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Proprioception refers to the ability to perceive the position and movement of body segments in space. The cortical aspects of the proprioceptive afference from the body can be investigated using corticokinematic coherence (CKC). CKC accurately quanti- fies the degree of coupling between cortical activity and limb kinematics, especially if precise proprioceptive stimulation of evoked movements is used. However, there is no evidence on how volitional muscle activation during proprioceptive stimulation affects CKC strength. Twenty-five healthy volunteers (28.8 ± 7 yr, 11 females) participated in the experiment, which included electroencephalographic (EEG), electromyographic (EMG), and kinematic recordings. Ankle-joint rotations (2-Hz) were elicited through a movement actuator in two conditions: passive condition with relaxed ankle and active condition with constant 5-Nm plantar flexion exerted during the stimulation. In total, 6 min of data were recorded per condition. CKC strength was defined as the maximum coherence value among all the EEG channels at the 2-Hz movement frequency for each condition separately. Both conditions resulted in significant CKC peaking at the Cz electrode over the foot area of the primary sensorimotor (SM1) cortex. Stronger CKC was found for the active (0.13 ± 0.14) than the passive (0.03 ± 0.04) condition (P < 0.01). The results indicated that volitional activation of the muscles intensifies the neuronal proprioceptive processing in the SM1 cortex. This finding could be explained both by peripheral sensitization of the ankle joint proprioceptors and central modulation of the neuronal proprioceptive processing at the spinal and cortical levels. [ABSTRACT FROM AUTHOR]

Neural plasticity of the brain or its ability to reorganize following injury has likely coincided with the successful clinical correction of severe deformity by facial transplantation since 2005. In this study, we present the cortical reintegration outcomes following syngeneic hemifacial vascularized composite allograft (VCA) in a small animal model. Specifically, changes in the topographic organization and unit response properties of the rodent whisker-barrel somatosensory system were assessed following hemifacial VCA. Clear differences emerged in the barrel-cortex system when comparing naïve and hemiface transplanted animals. Neurons in the somatosensory cortex of transplanted rats had decreased sensitivity albeit increased directional sensitivity compared with naïve rats and evoked responses in transplanted animals were more temporally dispersed. In addition, receptive fields were often topographically mismatched with the indication that the mismatched topography reorganized within adjacent barrel (same row-arc bias following hemifacial transplant). These results suggest subcortical changes in the thalamus and/or brainstem play a role in hemifacial transplantation cortical plasticity and demonstrate the discrete and robust data that can be derived from this clinically relevant small animal VCA model for use in optimizing postsurgical outcomes. [ABSTRACT FROM AUTHOR]

Speech perception is known to be a multimodal process, relying not only on auditory input but also on the visual system and possibly on the motor system as well. To date there has been little work on the potential involvement of the somatosensory system in speech perception. In the present review, we identify the somatosensory system as another contributor to speech perception. First, we argue that evidence in favor of a motor contribution to speech perception can just as easily be interpreted as showing somatosensory involvement. Second, physiological and neuroanatomical evidence for auditory-somatosensory interactions across the auditory hierarchy indicates the availability of a neural infrastructure that supports somatosensory involvement in auditory processing in general. Third, there is accumulating evidence for somatosensory involvement in the context of speech specifically. In particular, tactile stimulation modifies speech perception, and speech auditory input elicits activity in somatosensory cortical areas. Moreover, speech sounds can be decoded from activity in somatosensory cortex; lesions to this region affect perception, and vowels can be identified based on somatic input alone. We suggest that the somatosensory involvement in speech perception derives from the somatosensory-auditory pairing that occurs during speech production and learning. By bringing together findings from a set of studies that have not been previously linked, the present article identifies the somatosensory system as a presently unrecognized contributor to speech perception. [ABSTRACT FROM AUTHOR]

Seminal neurophysiological studies in the 1940s discovered two somatomotor maps in the cerebellum--an inverted anterior lobe map and an upright posterior lobe map. Both maps have been confirmed in the human using noninvasive neuroimaging with additional hints of a third map within and near to the cerebellar vermis. Here, we sought direct evidence for the third somatomotor map by using intensive, repeated functional MRI (fMRI) scanning of individuals performing movements across multiple body parts (tongue, hands, glutes, and feet). An initial discovery sample (n = 4, 4 sessions per individual including 576 separate blocks of body movements) yielded evidence for the two established cerebellar somatomotor maps, as well as evidence for a third discontinuous foot representation within the vermis. When the left versus right foot movements were directly contrasted, the third representation could be clearly distinguished from the second representation in multiple individuals. Functional connectivity from seed regions in the third somatomotor representation confirmed anatomically specific connectivity with the cerebral cortex, paralleling the patterns observed for the two well-established maps. All results were prospectively replicated in an independent dataset with new individuals (n = 4). These collective findings provide direct support for a third somatomotor representation in the vermis of the cerebellum that may be part of a third map. We discuss the relations of this candidate third map to the broader topography of the cerebellum as well as its implications for understanding the specific organization of the human cerebellar vermis where distinct zones appear functionally specialized for somatomotor and visual domains. [ABSTRACT FROM AUTHOR]

Interactions with objects involve simultaneous contact with multiple, not necessarily adjacent, skin regions. Although advances have been made in understanding the capacity to selectively attend to a single tactile element among distracting stimulations, here, we examine how multiple stimulus elements are explicitly integrated into an overall tactile percept. Across four experiments, participants averaged the direction of two simultaneous tactile motion trajectories of varying discrepancy delivered to different fingerpads. Averaging performance differed between within- and between-hands conditions in terms of sensitivity and precision but was unaffected by somatotopic proximity between stimulated fingers. First, precision was greater in between-hand compared with within-hand conditions, demonstrating a bimanual perceptual advantage in multi-touch integration. Second, sensitivity to the average direction was influenced by the discrepancy between individual motion signals, but only for within-hand conditions. Overall, our experiments identify key factors that influence perception of simultaneous tactile events. In particular, we show that multi-touch integration is constrained by hand-specific rather than digit-specific mechanisms. NEW & NOTEWORTHY Object manipulation involves encoding spatially and temporally extended tactile signals, yet most studies emphasize minimal units of tactile perception (e.g., selectivity). Instead, we asked participants to average two tactile motion trajectories delivered simultaneously to two different fingerpads. Our results show strong integration between multiple tactile inputs, but subject to limitations for inputs delivered within a hand. As such, the present study establishes a paradigm for studying unified experience of touch despite distinct stimulus elements. [ABSTRACT FROM AUTHOR]

