Active Whisking Research Articles

At the onset of active whisking, the input layer of barrel cortex exhibits a 24 h window of increased excitability that depends on prior experience.

The development of motor control over sensory organs is a critical milestone in sensory processing, enabling active exploration and shaping of the sensory environment. However, whether the onset of sensory organ motor control directly influences the development of corresponding sensory cortices remains unknown. Here, we exploit the late onset of whisking behavior in mice to address this question in the somatosensory system. Using ex vivo electrophysiology, we discovered a transient increase in the intrinsic excitability of excitatory neurons in layer IV of the barrel cortex, which processes whisker input, precisely coinciding with the onset of active whisking at postnatal day 14 (P14). This increase in neuronal gain was specific to layer IV, independent of changes in synaptic strength, and required prior sensory experience. Strikingly, the effect was not observed in layer II/III of the barrel cortex or in the visual cortex upon eye opening, suggesting a unique interaction between the development of active sensing and the thalamocortical input layer in the somatosensory system. Predictive modeling indicated that changes in active membrane conductances alone could reliably distinguish P14 neurons in control but not whisker-deprived hemispheres. Our findings demonstrate an experience-dependent, lamina-specific refinement of neuronal excitability tightly linked to the emergence of active whisking. This transient increase in the gain of the thalamic input layer coincides with a critical period for synaptic plasticity in downstream layers, suggesting a role in facilitating cortical maturation and sensory processing. Together, our results provide evidence for a direct interaction between the development of motor control and sensory cortex, offering new insights into the experience-dependent development and refinement of sensory systems. These findings have broad implications for understanding the interplay between motor and sensory development, and how the mechanisms of perception cooperate with behavior.

Relating Pupil Diameter and Blinking to Cortical Activity and Hemodynamics across Arousal States.

Arousal state affects neural activity and vascular dynamics in the cortex, with sleep associated with large changes in the local field potential and increases in cortical blood flow. We investigated the relationship between pupil diameter and blink rate with neural activity and blood volume in the somatosensory cortex in male and female unanesthetized, head-fixed mice. We monitored these variables while the mice were awake, during periods of rapid eye movement (REM), and non-rapid eye movement (NREM) sleep. Pupil diameter was smaller during sleep than in the awake state. Changes in pupil diameter were coherent with both gamma-band power and blood volume in the somatosensory cortex, but the strength and sign of this relationship varied with arousal state. We observed a strong negative correlation between pupil diameter and both gamma-band power and blood volume during periods of awake rest and NREM sleep, although the correlations between pupil diameter and these signals became positive during periods of alertness, active whisking, and REM. Blinking was associated with increases in arousal and decreases in blood volume when the mouse was asleep. Bilateral coherence in gamma-band power and in blood volume dropped following awake blinking, indicating a reset of neural and vascular activity. Using only eye metrics (pupil diameter and eye motion), we could determine the arousal state of the mouse ('Awake,' 'NREM,' 'REM') with >90% accuracy with a 5 s resolution. There is a strong relationship between pupil diameter and hemodynamics signals in mice, reflecting the pronounced effects of arousal on cerebrovascular dynamics.SIGNIFICANCE STATEMENT Determining arousal state is a critical component of any neuroscience experiment. Pupil diameter and blinking are influenced by arousal state, as are hemodynamics signals in the cortex. We investigated the relationship between cortical hemodynamics and pupil diameter and found that pupil diameter was strongly related to the blood volume in the cortex. Mice were more likely to be awake after blinking than before, and blinking resets neural activity. Pupil diameter and eye motion can be used as a reliable, noninvasive indicator of arousal state. As mice transition from wake to sleep and back again over a timescale of seconds, monitoring pupil diameter and eye motion permits the noninvasive detection of sleep events during behavioral or resting-state experiments.

