the effect of background strain, sex and age on whisker movements needs to be better understood., (Copyright © 2019 Elsevier B.V. All rights reserved.)

This paper reports on an analytical study of the intrinsic shapes of 523 whiskers from 15 rats. We show that the variety of whiskers on a rat's cheek, each of which has different lengths and shapes, can be described by a simple mathematical equation such that each whisker is represented as an interval on the Euler spiral. When all the representative curves of mystacial vibrissae for a single rat are assembled together, they span an interval extending from one coiled domain of the Euler spiral to the other. We additionally find that each whisker makes nearly the same angle of 47 ∘ with the normal to the spherical virtual surface formed by the tips of whiskers, which constitutes the rat's tactile sensory shroud or "search space." The implications of the linear curvature model for gaining insight into relationships between growth, form, and function are discussed., (Copyright © 2020 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works. Distributed under a Creative Commons Attribution NonCommercial License 4.0 (CC BY-NC).)

While most mammals have whiskers, some tactile specialists-mainly small, nocturnal, and arboreal species-can actively move their whiskers in a symmetrical, cyclic movement called whisking. Whisking enables mammals to rapidly, tactually scan their environment to efficiently guide locomotion and foraging in complex habitats. The muscle architecture that enables whisking is preserved from marsupials to primates, prompting researchers to suggest that a common ancestor might have had moveable whiskers. Studying the evolution of whisker touch sensing is difficult, and we suggest that measuring an aspect of skull morphology that correlates with whisking would enable comparisons between extinct and extant mammals. We find that whisking mammals have larger infraorbital foramen (IOF) areas, which indicates larger infraorbital nerves and an increase in sensory acuity. While this relationship is quite variable and IOF area cannot be used to solely predict the presence of whisking, whisking mammals all have large IOF areas. Generally, this pattern holds true regardless of an animal's substrate preferences or activity patterns. Data from fossil mammals and ancestral character state reconstruction and tracing techniques for extant mammals suggest that whisking is not the ancestral state for therian mammals. Instead, whisking appears to have evolved independently as many as seven times across the clades Marsupialia, Afrosoricida, Eulipotyphla, and Rodentia, with Xenarthra the only placental superordinal clade lacking whisking species. However, the term whisking only captures symmetrical and rhythmic movements of the whiskers, rather than all possible whisker movements, and early mammals may still have had moveable whiskers. Anat Rec, 2018. © 2018 American Association for Anatomy., (© 2018 American Association for Anatomy.)

Background: Previous studies have demonstrated that analysing whisker movements and locomotion allows us to quantify the behavioural consequences of sensory, motor and cognitive deficits in rodents. Independent whisker and feet trackers exist but there is no fully-automated, open-source software and hardware solution, that measures both whisker movements and gait., New Method: We present the LocoWhisk arena and new accompanying software (ARTv2) that allows the automatic detection and measurement of both whisker and gait information from high-speed video footage., Results: We demonstrate the new whisker and foot detector algorithms on high-speed video footage of freely moving small mammals, and show that whisker movement and gait measurements collected in the LocoWhisk arena are similar to previously reported values in the literature., Comparison With Existing Method(s): We demonstrate that the whisker and foot detector algorithms, are comparable in accuracy, and in some cases significantly better, than readily available software and manual trackers., Conclusion: The LocoWhisk system enables the collection of quantitative data from whisker movements and locomotion in freely behaving rodents. The software automatically records both whisker and gait information and provides added statistical tools to analyse the data. We hope the LocoWhisk system and software will serve as a solid foundation from which to support future research in whisker and gait analysis., (Copyright © 2019 Elsevier B.V. All rights reserved.)

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., (© 2018 The Author(s).)