Proprioceptive afference can be investigated using corticokinematic coherence (CKC), which indicates coupling between limb kinematics and cortical activity. CKC has been quantified using proprioceptive stimulation (movement actuators) with fixed interstimulus interval (ISI). However, it is unclear how regularity of the stimulus sequence (jitter) affects CKC strength. Eighteen healthy volunteers (16 right-handed, 27.8 ± 5.0 yr, 7 females) participated in magnetoencephalography (MEG) session in which their right index finger was continuously moved at ∼3 Hz with Constant 333 ms ISI or with 20% Jitter (ISI 333 ± 66 ms) using a pneumatic-movement actuator. Three minutes of data per condition were collected. Finger kinematics were recorded with a three-axis accelerometer. CKC strength was defined as the peak coherence value in the Rolandic MEG gradiometer pair contralateral to the movement at 3 Hz. Both conditions resulted in significant coherence peaking in the gradiometers over the primary sensorimotor cortex. Constant stimulation yielded stronger CKC at 3 Hz (0.78 ± 0.11 vs. 0.66 ± 0.13, P < 0.001) and its first harmonic (0.60 ± 0.19 vs. 0.27 ± 0.11, P < 0.001) than irregular stimulation. Similarly, the respective sustained-movement evoked field was also stronger for constant stimulation. The results emphasize the importance of temporal stability of the proprioceptive stimulation sequence when quantifying CKC strength. The weaker CKC during irregular stimulation can be explained with temporal and thus spectral scattering of the paired peripheral and cortical events beyond the mean stimulation frequency. This impairs the signal-to-noise ratio of respective MEG signal and thus CKC strength. When accurately estimating and following changes in CKC strength, we suggest using precise movement actuators with constant stimulation sequence. NEW & NOTEWORTHY Cortical proprioceptive processing can be investigated using corticokinematic coherence (CKC). The findings show that CKC method is sensitive to temporal stability in the stimulation sequence. Although both regular and irregular sequences resulted in robust coherence, the regular stimulation sequence with pneumatic movement actuator is recommended to maximize coherence strength and reproducibility to allow better comparability between groups or populations. [ABSTRACT FROM AUTHOR]

Whisker deflection evokes sparse, low-probability spiking among L2/3 pyramidal cells in rodent somatosensory cortex (S1), with spiking distributed nonuniformly between more and less responsive cells. The cellular and local circuit factors that determine whisker responsiveness across neurons are unclear. To identify these factors, we used two-photon calcium imaging and loose-seal recording to identify more and less responsive L2/3 neurons in S1 slices in vitro, during feedforward recruitment of the L2/3 network by L4 stimulation. We observed a broad gradient of spike recruitment thresholds within local L2/3 populations, with low- and high-threshold cells intermixed. This recruitment gradient was significantly correlated across different L4 stimulation sites, and between L4-evoked and whisker-evoked responses in vivo, indicating that a substantial component of responsiveness is independent of tuning to specific feedforward inputs. Low- and high-threshold L2/3 pyramidal cells differed in L4-evoked excitatory synaptic conductance and intrinsic excitability, including spike threshold and the likelihood of doublet spike bursts. A gradient of intrinsic excitability was observed across neurons. Cells that spiked most readily to L4 stimulation received the most synaptic excitation but had the lowest intrinsic excitability. Low- and high-threshold cells did not differ in dendritic morphology, passive membrane properties, or L4-evoked inhibitory conductance. Thus multiple gradients of physiological properties exist across L2/3 pyramidal cells, with excitatory synaptic input strength best predicting overall spiking responsiveness during network recruitment.

Corticokinematic coherence (CKC) is the phase coupling between limb kinematics and cortical neurophysiological signals, reflecting cortical processing of proprioceptive afference, and it is reproducible when estimated with magnetoencephalography (MEG). However, feasibility and reproducibility of CKC based on electroencephalography (EEG) is still unclear and is the primary object of the present report. Thirteen healthy right-handed volunteers (seven females, 21.7 ± 4.3 yr) participated in two combined MEG/EEG sessions 12.6 ± 1.3 mo apart. Participants' dominant and nondominant index finger was continuously moved at 3 Hz for 4 min separately using a pneumatic-movement actuator. Coherence was computed between finger acceleration and three derivations of EEG signals: 1) average reference, 2) bipolar derivations, and 3) surface Laplacian. CKC strength was defined as the peak coherence value at movement frequency. Intraclass-correlation coefficient values (0.74-0.93) indicated excellent intersession reproducibility for CKC strength for all derivations and moved fingers. CKC strength obtained with EEG was approximately two times lower compared with MEG, but the values were positively correlated across the participants. CKC strength was significantly (P < 0.01) higher for bipolar (session 1: 0.19 ± 0.09; session 2: 0.20 ± 0.10) and surface Laplacian (session 1: 0.22 ± 0.09; session 2: 0.21 ± 0.09) derivations than for the average reference (session 1: 0.10 ± 0.04; session 2: 0.11 ± 0.05). We demonstrated that CKC is a feasible and reproducible tool to monitor proprioception using EEG recordings, although the strength of CKC was twice lower for EEG compared with MEG. Laplacian and bipolar (CP3-C1/CP3-C3 and CP4-C2/C4-FC2) EEG derivation(s) are recommended for future research and clinical use of CKC method. NEW & NOTEWORTHY The most important message in this report is that the corticokinematic coherence (CKC) method is a feasible and reproducible tool to quantify, map, and follow cortical proprioceptive ("the movement sense") processing using EEG that is more widely available for CKC recordings than previously used magnetoencephalography designs, in basic research, but especially in clinical environments. We provide useful recommendations for optimal EEG derivations for cost-effective experimental designs, making it possible to scale up in sample size in future studies. [ABSTRACT FROM AUTHOR]

In a prior study using laser scanning photostimulation, we found a pronounced cell type-specific mediolateral asymmetry in the local synaptic connectivity in the superficial laminae of the spinal dorsal horn (Kosugi M, Kato G, Lukashov S, Pendse G, Puskar Z, Kozsurek M, Strassman AM. J Physiol 591: 1935-1949, 2013). To obtain information on dorsal horn organization that might complement findings from microelectrode studies, voltage-sensitive dye imaging was used in the present study to examine patterns of activity evoked by focal electrical stimulation, in the presence and absence of synaptic blocking agents, at different positions in transverse, parasagittal, and horizontal slices of the dorsal horn of 2- to 3-wk -old male rats. A pronounced difference in responsiveness was found between medial and lateral dorsal horn, in that medial sites in the superficial dorsal horn showed much larger synaptic responses to focal stimulation than lateral sites. This difference appeared to be a result of a difference in the intrinsic elements of the dorsal horn, rather than a difference in the inputs from the white matter, because the stimulus intensities were subthreshold for evoking synaptic responses from stimulation at sites in the white matter, although it is also possible that the greater responsiveness is due, at least in part, to activation of Aβ primary afferent fibers that pass through the medial dorsal horn. The results raise the possibility of differences between medial and dorsal horn that need to be taken into account in the interpretation of studies of dorsal horn organization. NEW & NOTEWORTHY We used voltage-sensitive dye imaging to obtain information on spatial aspects of dorsal horn organization that are difficult to examine with single-cell approaches because of the limitations of microelectrode sampling. The most noteworthy finding was a previously unreported, extreme difference between medial and lateral dorsal horn in responsiveness to focal stimulation that appears to result, at least in part, from a greater degree of excitability or local connectivity in medial dorsal horn. [ABSTRACT FROM AUTHOR]

Bodies change continuously, but we do not know if and how these changes affect somatosensory cortex. We address this issue in the whisker-barrel-cortex-pathway. We ask how outgrowing whiskers are mapped onto layer 4 barrel neuron responses. Half of whisker follicles contained dual whiskers, a shorter presumably outgrowing whisker (referred to as young whisker) and a longer one (referred to as old whisker). Young whiskers were much thinner than old ones but were inserted more deeply into the whisker follicle. Both whiskers were embedded in one outer root sheath surrounded by a common set of afferent nerve fibers. We juxtacellularly identified layer 4 barrel neurons representing dual whiskers with variable whisker length differences in anesthetized rats. Strength and latency of neuronal responses were strongly correlated for deflections of young and old whiskers but were not correlated with whisker length. The direction preferences of young and old whiskers were more similar than expected by chance. Old whiskers evoked marginally stronger and slightly shorter latency spike and local field potential responses than young whiskers. Our data suggest a conservative rewiring mechanism, which connects young whiskers to existing peripheral sensors. The fact that layer 4 barrel neurons retain their response properties is remarkable given the different length, thickness, and insertion depth of young and old whiskers. Retention of cortical response properties might be related to the placement of young and old whisker in one common outer root sheath and may contribute to perceptual stability across whisker replacement. [ABSTRACT FROM AUTHOR]