Olfactory modulation of barrel cortex activity during active whisking and passive whisker stimulation

Rodents depend on olfaction and touch to meet many of their fundamental needs. However, the impact of simultaneous olfactory and tactile inputs on sensory representations in the cortex remains elusive. To study these interactions, we recorded large populations of barrel cortex neurons using 2-photon calcium imaging in head-fixed mice during olfactory and tactile stimulation. Here we show that odors bidirectionally alter activity in a small but significant population of barrel cortex neurons through at least two mechanisms, first by enhancing whisking, and second by a central mechanism that persists after whisking is abolished by facial nerve sectioning. Odor responses have little impact on tactile information, and they are sufficient for decoding odor identity, while behavioral parameters like whisking, sniffing, and facial movements are not odor identity-specific. Thus, barrel cortex activity encodes specific olfactory information that is not linked with odor-induced changes in behavior.

The Role of Inhibitory Interneurons in Circuit Assembly and Refinement Across Sensory Cortices

Sensory information is transduced into electrical signals in the periphery by specialized sensory organs, which relay this information to the thalamus and subsequently to cortical primary sensory areas. In the cortex, microcircuits constituted by interconnected pyramidal cells and inhibitory interneurons, distributed throughout the cortical column, form the basic processing units of sensory information underlying sensation. In the mouse, these circuits mature shortly after birth. In the first postnatal week cortical activity is characterized by highly synchronized spontaneous activity. While by the second postnatal week, spontaneous activity desynchronizes and sensory influx increases drastically upon eye opening, as well as with the onset of hearing and active whisking. This influx of sensory stimuli is fundamental for the maturation of functional properties and connectivity in neurons allocated to sensory cortices. In the subsequent developmental period, spanning the first five postnatal weeks, sensory circuits are malleable in response to sensory stimulation in the so-called critical periods. During these critical periods, which vary in timing and duration across sensory areas, perturbations in sensory experience can alter cortical connectivity, leading to long-lasting modifications in sensory processing. The recent advent of intersectional genetics, in vivo calcium imaging and single cell transcriptomics has aided the identification of circuit components in emergent networks. Multiple studies in recent years have sought a better understanding of how genetically-defined neuronal subtypes regulate circuit plasticity and maturation during development. In this review, we discuss the current literature focused on postnatal development and critical periods in the primary auditory (A1), visual (V1), and somatosensory (S1) cortices. We compare the developmental trajectory among the three sensory areas with a particular emphasis on interneuron function and the role of inhibitory circuits in cortical development and function.

Contribution of Interneuron Subtype-Specific GABAergic Signaling to Emergent Sensory Processing in Mouse Somatosensory Whisker Barrel Cortex.

Mammalian neocortex is important for conscious processing of sensory information with balanced glutamatergic and GABAergic signaling fundamental to this function. Yet little is known about how this interaction arises despite increasing insight into early GABAergic interneuron (IN) circuits. To study this, we assessed the contribution of specific INs to the development of sensory processing in the mouse whisker barrel cortex, specifically the role of INs in early speed coding and sensory adaptation. In wild-type animals, both speed processing and adaptation were present as early as the layer 4 critical period of plasticity and showed refinement over the period leading to active whisking onset. To test the contribution of IN subtypes, we conditionally silenced action-potential-dependent GABA release in either somatostatin (SST) or vasoactive intestinal peptide (VIP) INs. These genetic manipulations influenced both spontaneous and sensory-evoked cortical activity in an age- and layer-dependent manner. Silencing SST + INs reduced early spontaneous activity and abolished facilitation in sensory adaptation observed in control pups. In contrast, VIP + IN silencing had an effect towards the onset of active whisking. Silencing either IN subtype had no effect on speed coding. Our results show that these IN subtypes contribute to early sensory processing over the first few postnatal weeks.

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.