Background: Motor dysfunction is a major component of the Huntington's disease (HD) phenotype, both in patients and animal models. Motor function in mice is usually measured using tests that involve a novel environment, or require a degree of learning, which creates potential confounds in animals, such as anxiety and/or learning., New Method: We propose that studying whisker control provides a more naturalistic way to measure motor function in HD mice. To this end we tested three strains of HD mice; R6/2 (CAG250), zQ175 and Hdh (CAG50, 150 and 250) mice., Results: We discovered a clear and progressive whisking deficit in the most severe model, the R6/2 CAG250 mouse. At 10 weeks, R6/2 mice showed an increase in whisking movements, which may be a correlate of the hyperkinesia seen in HD patients. By 18 weeks the R6/2 mice showed a reduction in whisking movements. Hdh Q250 mice showed a hyperkinetic profile at 10 weeks, approximately 4 months before other motor deficits have previously been reported in these mice. Q175 mice showed very little change in whisking behaviour, apart from a transient increase in retraction velocity at 10 weeks., Comparisons With Existing Methods: Our findings suggest that whisking may be a more sensitive test of motor function in HD mice than more commonly used methods, such as the rotarod., Conclusions: Our data suggest that whisking deficits represent a novel way of assessing the progression of the motor phenotype, and are early indicators for reversal of phenotype studies, such as drug trials., (Copyright © 2017 Elsevier B.V. All rights reserved.)

All mammals (apart from apes and humans) have whiskers that make use of a similar muscle arrangement. Whisker specialists, such as rats and mice, tend to be nocturnal and arboreal, relying on their whisker sense of touch to guide exploration around tree canopies at night. As such, nocturnal arboreal rodents have many whiskers that are organised into a grid-like pattern, and moved using a complex array of muscles. Indeed, most arboreal, nocturnal mammals tend to have specialised whiskers that are longer and arranged in a dense, regular grid, compared with terrestrial, diurnal mammals. The guinea pig diverged early from murid rodents (around 75 million years ago), and are ground-dwelling, diurnal animals. It would be predicted that, as a terrestrial mammal, they may have less whiskers and a reduced muscle architecture compared to arboreal, nocturnal rodents. We examined the mystacial whisker layout, musculature and movement capacity of Guinea pig (Cavia porcellus) whiskers and found that they did indeed have a disorganized whisker layout, with a fortification around the eye area. In addition, there was a reduction in musculature, especially in the intrinsic muscles. Despite guinea pigs not cyclically moving their whiskers, the mystacial musculature was still very similar to that of murid rodents. We suggest that the conserved presence of whisker layout and musculature, even in visual mammals such as primates and guinea pigs, may indicate that whiskers still play an important role in these animals, including protecting the eyes and being involved in tactile social behaviors. Anat Rec, 300:527-536, 2017. © 2016 Wiley Periodicals, Inc., (© 2016 Wiley Periodicals, Inc.)

Dormouse numbers are declining in the UK due to habitat loss and fragmentation. We know that dormice are nocturnal, arboreal, and avoid crossing open spaces between habitats, yet how they navigate around their canopy is unknown. As other rodents use whisker touch sensing to navigate and explore their environment, this study investigates whether Hazel dormice (Muscardinus avellanarius) employ their whiskers to cross between habitats. We analysed high-speed video footage of dormice exploring freely in flat and climbing arenas in near darkness and using infrared light illumination. We confirm that, like rats and mice, dormice move their whiskers back and forth continuously (~10 Hz) in a motion called whisking and recruit them to explore small gaps (<10 cm) by increasing the amplitude and frequency of whisking and also the asymmetry of movement between the left and right whisker fields. When gaps between platforms are larger than 10-15 cm, dormice spend more time travelling on the floor. These findings suggest that dormice can actively and purposively move their whiskers to gather relevant information from their canopy at night. As this species is vulnerable to threats on the ground, we also provide evidence that joining habitat patches between dormouse populations is important for promoting natural behaviours and movement between patches.