The present case study details sensations elicited by electrical stimulation of peripheral nerve axons using an implanted nerve cuff electrode, in a participant with a transhumeral amputation. The participant uses an osseointegrated electromechanical interface, which enables skeletal attachment of the prosthesis and long-term, stable, bidirectional communication between the implanted electrodes and prosthetic arm. We focused on evoking somatosensory percepts, where we tracked and quantified the evolution of perceived sensations in the missing hand, which were evoked from electrical stimulation of the nerve, for over 2 yr. These sensations included small, pointlike areas of either vibration or pushing, to larger sensations over wider areas, indicating the recruitment of a few and many afferents, respectively. Furthermore, we used a two-alternative forced choice paradigm to measure the level of discrimination between trains of brief electrical stimuli, to gauge what the participant could reliably distinguish between. At best, the participant was able to distinguish a 0.5-Hz difference and on average acquired a 3.8-Hz just-noticeable difference at a more stringent psychophysical level. The current work shows the feasibility for long-term sensory feedback in prostheses, via electrical axonal stimulation, where small and relatively stable percepts were felt that may be used to deliver graded sensory feedback. This opens up opportunities for signaling feedback during movements (e.g., for precision grip), but also for conveying more complex cutaneous sensations, such as texture. NEW & NOTEWORTHY We demonstrate the long-term stability and generation of sensations from electrical peripheral nerve stimulation in an amputee, through an osseointegrated implant. We find that perceived tactilelike sensations could be generated for over 2 yr, in the missing hand. This is useful for prosthetic development and the implementation of feedback in artificial body parts. [ABSTRACT FROM AUTHOR]

Roughness is the most salient perceptual dimension of surface texture but has no well-defined physical basis. We seek to determine the neural determinants of tactile roughness in the somatosensory nerves. Specifically, we record the patterns of activation evoked in tactile nerve fibers of anesthetized Rhesus macaques to a large and diverse set of natural textures and assess what aspect of these patterns of activation can account for psychophysical judgments of roughness, obtained from human observers. We show that perceived roughness is determined by the variation in the population response, weighted by fiber type. That is, a surface will feel rough to the extent that the activity varies across nerve fibers and varies across time within nerve fibers. We show that this variation-based neural code can account not only for magnitude estimates of roughness but also for roughness discrimination performance. NEW & NOTEWORTHY Our sense of touch endows us with an exquisite sensitivity to the microstructure of surfaces, the most salient aspect of which is roughness. We analyze the responses evoked in tactile fibers of monkeys by natural textures and compare them to judgments of roughness obtained for the same textures from human observers. We then describe how texture signals from three populations of nerve fibers are integrated to culminate in a percept of roughness. [ABSTRACT FROM AUTHOR]

Attention-deficit hyperactivity disorder (ADHD) is characterized by an inability to concentrate, heightened activity, and hypermotoric behavior, but sensory (e.g., tactile) problems are common. The literature on tactile impairments in ADHD is limited, with most work employing clinical observations or questionnaires. We studied tactile processing in children with ADHD and hypothesized that children with ADHD would show reduced performance in tasks closely linked to inhibition. Sixty-seven children with ADHD and 62 typically developing children (TDC) performed a battery of tasks grouped in domains: simple and choice reaction time; static and dynamic detection threshold (probing feedforward inhibition); amplitude discrimination without adaptation and with dual and single-site adaptation (probing lateral inhibition and adaptation); sequential and simultaneous frequency discrimination (previously linked to GABA); and temporal order judgment with and without a synchronous carrier stimulus. Children with ADHD could discriminate different amplitudes without adaptation, suggesting lateral inhibition is intact, but were negatively affected in all adaptation conditions, whereas TDC were only affected during single-site adaptation. Children with ADHD also showed normal frequency discrimination. Children with ADHD showed slower reaction times and higher detection threshold, likely driven by IQ and inattention, because reaction time and detection thresholds correlated with IQ and subtle motor signs. Children with ADHD showed a pattern of altered tactile processing on specific tasks, suggesting that higher cognitive function and cortical mechanisms related to adaptation are affected in ADHD, but no clear conclusion can be drawn toward impaired inhibition. NEW & NOTEWORTHY This manuscript presents the first tactile psychophysical study testing different aspects of tactile processing in attention-deficit hyperactivity disorder (ADHD), using large cohort sizes of 67 children with ADHD and 65 Typically Developing Children. This study demonstrates impaired tactile processing in children with ADHD, on some, but not all tasks (showing this is not just due to attention), related to impaired cortical mechanisms. Furthermore, both IQ and soft motor skill abnormalities (common in ADHD) are correlated with tactile abnormalities. [ABSTRACT FROM AUTHOR]

The bAP-Ca2+ signal amplitudes in some apical tuft branches randomly vary from moment to moment. In repetitive measurements, successful AP invasions are followed by complete failures. Passive spread of voltage from the apical trunk into the tuft occasionally reaches the threshold for local Na+ spike, resulting in stronger Ca2+ influx. During a burst of three somatic APs, the peak of dendritic Ca2+ in the apical tuft occurs with a delay of 20-50 ms depending on AP frequency. In cortical pyramidal neurons, backpropagating action potentials (bAPs) supply Ca2+ to synaptic contacts on dendrites. To determine whether the efficacy of AP backpropagation into apical tuft dendrites is stable over time, we performed dendritic Ca2+ and voltage imaging in rat brain slices. We found that the amplitude of bAP-Ca2+ in apical tuft branches was unstable, given that it varied from trial to trial (termed “bAP-Ca2+ flickering”). Small perturbations in dendritic physiology, such as spontaneous synaptic inputs, channel inactivation, or temperature-induced changes in channel kinetics, can cause bAP flickering. In the tuft branches, the density of Na+ and K+ channels was sufficient to support local initiation of fast spikelets by glutamate iontophoresis. We quantified the time delay between the somatic AP burst and the peak of dendritic Ca2+ transient in the apical tuft, because this delay is important for induction of spike-timing dependent plasticity. Depending on the frequency of the somatic AP triplets, Ca2+ signals peaked in the apical tuft 20–50 ms after the 1st AP in the soma. Interestingly, at low frequency (<20 Hz), the Ca2+ peaked sooner than at high frequency, because only the 1st AP invaded tuft. Activation of dendritic voltage-gated Ca2+ channels is sensitive to the duration of the dendritic voltage transient. In apical tuft branches, small changes in the duration of bAP voltage waveforms cause disproportionately large increases in dendritic Ca2+ influx (bAP-Ca2+ flickering). The stochastic nature of bAP-Ca2+ adds a new perspective on the mechanisms by which pyramidal neurons combine inputs arriving at different cortical layers. NEW & NOTEWORTHY The bAP-Ca2+ signal amplitudes in some apical tuft branches randomly vary from moment to moment. In repetitive measurements, successful AP invasions are followed by complete failures. Passive spread of voltage from the apical trunk into the tuft occasionally reaches the threshold for local Na+ spike, resulting in stronger Ca2+ influx. During a burst of three somatic APs, the peak of dendritic Ca2+ in the apical tuft occurs with a delay of 20-50 ms depending on AP frequency. [ABSTRACT FROM AUTHOR]