Developmental divergence of sensory stimulus representation in cortical interneurons

Vasocative-intestinal-peptide (VIP+) and somatostatin (SST+) interneurons are involved in modulating barrel cortex activity and perception during active whisking. Here we identify a developmental transition point of structural and functional rearrangements onto these interneurons around the start of active sensation at P14. Using in vivo two-photon Ca2+ imaging, we find that before P14, both interneuron types respond stronger to a multi-whisker stimulus, whereas after P14 their responses diverge, with VIP+ cells losing their multi-whisker preference and SST+ neurons enhancing theirs. Additionally, we find that Ca2+ signaling dynamics increase in precision as the cells and network mature. Rabies virus tracings followed by tissue clearing, as well as photostimulation-coupled electrophysiology reveal that SST+ cells receive higher cross-barrel inputs compared to VIP+ neurons at both time points. In addition, whereas prior to P14 both cell types receive direct input from the sensory thalamus, after P14 VIP+ cells show reduced inputs and SST+ cells largely shift to motor-related thalamic nuclei.

Temporal refinement of sensory-evoked activity across layers in developing mouse barrel cortex.

Rhythmic whisking behavior in rodents fully develops during a critical period about 2weeks after birth, in parallel with the maturation of other sensory modalities and the onset of exploratory locomotion. How whisker-related sensory processing develops during this period in the primary somatosensory cortex (S1) remains poorly understood. Here, we characterized neuronal activity evoked by single- or dual-whisker stimulation patterns in developing S1, before, during and after the occurrence of active whisking. Employing multi-electrode recordings in all layers of barrel cortex in urethane-anesthetized mice, we find layer-specific changes in multi-unit activity for principal and neighboring barrel columns. While whisker stimulation evoked similar early responses (0-50ms post-stimulus) across development, the late response (50-150ms post-stimulus) decreased in all layers with age. Furthermore, peak onset times and the duration of the late response decreased in all layers across age groups. Responses to paired-pulse stimulation showed increases in spiking precision and in paired-pulse ratios in all cortical layers during development. Sequential activation of two neighboring whiskers with varying stimulus intervals evoked distinct response profiles in the activated barrel columns, depending on the direction and temporal separation of the stimuli. In conclusion, our findings indicate that the temporal sharpening of sensory-evoked activity coincides with the onset of active whisking.

Whisker exploration behaviours in the 5xFAD mouse are affected by sex and retinal degeneration.

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.

Decoding cortical brain states from widefield calcium imaging data using visibility graph.

Widefield optical imaging of neuronal populations over large portions of the cerebral cortex in awake behaving animals provides a unique opportunity for investigating the relationship between brain function and behavior. In this paper, we demonstrate that the temporal characteristics of calcium dynamics obtained through widefield imaging can be utilized to infer the corresponding behavior. Cortical activity in transgenic calcium reporter mice (n=6) expressing GCaMP6f in neocortical pyramidal neurons is recorded during active whisking (AW) and no whisking (NW). To extract features related to the temporal characteristics of calcium recordings, a method based on visibility graph (VG) is introduced. An extensive study considering different choices of features and classifiers is conducted to find the best model capable of predicting AW and NW from calcium recordings. Our experimental results show that temporal characteristics of calcium recordings identified by the proposed method carry discriminatory information that are powerful enough for decoding behavior.

Radial Distance Estimation with Tapered Whisker Sensors.

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.

Bilateral Discrimination of Tactile Patterns without Whisking in Freely Running Rats.