Nicotine, an important component of cigarette smoke, is a neurotransmitter that contributes to stress, depression and anxiety in smokers. In rodents, it increases anxiety and reduces exploratory behaviours. However, so far, the measurements of exploratory behaviour in rodents have only been semi-quantitative and lacking in sufficient detail to characterise the temporal effect of smoking cessation. As rodents, such as mice and rats, primarily use whiskers to explore their environment, we studied the effect of 3 months smoking with 1 and 2 weeks smoking cessation on whisker movements in mice, using high-speed video camera footage and image analysis. Both protraction and retraction whisker velocities were increased in smoking mice (p<0.001) and returned to normal following just one week of smoking cessation. In addition, locomotion speeds were decreased in smoking mice, and returned to normal following smoking cessation. Lung function was also impacted by smoking and remained impaired even following smoking cessation. We suggest that the increased whisker velocities in the smoking mice reflect reduced exploration and impeded tactile performance. The increase in whisker velocity with smoking, and its reduction following smoking cessation, also lends support to acetylcholine being involved in awareness, attention and alertness pathways. It also shows that smoking-induced behavioural changes can be reversed with smoking cessation, which may have implications for human smokers., (Copyright © 2016 Elsevier B.V. All rights reserved.)

Studies in pinniped whisker use have shown that their whiskers are extremely sensitive to tactile and hydrodynamic signals. While pinnipeds position their whiskers on to objects and have some control over their whisker protractions, it has always been thought that head movements are more responsible for whisker positioning than the movement of the whiskers themselves. This study uses ball balancing, a dynamic sensorimotor skill that is often used in human and robotic coordination studies, to promote sea lion whisker movements during the task. For the first time, using tracked video footage, we show that sea lion whisker movements respond quickly (26.70 ms) and mirror the movement of the ball, much more so than the head. We show that whisker asymmetry and spread are both altered to help sense and control the ball during balancing. We believe that by designing more dynamic sensorimotor tasks we can start to characterise the active nature of this specialised sensory system in pinnipeds.

During exploration, rats and other small mammals make rhythmic back-and-forth sweeps of their long facial whiskers (macrovibrissae) [1-3]. These "whisking" movements are modulated by head movement [4] and by vibrissal sensory input [5, 6] and hence are often considered "active" in the Gibsonian sense of being purposive and information seeking [7, 8]. An important hallmark of active sensing is the modification of the control strategy according to context [9]. Using a task in which rats were trained to run circuits for food, we tested the hypothesis that whisker control, as measured by high-speed videography, changes with contextual variables such as environment familiarity, risk of collision, and availability of visual cues. In novel environments, functionally blind rats moved at slow speeds and performed broad whisker sweeps. With greater familiarity, however, they moved more rapidly, protracted their whiskers further, and showed decreased whisking amplitude. These findings indicate a strategy change from using the vibrissae to explore nearby surfaces to using them primarily for "look ahead." In environments with increased risk of collision, functionally blind animals moved more slowly but protracted their whiskers further. Sighted animals also showed changes in whisker control strategy with increased familiarity, but these changes were different to those of the functionally blind strain. Sighted animals also changed their vibrissal behavior when visual cues were subsequently removed (by being placed in darkness). These contextual influences provide strong evidence of active control and demonstrate that the vibrissal system provides an accessible model of purposive behavior in mammals., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

The transgenic SOD1(G93A) mouse is a model of human amyotrophic lateral sclerosis (ALS) and recapitulates many of the pathological hallmarks observed in humans, including motor neuron degeneration in the brain and the spinal cord. In mice, neurodegeneration particularly impacts on the facial nuclei in the brainstem. Motor neurons innervating the whisker pad muscles originate in the facial nucleus of the brain stem, with contractions of these muscles giving rise to "whisking" one of the fastest movements performed by mammals. A longitudinal study was conducted on SOD1(G93A) mice and wild-type litter mate controls, comparing: (i) whisker movements using high-speed video recordings and automated whisker tracking, and (ii) facial nucleus degeneration using MRI. Results indicate that while whisking still occurs in SOD1(G93A) mice and is relatively resistant to neurodegeneration, there are significant disruptions to certain whisking behaviours, which correlate with facial nuclei lesions, and may be as a result of specific facial muscle degeneration. We propose that measures of mouse whisker movement could potentially be used in tandem with measures of limb dysfunction as biomarkers of disease onset and progression in ALS mice and offers a novel method for testing the efficacy of novel therapeutic compounds., (Copyright © 2013 Elsevier B.V. All rights reserved.)