Neuronal networks within the spinal cord directly control rhythmic movements of the arms/forelimbs and legs/hindlimbs during locomotion in mammals. For an effective locomotion, these networks must be flexibly coordinated to allow for various gait patterns and independent use of the arms/forelimbs. This coordination can be accomplished by mechanisms intrinsic to the spinal cord, somatosensory feedback from the limbs, and various supraspinal pathways. Incomplete spinal cord injury disrupts some of the pathways and structures involved in interlimb coordination, often leading to a disruption in the coordination between the arms/forelimbs and legs/hindlimbs in animal models and in humans. However, experimental spinal lesions in animal models to uncover the mechanisms coordinating the limbs have limitations due to compensatory mechanisms and strategies, redundant systems of control, and plasticity within remaining circuits. The purpose of this review is to provide a general overview and critical discussion of experimental studies that have investigated the neural mechanisms involved in coordinating the arms/forelimbs and legs/hindlimbs during mammalian locomotion. [ABSTRACT FROM AUTHOR]

Supragranular layers of sensory cortex are known to exhibit sparse firing. In rodent vibrissal cortex, a small fraction of neurons in layer 2 and 3 (L2/3) respond to whisker stimulation. In this study, we combined whole cell recording and two-photon imaging in anesthetized mice and quantified the synaptic response and spiking profile of L2/3 neurons. Previous literature has shown that neurons across layers of vibrissal cortex are tuned to the velocity of whisker movement. We therefore used a broad range of stimuli that included the standard range of velocities (0 –1.2 deg/ms) and extended to a “sharp” high-velocity deflection (3.8 deg/ms). Consistent with previous literature, whole cell recording revealed a sparse response to the standard range of velocities: although all recorded cells showed tuning to velocity in their postsynaptic potentials, only a small fraction produced stimulus-evoked spikes. In contrast, the sharp stimulus evoked reliable spiking in the majority of neurons. The action potential threshold of spikes evoked by the sharp stimulus was significantly lower than that of the spontaneous spikes. Juxtacellular recordings confirmed that application of sharp stimulus to single or multiple whiskers produced temporally precise spiking with minimal trial-to-trial spike count variability (Fano factors equal or close to the theoretical minimum). Two-photon imaging further confirmed that most neurons that were not responsive to the standard deflections responded to the sharp stimulus. Altogether, our results indicate that sparseness in L2/3 cortex depends on the choice of stimulus: strong single- or multiwhisker stimulation can induce the transition from sparse to “dense” population response. [ABSTRACT FROM AUTHOR]

Our ability to process temporal frequency information by touch underlies our capacity to perceive and discriminate surface textures. Auditory signals, which also provide extensive temporal frequency information, can systematically alter the perception of vibrations on the hand. How auditory signals shape tactile processing is unclear; perceptual interactions between contemporaneous sounds and vibrations are consistent with multiple neural mechanisms. Here we used a crossmodal adaptation paradigm, which separated auditory and tactile stimulation in time, to test the hypothesis that tactile frequency perception depends on neural circuits that also process auditory frequency. We reasoned that auditory adaptation effects would transfer to touch only if signals from both senses converge on common representations. We found that auditory adaptation can improve tactile frequency discrimination thresholds. This occurred only when adaptor and test frequencies overlapped. In contrast, auditory adaptation did not influence tactile intensity judgments. Thus auditory adaptation enhances touch in a frequency- and feature-specific manner. A simple network model in which tactile frequency information is decoded from sensory neurons that are susceptible to auditory adaptation recapitulates these behavioral results. Our results imply that the neural circuits supporting tactile frequency perception also process auditory signals. This finding is consistent with the notion of supramodal operators performing canonical operations, like temporal frequency processing, regardless of input modality. [ABSTRACT FROM AUTHOR]

Imperceptible tactile noise applied to the skin of the feet enhances posture-correcting cutaneous reflexes. This sensory augmentation technique, stochastic resonance (SR), has not been tested in the less-sensitive hairy skin of the leg for its reflex-enhancement ability. The objectives of this study were to determine whether calf skin stimulation produces cutaneous reflexes and whether noise can modify the reflex. In 20 participants, electrotactile pulse trains were applied at the calf while participants performed submaximal isometric knee extension. To test SR, five different levels of vibrotactile noise were applied simultaneously to the test input. Muscle activity from the vastus lateralis (VL) was analyzed 60-110 ms after stimulation. Reflex ratios were calculated by dividing the reflex peak activity by the prestimulation background muscle activity. A significant reflex response was evoked in 16/20 participants (5.41 ± 2.6% of background muscle activity); these responses varied between individuals with eight being facilitatory and eight being inhibitory. In half of the participants, a new reflex appeared at some level of added noise ( n = 10). The average reflex ratio of the study population was significantly higher at the "optimal" noise level (8.61 ± 4.5) than at "baseline" (4.70 ± 5.6) ( P = 0.002); the optimal level varied across participants. These results suggest that cutaneous reflexes exist at the VL in response to calf skin stimulation and that SR can change cutaneous reflexes at the leg. This study provides an important first step toward SR application in clinical populations with sensory loss such as individuals with lower extremity amputation. NEW & NOTEWORTHY Our work showed that cutaneous reflexes, known to be present in response to foot sole stimulation, can also be evoked by stimulation of hairy leg skin. In addition, we demonstrated that adding tactile noise can enhance this reflex response. These findings demonstrate proof-of-concept for potential future applications where tactile stimulation, applied to the leg of an individual with amputation, can enhance postural-relevant reflexes. Improving postural control may reduce the risk of falls in this high-risk population.

Vestibular information is known to be important for postural stability on tilting surfaces, but the relative importance of vestibular information across a wide range of surface tilt velocities is less clear. We compared how tilt velocity influences postural orientation and stability in nine subjects with bilateral vestibular loss and nine age-matched, control subjects. Subjects stood on a force platform that tilted 6 deg, toes-up at eight velocities (0.25 to 32 deg/s), with and without vision. Results showed that visual information effectively compensated for lack of vestibular information at all tilt velocities. However, with eyes closed, subjects with vestibular loss were most unstable within a critical tilt velocity range of 2 to 8 deg/s. Subjects with vestibular deficiency lost their balance in more than 90% of trials during the 4 deg/s condition, but never fell during slower tilts (0.25-1 deg/s) and fell only very rarely during faster tilts (16-32 deg/s). At the critical velocity range in which falls occurred, the body center of mass stayed aligned with respect to the surface, onset of ankle dorsiflexion was delayed, and there was delayed or absent gastrocnemius inhibition, suggesting that subjects were attempting to actively align their upper bodies with respect to the moving surface instead of to gravity. Vestibular information may be critical for stability at velocities of 2 to 8 deg/s because postural sway above 2 deg/s may be too fast to elicit stabilizing responses through the graviceptive somatosensory system, and postural sway below 8 deg/s may be too slow for somatosensory-triggered responses or passive stabilization from trunk inertia. [ABSTRACT FROM AUTHOR]