A majority of whisker discrimination tasks in rodents are performed on head-fixed animals to facilitate tracking or control of the sensory inputs. However, head fixation critically restrains the behavior and thus the incoming stimuli compared with those occurring in natural conditions. In this study, we investigated whether freely behaving rats can discriminate fine tactile patterns while running, in particular when stimuli are presented simultaneously on both sides of the snout. We developed a two-alternative forced-choice task in an automated modified T-maze. Stimuli were either a surface with no bars (smooth) or with vertical bars spaced irregularly or regularly. While running at full speed, rats encountered simultaneously the two discriminanda placed on the two sides of the central aisle. Rats learned to recognize regular bars versus a smooth surface in 8 weeks. They solved the task while running at an average speed of 1 m/s, so that the contact with the stimulus lasted <1 typical whisking cycle, precluding the use of active whisking. Whisker-tracking analysis revealed an asymmetry in the position of the whiskers: they oriented toward the rewarded stimulus during successful trials as early as 60 ms after the first possible contact. We showed that the whiskers and activity in the primary somatosensory cortex are involved during the discrimination process. Finally, we identified irregular patterns of bars that the rats can discriminate from the regular one. This novel task shows that freely moving rodents can make simultaneous bilateral tactile discrimination without whisking.SIGNIFICANCE STATEMENT The whisker system of rodents is a widely used model to study tactile processing. Rats show remarkable abilities in discriminating surfaces by actively moving their whiskers (whisking) against stimuli, typically sampling them several times. This motor strategy affects considerably the way that tactile information is acquired and thus the way that neuronal networks process the information. However, when rats run at high speed, they protract their whiskers in front of the snout without large movements. Here, we investigated whether rats are able to discriminate regular and irregular patterns of vertical bars while running without whisking. We found that the animals can perform a bilateral simultaneous discrimination without whisking and that this involves both whiskers and barrel cortex activity.

Vibrissal paralysis produces increased corticosterone levels and impairment of spatial memory retrieval

This research was aimed at establishing how the absence of active whisking in rats affects acquisition and recovery of spatial memory. The mystacial vibrissae were irreversibly paralyzed by cutting the facial nerve’s mandibular and buccal branches bilaterally in the facial nerve lesion group (N=14); control animals were submitted to sham-surgery (N=15). Sham-operated (N=11) and facial nerve-lesioned (N=10) animals were trained (one session, eight acquisition trials) and tested 24h later in a circular Barnes maze. It was found that facial nerve lesioned-animals adequately acquired the spatial task, but had impaired recovery of it when tested 24h after training as compared to control ones. Plasma corticosterone levels were measured after memory testing in four randomly chosen animals of each trained group and after a single training trial in the maze in additional facial nerve-lesioned (N=4) and sham-operated animals (N=4). Significant differences respecting the elevation of corticosterone concentration after either a single training trial or memory testing indicated that stress response was enhanced in facial nerve-lesioned animals as compared to control ones. Increased corticosterone levels during training and testing might have elicited the observed whisker paralysis-induced spatial memory retrieval impairment.

The cerebellum linearly encodes whisker position during voluntary movement.

Active whisking is an important model sensorimotor behavior, but the function of the cerebellum in the rodent whisker system is unknown. We have made patch clamp recordings from Purkinje cells in vivo to identify whether cerebellar output encodes kinematic features of whisking including the phase and set point. We show that Purkinje cell spiking activity changes strongly during whisking bouts. On average, the changes in simple spike rate coincide with or slightly precede movement, indicating that the synaptic drive responsible for these changes is predominantly of efferent (motor) rather than re-afferent (sensory) origin. Remarkably, on-going changes in simple spike rate provide an accurate linear read-out of whisker set point. Thus, despite receiving several hundred thousand discrete synaptic inputs across a non-linear dendritic tree, Purkinje cells integrate parallel fiber input to generate precise information about whisking kinematics through linear changes in firing rate.

Pathway-specific reorganization of projection neurons in somatosensory cortex during learning.

In the mammalian brain, sensory cortices exhibit plasticity during task learning, but how this alters information transferred between connected cortical areas remains unknown. We found that divergent subpopulations of cortico-cortical neurons in mouse whisker primary somatosensory cortex (S1) undergo functional changes reflecting learned behavior. We chronically imaged activity of S1 neurons projecting to secondary somatosensory (S2) or primary motor (M1) cortex in mice learning a texture discrimination task. Mice adopted an active whisking strategy that enhanced texture-related whisker kinematics, correlating with task performance. M1-projecting neurons reliably encoded basic kinematics features, and an additional subset of touch-related neurons was recruited that persisted past training. The number of S2-projecting touch neurons remained constant, but improved their discrimination of trial types through reorganization while developing activity patterns capable of discriminating the animal's decision. We propose that learning-related changes in S1 enhance sensory representations in a pathway-specific manner, providing downstream areas with task-relevant information for behavior.