Facial vibrissae, or whiskers, are found in nearly all extant mammal species and are likely to have been present in early mammalian ancestors. A sub-set of modern mammals, including many rodents, move their long mystacial whiskers back and forth at high speed whilst exploring in a behaviour known as 'whisking'. It is not known whether the vibrissae of early mammals moved in this way. The grey short-tailed opossum, Monodelphis domestica, is considered a useful species from the perspective of tracing the evolution of modern mammals. Interestingly, these marsupials engage in whisking bouts similar to those seen in rodents. To better assess the likelihood that active vibrissal sensing was present in ancestral mammals, we examined the vibrissal musculature of the opossum using digital miscroscopy to see whether this resembles that of rodents. Although opossums have fewer whiskers than rats, our investigation found that they have a similar vibrissal musculature. In particular, in both rats and opossums, the musculature includes both intrinsic and extrinsic muscles with the intrinsic muscles positioned as slings linking pairs of large vibrissae within rows. We identified some differences in the extrinsic musculature which, interestingly, matched with behavioural data obtained through high-speed video recording, and indicated additional degrees of freedom for positioning the vibrissae in rats. These data show that the whisker movements of opossum and rat exploit similar underlying mechanisms. Paired with earlier results suggesting similar patterns of vibrissal movement, this strongly implies that early therian (marsupial and placental) mammals were whisking animals that actively controlled their vibrissae.

Adult rats sweep their large facial whiskers (macrovibrissae) back and forth in a rhythmic pattern known as "whisking". Here we examine how these whisker movements develop in relation to other aspects of exploratory behavior, particularly locomotion. We analyzed 963 high-speed video recordings of neonatal rats, from P1 (Post-natal day 1) to P21, and measured the emergence of whisker control and of head, body, and limb movements. Prior to P11, whisker movements were largely limited to unilateral retractions accompanying head turns. Between P11 and P13 bilateral whisking emerged alongside increased forward locomotion and improved control of the head. Contact-induced modulations of whisking symmetry, synchrony, and whisker spread emerge shortly thereafter but continue to develop until at least P18, coinciding with the emergence of adult-like locomotion patterns such as rearing. Overall, whisking develops alongside increasing locomotor competence indicating that active vibrissal sensing plays an important role in the exploratory behavior of the developing animal., (Copyright © 2011 Wiley Periodicals, Inc.)

In rats, the long facial whiskers (mystacial macrovibrissae) are repetitively and rapidly swept back and forth during exploration in a behaviour known as 'whisking'. In this paper, we summarize previous evidence from rats, and present new data for rat, mouse and the marsupial grey short-tailed opossum (Monodelphis domestica) showing that whisking in all three species is actively controlled both with respect to movement of the animal's body and relative to environmental structure. Using automatic whisker tracking, and Fourier analysis, we first show that the whisking motion of the mystacial vibrissae, in the horizontal plane, can be approximated as a blend of two sinusoids at the fundamental frequency (mean 8.5, 11.3 and 7.3 Hz in rat, mouse and opossum, respectively) and its second harmonic. The oscillation at the second harmonic is particularly strong in mouse (around 22 Hz) consistent with previous reports of fast whisking in that species. In all three species, we found evidence of asymmetric whisking during head turning and following unilateral object contacts consistent with active control of whisker movement. We propose that the presence of active vibrissal touch in both rodents and marsupials suggests that this behavioural capacity emerged at an early stage in the evolution of therian mammals.

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.

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.