The dorsal column nuclei (DCN) of the brain stem contain secondary afferent neurons, which process ascending somatosensory information. Most of the known physiology of the DCN in primates has been acquired in acute experiments with anesthetized animals. Here, we developed a technique to implant a multielectrode array (MEA) chronically in the DCN of macaque monkeys to enable experiments with the animals awake. Two monkeys were implanted with brain-stem MEAs for 2-5 mo with no major adverse effects. Responses of the cuneate and gracile nuclei were quantified at the level of both field potentials and single units. Tactile receptive fields (RFs) were identified for 315 single units. A subset of these units had very regular spiking patterns with spike frequencies predominantly in the alpha band (8-14 Hz). The stability of the neuronal recordings was assessed with a novel analysis that identified units by their mean spike waveform and by the spike-triggered average of activity on all other electrodes in the array. Fifty-six identified neurons were observed over two or more sessions and in a few cases for as long as 1 mo. RFs of stable neurons were largely consistent across days. The results demonstrate that a chronic DCN implant in a macaque can be safe and effective, yielding high-quality unit recording for several months. The unprecedented access to these nuclei in awake primates should lead to a better understanding of their role in sensorimotor behavior. [ABSTRACT FROM AUTHOR]

In the rodent whisker system, a key model for neural processing and behavioral choices during active sensing, whisker motion is increasingly recognized as only part of a broader motor repertoire employed by rodents during active touch. In particular, recent studies suggest whisker and head motions are tightly coordinated. However, conditions governing the selection and temporal organization of such coordinated sensing strategies remain poorly understood. We videographically reconstructed head and whisker motions of freely moving mice searching for a randomly located rewarded aperture, focusing on trials in which animals appeared to rapidly "correct" their trajectory under tactile guidance. Mice orienting after unilateral contact repositioned their whiskers similarly to previously reported head-turning asymmetry. However, whisker repositioning preceded head turn onsets and was not bilaterally symmetric. Moreover, mice selectively employed a strategy we term contact maintenance, with whisking modulated to counteract head motion and facilitate repeated contacts on subsequent whisks. Significantly, contact maintenance was not observed following initial contact with an aperture boundary, when the mouse needed to make a large corrective head motion to the front of the aperture, but only following contact by the same whisker field with the opposite aperture boundary, when the mouse needed to precisely align its head with the reward spout. Together these results suggest that mice can select from a diverse range of sensing strategies incorporating both knowledge of the task and whisk-by-whisk sensory information and, moreover, suggest the existence of high level control (not solely reflexive) of sensing motions coordinated between multiple body parts. [ABSTRACT FROM AUTHOR]

The ability to resolve the orientation of edges is crucial to daily tactile and sensorimotor function, yet the means by which edge perception occurs is not well understood. Primate cortical area 3b neurons have diverse receptive field (RF) spatial structures that may participate in edge orientation perception. We evaluated five candidate RF models for macaque area 3b neurons, previously recorded while an oriented bar contacted the monkey's fingertip. We used a Bayesian classifier to assign each neuron a best-fit RF structure. We generated predictions for human performance by implementing an ideal observer that optimally decoded stimulus-evoked spike counts in the model neurons. The ideal observer predicted a saturating reduction in bar orientation discrimination threshold with increasing bar length. We tested 24 humans on an automated, precision-controlled bar orientation discrimination task and observed performance consistent with that predicted. We next queried the ideal observer to discover the RF structure and number of cortical neurons that best matched each participant's performance. Human perception was matched with a median of 24 model neurons firing throughout a 1-s period. The 10 lowest-performing participants were fit with RFs lacking inhibitory sidebands, whereas 12 of the 14 higher-performing participants were fit with RFs containing inhibitory sidebands. Participants whose discrimination improved as bar length increased to 10 mm were fit with longer RFs; those who performed well on the 2-mm bar, with narrower RFs. These results suggest plausible RF features and computational strategies underlying tactile spatial perception and may have implications for perceptual learning. [ABSTRACT FROM AUTHOR]

Central auditory circuits are influenced by the somatosensory system, a relationship that may underlie tinnitus generation. In the guinea pig dorsal cochlear nucleus (DCN), pairing spinal trigeminal nucleus (Sp5) stimulation with tones at specific intervals and orders facilitated or suppressed subsequent tone-evoked neural responses, reflecting spike timing-dependent plasticity (STDP). Furthermore, after noiseinduced tinnitus, bimodal responses in DCN were shifted from Hebbian to anti-Hebbian timing rules with less discrete temporal windows, suggesting a role for bimodal plasticity in tinnitus. Here, we aimed to determine if multisensory STDP principles like those in DCN also exist in primary auditory cortex (A1), and whether they change following noise-induced tinnitus. Tone-evoked and spontaneous neural responses were recorded before and 15 min after bimodal stimulation in which the intervals and orders of auditory-somatosensory stimuli were randomized. Tone-evoked and spontaneous firing rates were influenced by the interval and order of the bimodal stimuli, and in sham-controls Hebbian-like timing rules predominated as was seen in DCN. In noise-exposed animals with and without tinnitus, timing rules shifted away from those found in sham-controls to more anti-Hebbian rules. Only those animals with evidence of tinnitus showed increased spontaneous firing rates, a purported neurophysiological correlate of tinnitus in A1. Together, these findings suggest that bimodal plasticity is also evident in A1 following noise damage and may have implications for tinnitus generation and therapeutic intervention across the central auditory circuit. [ABSTRACT FROM AUTHOR]

Measuring the excitability of individual axons is complicated by the prohibitive difficulty in obtaining intracellular recordings. Here, we present an innovative methodology that enables local excitability to be measured anywhere in a channelrhodopsin (ChR2)-expressing neuron. The approach hinges on activating ChR2 in a spatially and temporally precise manner while recording the resulting spike train from a remote site. We validated this approach in primary afferent neurons (PANs). Initial encoding of somatosensory stimuli relies on transduction of the physical stimulus into a receptor potential and transformation of the receptor potential into a spike train; the transformation process depends on the excitability of the most distal PAN endings but, as explained above, is extraordinarily difficult to study in situ using traditional methods. Using ChR2-based photoactivation, we show 1) that excitability differs between the distal endings and more proximal portions of PAN axons, 2) that the transformation process differs between PANs, and 3) that the transformation process is directly affected by inflammation. Beyond presenting an innovative method by which to study axonal excitability, this study has validated its utility in helping to decipher the earliest stages of somatosensory encoding. [ABSTRACT FROM AUTHOR]

The function of rodents' whisker somatosensory system is to transform tactile cues, in the form of vibrissa vibrations, into neuronal responses. It is well established that rodents can detect numerous tactile stimuli and tell them apart. However, the transformation of tactile stimuli obtained through whisker movements to neuronal responses is not well-understood. Here we examine the role of whisker velocity in tactile information transmission and its coding mechanisms. We show that in anaesthetized rats, whisker velocity is related to the radial distance of the object contacted and its own velocity. Whisker velocity is accurately and reliably coded in first-order neurons in parallel, by both the relative time interval between velocity-independent first spike latency of rapidly adapting neurons and velocity-dependent first spike latency of slowly adapting neurons. At the same time, whisker velocity is also coded, although less robustly, by the firing rates of slowly adapting neurons. Comparing first- and second-order neurons, we find similar decoding efficiencies for whisker velocity using either temporal or rate-based methods. Both coding schemes are sufficiently robust and hardly affected by neuronal noise. Our results suggest that whisker kinematic variables are coded by two parallel coding schemes and are disseminated in a similar way through various brain stem nuclei to multiple brain areas. [ABSTRACT FROM AUTHOR]