Modulation of artificial whisking related signals in barrel cortex.

Rats use rhythmic whisker movements, called active whisking, to sense the environment, which include whisker protractions followed by retractions at various frequencies. Using a proxy of active whisking in anesthetized rats, called artificial whisking, which is induced by electrically stimulating the facial motor nerve, we characterized the neural responses evoked in the barrel cortex by whisking in air (without contact) and on a surface (with contact). Neural responses were compared between distinct network states consisting of cortical deactivation (synchronized slow oscillations) and activation (desynchronized state) produced by neuromodulation (cholinergic or noradrenergic stimulation in neocortex or thalamus). Here we show that population responses in the barrel cortex consist of a robust signal driven by the onset of the whisker protraction followed by a whisking retraction signal that emerges during low frequency whisking on a surface. The whisking movement onset signal is suppressed by increasing whisking frequency, is controlled by cortical synaptic inhibition, is suppressed during cortical activation states, is little affected by whisking on a surface, and is ubiquitous in ventroposterior medial (VPM) thalamus, barrel cortex, and superior colliculus. The whisking retraction signal codes the duration of the preceding whisker protraction, is present in thalamocortical networks but not in superior colliculus, and is robust during cortical activation; a state associated with natural exploratory whisking. The expression of different whisking signals in forebrain and midbrain may define the sensory processing abilities of those sensorimotor circuits. Whisking related signals in the barrel cortex are controlled by network states that are set by neuromodulators.

The role of environmental feedback in a brain state switch from passive to active sensing

Coherent behaviour emerges from mutual interaction between the brain, body and environment across multiple timescales and not from within the brain alone [1,2]. For example sensation is actively shaped by dynamical interaction of the brain and environment through motor actions such as sniffing, saccading, and touching. The onset of active sensing is often concomitant with qualitative changes in neural dynamics [3] and responses to sensory input [4,5]. Indeed, some neural responses are uniquely sensitive to the presence or absence of dynamical sensory feedback [6]. However, understanding how active sensing strategies impact on neural dynamics and sensory responses is an open challenge. Whisking behaviour in rodents has been the central model system for studying neural mechanisms of active sensing. During quiet wakefulness the membrane potential of neurons in the barrel cortex exhibit high power, low frequency fluctuations and nearby neurons become highly correlated [3]. As active whisking onsets, the brain state of the barrel cortex qualitatively changes: low frequency fluctuations are suppressed and nearby neurons decorrelate [3]. Interestingly both correlation and low frequency LFP are partially restored during periods of active touch, i.e., when the mouse palpate towards, and repeatedly contacts with an object [3]. The brain state transition is also concomitant with changes in responses of barrel cortex neurons to whisker stimulation. The sensitivity of neurons to whisker perturbation drops with the onset of whisking [5]. However, robust and repeatable whisker response are recovered for more naturalistic stimuli, i.e., active touch [5]. Here we propose a theory, and construct a model, of active whisking that explains the core phenomenology through the dynamical interaction between the brain and the environment. The three core assumptions underlying the theory are: 1. Strong low frequency membrane fluctuations, intra-neural correlations between nearby neurons, and sensory responses to brief whisker deflection associative with quiet attentive state arise as result of network dynamics that are close to a dynamical instability. 2. Reafferent input (sensory feedback related to self-action) during whisking behaviour provides negative feedback to sensory neurons that stabilize cortical dynamics reducing low frequency fluctuations, intra neural correlations, sensory responses to brief whisker deflection, while increasing the correlation between cortical activity and whisker position. 3. Interrupting the reafferent signal, via a whisker touch event, temporally destabilise the cortex and enhances ex-afferent input (external sensory input coding whisker contacts). Our theory and model suggests that sensory feedback via reafference signals is sufficient to account for the changes in cortical dynamics that appear with the onset of active whisking and provides a novel mechanistic account for sensory processing during active whisking. We discuss how this feedback stabilization mechanism coexists and interacts with other internal mechanisms that modulate cortical dynamics within the brain [3,7]. We quantify how this mechanism of active touch enhances the signal-to noise ratio of the input to the cortex. Finally we discuss the wider implications of these results for experimental work attempting to characterise neural activity and responses in an open-loop (in the absence of sensory/motor feedback) condition.