The brain's ability to integrate information from different senses is acquired only after extensive sensory experience. However, whether early life experience instantiates a general integrative capacity in multisensory neurons or one limited to the particular cross-modal stimulus combinations to which one has been exposed is not known. By selectively restricting either visual-nonvisual or auditory-nonauditory experience during the first few months of life, the present study found that trisensory neurons in cat superior colliculus (as well as their bisensory counterparts) became adapted to the cross-modal stimulus combinations specific to each rearing environment. Thus, even at maturity, trisensory neurons did not integrate all cross-modal stimulus combinations to which they were capable of responding, but only those that had been linked via experience to constitute a coherent spatiotemporal event. This selective maturational process determines which environmental events will become the most effective targets for superior colliculus-mediated shifts of attention and orientation. [ABSTRACT FROM AUTHOR]

The goal of this review is to start to consolidate and distill the substantial body of research that comprises the published work of the late Professor Steven S. Hsiao. The studies of Hsiao began by demonstrating the receptive field properties of somatosensory neurons, progressed to describing cortical feature selectivity, and then eventually elevated the field to hopes of tapping into natural neural codes with artificial somatosensory feedback. With ongoing analogies to contemporaneous studies in visual neuroscience, the research results and writings of Hsiao have provided the fields of haptics and somatosensory neurophysiology with the conceptual tools needed to allow profound progress. Specifically, Hsiao suggested that slowly adapting tactile form perception could be restored with cortical microstimulation, rapidly adapting slip reflexes should be relegated to low-level, hard-wired prosthetic components, and Pacinian-corpuscle spatiotemporal population responses could potentially be decoded/encoded to provide information about interactions of hands and hand-held instruments with external objects. Future studies will be guided by these insightful reports from Hsiao. [ABSTRACT FROM AUTHOR]

We routinely identify objects with our hands, and the physical attributes of touched objects are often held in short-term memory to aid future decisions. However, the brain structures that selectively process tactile information to encode object shape are not fully identified. In this article we describe the areas within the human cerebral cortex that specialize in encoding, short-term memory, and decision-making related to the shape of objects explored with the hand. We performed event-related functional magnetic resonance imaging in subjects performing a shape discrimination task in which two sequentially presented objects had to be explored to determine whether they had the same shape or not. To control for low-level and nonspecific brain activations, subjects performed a temperature discrimination task in which they compared the temperature of two spheres. Our results show that although a large network of brain structures is engaged in somatosensory processing, it is the areas lining the intraparietal sulcus that selectively participate in encoding, maintaining, and deciding on tactile information related to the shape of objects. [ABSTRACT FROM AUTHOR]

Here, we investigate the neural mechanisms of detecting lumps embedded in artificial compliant tissues. We performed a combined psychophysical study of humans performing a passive lump detection task with a neurophysiological study in nonhuman primates (Macaca mulatta) where we recorded the responses of peripheral mechanoreceptive afferents to lumps embedded at various depths in intermediates (rubbers) of varying compliance. The psychophysical results reveal that human lump detection is greatly degraded by both lump depth and decreased compliance of the intermediate. The neurophysiology results reveal that only the slowly adapting type 1 (SA1) afferents provide a clear spatial representation of lumps at all depths and that the representation is affected by lump size, depth, and compliance of the intermediate. The rapidly adapting afferents are considerably less sensitive to the lump. We defined eight neural response measures that we hypothesized could explain the psychophysical behavior, including peak firing rate, spatial spread of neural activity, and additional parameters derived from these measures. We find that peak firing rate encodes the depth of the lump, and the neural spatial spread of the SA1 response encodes for lump size but not lump shape. We also find that the perception of lump size may be affected by the compliance of the intermediate. The results show that lump detection is based on a spatial population code of the SA1 afferents, which is distorted by the depth of the lump and compliance of the tissue. [ABSTRACT FROM AUTHOR]

Impaired responses to tactile stimulation are a commonly reported symptom among children with autism spectrum disorder (ASD). Furthermore, impairments in filtering or habituation to tactile input have been described in ASD. This study measured different aspects of tactile processing to investigate atypical touch sensitivity in children with ASD, methodology that has not been previously used in this population. Sixty-seven typically developing children (TDC) and 32 children with ASD (ages 8-12) completed vibrotactile tasks assessing: reaction time (RT); static and dynamic detection threshold (DT); amplitude discrimination with and without single-site adaptation; frequency discrimination; and temporal order judgment (TOJ) with and without concurrent stimulation. Children with ASD showed raised static detection thresholds and an absence of the effect of a dynamically increasing subthreshold stimulus on static detection threshold. Children with ASD also showed poorer amplitude discrimination than TDC, as well as decreased adaptation. There were no significant differences in frequency discrimination or TOJ performance between the groups. Differences in the effect of dynamic stimulation on detection threshold suggest impaired feed-forward inhibition in autism, which may be linked to poor sensory filtering. Increased baseline amplitude discrimination thresholds in ASD suggest that lateral inhibitory connections are weaker in ASD, and an absence of the effect of adaptation suggests impaired modulation of lateral inhibitory connections in ASD, which may relate to aberrant habituation. These results suggest a functional deficit in the somatosensory inhibitory system in autism. Understanding the specific mechanisms underlying sensory symptoms in autism may allow for more specific therapeutic or drug targeting in the near future. [ABSTRACT FROM AUTHOR]

We made simultaneous bilateral recordings of unit activity in the nucleus ventroposterior lateralis (VPL) in intact rats and after acute and chronic left thoracic hemisection. We observed an immediate bilateral decline in multireceptive units, reflecting a loss of nociceptive input on the lesion side and a loss of low-threshold inputs contralaterally. Unit properties were restored to normal by 6 wk. Mean spontaneous discharge frequency remained unchanged in left VPL at all intervals. Right VPL displayed a substantial increase in spontaneous discharge frequency at 2 and 4 wk, returning to normal by 6 wk. Activity in left VPL driven by Pinch or Brush of the right limb was unchanged except for an immediate decrease in the response to Pinch, which was reversed by 2 wk despite persistent left hemisection. In right VPL, the response to Pinch or Brush of the left hindlimb was enhanced at 2 and 4 wk but returned to normal by 6 wk. Behaviorally, the same rats displayed increased sensitivity to mechanical stimulation of the left hindlimb, but, unlike VPL activity, there was no significant behavioral recovery. Bursting cells were also observed bilaterally in VPL, but this did not match the restriction of scratches to the hindlimb contralateral to the hemisection considered to be evidence for neuropathic pain. The novel findings include recovery of responsiveness to Pinch on the side ipsilateral to the hemisection despite the lack of spinothalamic input as well as failure for the thalamus contralateral to hemisection to maintain its elevated responsiveness. [ABSTRACT FROM AUTHOR]

Linear receptive field (RF) models of area 3b neurons reveal a three-component structure: a central excitatory region flanked by two inhibitory regions that are spatially and temporally nonoverlapping with the excitation. Previous studies also report that there is an "infield" inhibitory region throughout the neuronal RF, which is a nonlinear interactive (second order) effect whereby stimuli lagging an input to the excitatory region are suppressed. Thus linear models may be inaccurate approximations of the neurons' true RFs. In this study, we characterize the RFs of area 3b neurons, using a second-order quadratic model. Data were collected from 80 neurons of two awake, behaving macaque monkeys while a random dot pattern was scanned simultaneously across the distal pads of digits D2, 3, and 4. We used an iterative method derived from matching pursuit to identify a set of linear and nonlinear terms with significant effects on the neuronal response. For most neurons (65/80), the linear component of the quadratic RF was characterized by a single excitatory region on the dominant digit. Interactions within the dominant digit were characterized by two quadratic filters that capture the spatial aspects of the interactive infield inhibition. Interactions between the dominant (most responsive) digit and its adjacent digit(s) formed the largest class of cross-digit interactions. The results demonstrate that a significant part of area 3b responses is due to nonlinear mechanisms, and furthermore, the data support the notion that area 3b neurons have "nonclassical RF"-like input from adjacent fingers, indicating that area 3b plays a role in integrating shape inputs across digits. [ABSTRACT FROM AUTHOR]

The dorsolateral part of the striatum (DLS) represents the initial stage for processing sensorimotor information in the basal ganglia. Although the DLS receives much of its input from the primary somatosensory (SI) cortex, peripheral somesthetic stimulation activates the DLS at latencies that are shorter than the response latencies recorded in the SI cortex. To identify the subcortical regions that transmit somesthetic information directly to the DLS, we deposited small quantities of retrograde tracers at DLS sites that displayed consistent time-locked responses to controlled whisker stimulation. The neurons that were retrogradely labeled by these injections were located mainly in the sensorimotor cortex and, to a lesser degree, in the amygdala and thalamus. Quantitative analysis of neuronal labeling in the thalamus indicated that the strongest thalamic input to the whisker-sensitive part of the DLS originates from the medial posterior nucleus (POm), a somesthetic-related region that receives inputs from the spinal trigeminal nucleus. Anterograde tracer injections in POm confirmed that this thalamic region projects to the DLS neuropil. In subsequent experiments, simultaneous recordings from POm and the DLS during whisker stimulation showed that POm consistently responds before the DLS. These results suggest that POm could transmit somesthetic information to the DLS, and this modality-specific thalamostriatal pathway may cooperate with the thalamostriatal projections that originate from the intralaminar nuclei. [ABSTRACT FROM AUTHOR]

During locomotion, motor cortical neurons projecting to the pyramidal tract (PTNs) discharge in close relation to strides. How their discharges vary based on the part of the body they influence is not well understood. We addressed this question with regard to joints of the forelimb in the cat. During simple and ladder locomotion, we compared the activity of four groups of PTNs with somatosensory receptive fields involving different forelimb joints: 1) 45 PTNs receptive to movements of shoulder, 2) 30 PTNs receptive to movements of elbow, 3) 40 PTNs receptive to movements of wrist, and 4) 30 nonresponsive PTNs. In the motor cortex, a relationship exists between the location of the source of afferent input and the target for motor output. On the basis of this relationship, we inferred the forelimb joint that a PTN influences from its somatosensory receptive field. We found that different PTNs tended to discharge differently during locomotion. During simple locomotion shoulderrelated PTNs were most active during late stance/early swing, and upon transition from simple to ladder locomotion they often increased activity and stride-related modulation while reducing discharge duration. Elbow-related PTNs were most active during late swing/early stance and typically did not change activity, modulation, or discharge duration on the ladder. Wrist-related PTNs were most active during swing and upon transition to the ladder often decreased activity and increased modulation while reducing discharge duration. These data suggest that during locomotion the motor cortex uses distinct mechanisms to control the shoulder, elbow, and wrist. [ABSTRACT FROM AUTHOR]

Somatosensory signals undergo substantial modulation in the dorsal column nuclei. We examined transmission of signals from forelimb afferents in primate cuneate and external cuneate nuclei. In anesthetized macaque monkeys, the median, ulnar, deep radial, and superficial radial nerves were electrically stimulated at 1.5-2 motor threshold with independent Poisson trains whereas extracellular recordings were made from 317 cells. Responses to peripheral stimulation included instances of both brief facilitation and long lasting suppression. A high proportion of cells (87%) responded to stimulation of two or more peripheral nerves, suggesting a large amount of convergence. Facilitated cells showed coherence with the peripheral stimulation across a broad frequency range; coherence was especially high in cells that responded with a burst of action potentials. Cells that responded with suppression also showed significant coherence, but this fell rapidly for frequencies above 25 Hz. Similar results were seen in both the main and external cuneate. When stimulation of one nerve was conditioned by a preceding nerve stimulus, the response to the second stimulus was attenuated for around 40 ms. This occurred independently of whether the first stimulus produced an initial facilitation or suppression or whether the same or a different nerve served as a conditioning stimulus. Mechanical stimulation of a receptive field suppressed responses to a second identical mechanical stimulus over a similar timescale. We conclude that the primate cuneate nucleus is capable of transmitting temporal information about stimuli with high fidelity; stimuli interact both temporally and spatially to modulate the onward transmission of information. [ABSTRACT FROM AUTHOR]

Behavioral studies have used visuo-tactile conflicts between a participant's body and a visually presented fake or virtual body to investigate the importance of bodily perception for self-consciousness (bodily self-consciousness). Illusory self-identification with a fake body and changes in tactile processing--modulation of visuo-tactile cross-modal congruency effects (CCEs)--were reported in previous findings. Although proprioceptive signals are deemed important for bodily self-consciousness, their contribution to the representation of the full body has not been studied. Here we investigated whether and how self-identification and tactile processing (CCE magnitude) could be modified by altering proprioceptive signals with 80-Hz vibrations at the legs. Participants made elevation judgments of tactile cues (while ignoring nearby lights) during synchronous and asynchronous stroking of a seen fake body. We found that proprioceptive signals during vibrations altered the magnitude of self-identification and mislocalization of touch (CCE) in a synchrony-dependent fashion: we observed an increase of self-identification and CCE magnitude during asynchronous stroking. In a second control experiment we studied whether proprioceptive signals per se, or those from the lower limbs in particular, were essential for these changes. We applied vibrations at the upper limbs (which provide no information about the position of the participant's body in space) and in this case observed no modulation of bodily self-consciousness or tactile perception. These data link proprioceptive signals from the legs that are conveyed through the dorsal column-medial lemniscal pathway to bodily self-consciousness. We discuss their integration with bodily signals from vision and touch for full-body representations. [ABSTRACT FROM AUTHOR]

The dorsolateral striatum (DLS) receives extensive projections from primary somatosensory cortex (SI), but very few studies have used somesthetic stimulation to characterize the sensory coding properties of DLS neurons. In this study, we used computer-controlled whisker deflections to characterize the extracellular responses of DLS neurons in rats lightly anesthetized with isoflurane. When multiple whiskers were synchronously deflected by rapid back-and-forth movements, whisker-sensitive neurons in the DLS responded to both directions of movement. The latency and magnitude of these neuronal responses displayed very little variation with changes in the rate (2, 5, or 8 Hz) of whisker stimulation. Simultaneous recordings in SI barrel cortex and the DLS revealed important distinctions in the neuronal responses of these serially connected brain regions. In contrast to DLS neurons, SI neurons were activated by the initial deflection of the whiskers but did not respond when the whiskers moved back to their original position. As the rate of whisker stimulation increased, SI responsiveness declined, and the latencies of the responses increased. In fact, when whiskers were deflected at 5 or 8 Hz, many neurons in the DLS responded before the SI neurons. These results and earlier anatomic findings suggest that a component of the sensory-induced response in the DLS is mediated by inputs from the thalamus. Furthermore, the lack of sensory adaptation in the DLS may represent a critical part of the neural mechanism by which the DLS encodes stimulus-response associations that trigger motor habits and other stimulus-evoked behaviors that are not contingent on rewarded outcomes. [ABSTRACT FROM AUTHOR]