Superior colliculus cells sensitive to active touch and texture during whisking

Rats sense the environment through rhythmic vibrissa protractions, called active whisking, which can be simulated in anesthetized rats by electrically stimulating the facial motor nerve. Using this method, we investigated barrel cortex field potential and superior colliculus single-unit responses during passive touch, whisking movement, active touch, and texture discrimination. Similar to passive touch, whisking movement is signaled during the onset of the whisker protraction by short-latency responses in barrel cortex that drive corticotectal responses in superior colliculus, and all these responses show robust adaptation with increases in whisking frequency. Active touch and texture are signaled by longer latency responses, first in superior colliculus during the rising phase of the protraction, likely driven by trigeminotectal inputs, and later in barrel cortex by the falling phase of the protraction. Thus, superior colliculus is part of a broader vibrissa neural network that can decode whisking movement, active touch, and texture.

Electrophysiological characterization of texture information slip-resistance dependent in the rat vibrissal nerve

BackgroundStudies in tactile discrimination agree that rats are able to learn a rough-smooth discrimination task by actively touching (whisking) objects with their vibrissae. In particular, we focus on recent evidence of how neurons at different levels of the sensory pathway carry information about tactile stimuli. Here, we analyzed the multifiber afferent discharge of one vibrissal nerve during active whisking. Vibrissae movements were induced by electrical stimulation of motor branches of the facial nerve. We used sandpapers of different grain size as roughness discrimination surfaces and we also consider the change of vibrissal slip-resistance as a way to improve tactile information acquisition. The amplitude of afferent activity was analyzed according to its Root Mean Square value (RMS). The comparisons among experimental situation were quantified by using the information theory.ResultsWe found that the change of the vibrissal slip-resistance is a way to improve the roughness discrimination of surfaces. As roughness increased, the RMS values also increased in almost all cases. In addition, we observed a better discrimination performance in the retraction phase (maximum amount of information).ConclusionsThe evidence of amplitude changes due to roughness surfaces and slip-resistance levels allows to speculate that texture information is slip-resistance dependent at peripheral level.

Three-dimensional axon morphologies of individual layer 5 neurons indicate cell type-specific intracortical pathways for whisker motion and touch

The cortical output layer 5 contains two excitatory cell types, slender- and thick-tufted neurons. In rat vibrissal cortex, slender-tufted neurons carry motion and phase information during active whisking, but remain inactive after passive whisker touch. In contrast, thick-tufted neurons reliably increase spiking preferably after passive touch. By reconstructing the 3D patterns of intracortical axon projections from individual slender- and thick-tufted neurons, filled in vivo with biocytin, we were able to identify cell type-specific intracortical circuits that may encode whisker motion and touch. Individual slender-tufted neurons showed elaborate and dense innervation of supragranular layers of large portions of the vibrissal area (total length, 86.8 ± 5.5 mm). During active whisking, these long-range projections may modulate and phase-lock the membrane potential of dendrites in layers 2 and 3 to the whisking cycle. Thick-tufted neurons with soma locations intermingling with those of slender-tufted ones display less dense intracortical axon projections (total length, 31.6 ± 14.3 mm) that are primarily confined to infragranular layers. Based on anatomical reconstructions and previous measurements of spiking, we put forward the hypothesis that thick-tufted neurons in rat vibrissal cortex receive input of whisker motion from slender-tufted neurons onto their apical tuft dendrites and input of whisker touch from thalamic neurons onto their basal dendrites. During tactile-driven behavior, such as object location, near-coincident input from these two pathways may result in increased spiking activity of thick-tufted neurons and thus enhanced signaling to their subcortical targets.