The rewarding attributes of foods containing fat are associated with the increase in fat consumption, but little is known of how the complex physical and chemical properties of orally ingested fats are represented and decoded in the brain nor how this impacts feeding behavior within the population. Here, functional MRI (fMRI) is used to assess the brain response to isoviscous, isosweet fat emulsions of increasing fat concentration and to investigate the correlation of behavioral and neuroimaging responses with taster status (TS). Cortical areas activated in response to fat, and those areas positively correlated with fat concentration, were identified. Significant responses that positively correlated with increasing fat concentration were found in the anterior insula, frontal operculum and secondary somatosensory cortex (SII), anterior cingulate cortex, and amygdala. Assessing the effect of TS revealed a strong correlation with self-reported preference of the samples and with cortical response in somatosensory areas [primary somatosensory cortex (SI), SII, and midinsula] and the primary taste area (anterior insula) and a trend in reward areas (amygdala and orbitofrontal cortex). This finding of a strong correlation with TS in somatosensory areas supports the theory of increased mechanosensory trigeminal innervation in high 6-n-propyl-2-thiouracil (PROP) tasters and has been linked to a higher risk of obesity. The interindividual differences in blood oxygenation level-dependent (BOLD) amplitude with TS indicates that segmenting populations by TS will reduce the heterogeneity of BOLD responses, improving signal detection power. [ABSTRACT FROM AUTHOR]

In a prior study using laser scanning photostimulation, we found a pronounced cell type-specific mediolateral asymmetry in the local synaptic connectivity in the superficial laminae of the spinal dorsal horn (Kosugi M, Kato G, Lukashov S, Pendse G, Puskar Z, Kozsurek M, Strassman AM. J Physiol 591: 1935-1949, 2013). To obtain information on dorsal horn organization that might complement findings from microelectrode studies, voltage-sensitive dye imaging was used in the present study to examine patterns of activity evoked by focal electrical stimulation, in the presence and absence of synaptic blocking agents, at different positions in transverse, parasagittal, and horizontal slices of the dorsal horn of 2- to 3-wk -old male rats. A pronounced difference in responsiveness was found between medial and lateral dorsal horn, in that medial sites in the superficial dorsal horn showed much larger synaptic responses to focal stimulation than lateral sites. This difference appeared to be a result of a difference in the intrinsic elements of the dorsal horn, rather than a difference in the inputs from the white matter, because the stimulus intensities were subthreshold for evoking synaptic responses from stimulation at sites in the white matter, although it is also possible that the greater responsiveness is due, at least in part, to activation of Aβ primary afferent fibers that pass through the medial dorsal horn. The results raise the possibility of differences between medial and dorsal horn that need to be taken into account in the interpretation of studies of dorsal horn organization. NEW & NOTEWORTHY We used voltage-sensitive dye imaging to obtain information on spatial aspects of dorsal horn organization that are difficult to examine with single-cell approaches because of the limitations of microelectrode sampling. The most noteworthy finding was a previously unreported, extreme difference between medial and lateral dorsal horn in responsiveness to focal stimulation that appears to result, at least in part, from a greater degree of excitability or local connectivity in medial dorsal horn.

Measuring the excitability of individual axons is complicated by the prohibitive difficulty in obtaining intracellular recordings. Here, we present an innovative methodology that enables local excitability to be measured anywhere in a channelrhodopsin (ChR2)-expressing neuron. The approach hinges on activating ChR2 in a spatially and temporally precise manner while recording the resulting spike train from a remote site. We validated this approach in primary afferent neurons (PANs). Initial encoding of somatosensory stimuli relies on transduction of the physical stimulus into a receptor potential and transformation of the receptor potential into a spike train; the transformation process depends on the excitability of the most distal PAN endings but, as explained above, is extraordinarily difficult to study in situ using traditional methods. Using ChR2-based photoactivation, we show 1) that excitability differs between the distal endings and more proximal portions of PAN axons, 2) that the transformation process differs between PANs, and 3) that the transformation process is directly affected by inflammation. Beyond presenting an innovative method by which to study axonal excitability, this study has validated its utility in helping to decipher the earliest stages of somatosensory encoding., (Copyright © 2015 the American Physiological Society.)

The brain's ability to integrate information from different senses is acquired only after extensive sensory experience. However, whether early life experience instantiates a general integrative capacity in multisensory neurons or one limited to the particular cross-modal stimulus combinations to which one has been exposed is not known. By selectively restricting either visual-nonvisual or auditory-nonauditory experience during the first few months of life, the present study found that trisensory neurons in cat superior colliculus (as well as their bisensory counterparts) became adapted to the cross-modal stimulus combinations specific to each rearing environment. Thus, even at maturity, trisensory neurons did not integrate all cross-modal stimulus combinations to which they were capable of responding, but only those that had been linked via experience to constitute a coherent spatiotemporal event. This selective maturational process determines which environmental events will become the most effective targets for superior colliculus-mediated shifts of attention and orientation., (Copyright © 2015 the American Physiological Society.)

We made simultaneous bilateral recordings of unit activity in the nucleus ventroposterior lateralis (VPL) in intact rats and after acute and chronic left thoracic hemisection. We observed an immediate bilateral decline in multireceptive units, reflecting a loss of nociceptive input on the lesion side and a loss of low-threshold inputs contralaterally. Unit properties were restored to normal by 6 wk. Mean spontaneous discharge frequency remained unchanged in left VPL at all intervals. Right VPL displayed a substantial increase in spontaneous discharge frequency at 2 and 4 wk, returning to normal by 6 wk. Activity in left VPL driven by Pinch or Brush of the right limb was unchanged except for an immediate decrease in the response to Pinch, which was reversed by 2 wk despite persistent left hemisection. In right VPL, the response to Pinch or Brush of the left hindlimb was enhanced at 2 and 4 wk but returned to normal by 6 wk. Behaviorally, the same rats displayed increased sensitivity to mechanical stimulation of the left hindlimb, but, unlike VPL activity, there was no significant behavioral recovery. Bursting cells were also observed bilaterally in VPL, but this did not match the restriction of scratches to the hindlimb contralateral to the hemisection considered to be evidence for neuropathic pain. The novel findings include recovery of responsiveness to Pinch on the side ipsilateral to the hemisection despite the lack of spinothalamic input as well as failure for the thalamus contralateral to hemisection to maintain its elevated responsiveness.

In rats, navigating through an environment requires continuous information about objects near the head. Sensory information such as object location and surface texture are encoded by spike firing patterns of single neurons within rat barrel cortex. Although there are many studies using single-unit electrophysiology, much less is known regarding the spatiotemporal pattern of activity of populations of neurons in barrel cortex in response to whisker stimulation. To examine cortical response at the population level, we used voltage-sensitive dye (VSD) imaging to examine ensemble spatiotemporal dynamics of barrel cortex in response to stimulation of single or two adjacent whiskers in urethane-anesthetized rats. Single whisker stimulation produced a poststimulus fluorescence response peak within 12-16 ms in the barrel corresponding to the stimulated whisker (principal whisker). This fluorescence subsequently propagated throughout the barrel field, spreading anisotropically preferentially along a barrel row. After paired whisker stimulation, the VSD signal showed sublinear summation (less than the sum of 2 single whisker stimulations), consistent with previous electrophysiological and imaging studies. Surprisingly, we observed a spatial shift in the center of activation occurring over a 10- to 20-ms period with shift magnitudes of 1-2 barrels. This shift occurred predominantly in the posteromedial direction within the barrel field. Our data thus reveal previously unreported spatiotemporal patterns of barrel cortex activation. We suggest that this nontopographical shift is consistent with known functional and anatomic asymmetries in barrel cortex and that it may provide an important insight for understanding barrel field activation during whisking behavior.