Passive Stimulation Research Articles

Predicting successful tactile mapping of virtual objects.

Improving spatial ability of blind and visually impaired people is the main target of orientation and mobility (O&M) programs. In this study, we use a minimalistic mouse-shaped haptic device to show a new approach aimed at evaluating devices providing tactile representations of virtual objects. We consider psychophysical, behavioral, and subjective parameters to clarify under which circumstances mental representations of spaces (cognitive maps) can be efficiently constructed with touch by blindfolded sighted subjects. We study two complementary processes that determine map construction: low-level perception (in a passive stimulation task) and high-level information integration (in an active exploration task). We show that jointly considering a behavioral measure of information acquisition and a subjective measure of cognitive load can give an accurate prediction and a practical interpretation of mapping performance. Our simple TActile MOuse (TAMO) uses haptics to assess spatial ability: this may help individuals who are blind or visually impaired to be better evaluated by O&M practitioners or to evaluate their own performance.

Behavioural and neurophysiological markers reveal differential sensitivity to homeostatic interactions between centrally and peripherally applied passive stimulation

Repetitive transcranial magnetic stimulation (rTMS) is an effective tool for inducing functional plastic changes in the brain. rTMS can also potentiate the effects of other interventions such as tactile coactivation, a form of repetitive stimulation, when both are applied simultaneously. In this study, we investigated the interaction of these techniques in affecting tactile acuity and cortical excitability, measured with somatosensory evoked potentials after paired median nerve stimulation. We first applied a session of 5-Hz rTMS, followed by a session of tactile repetitive stimulation, consisting of intermittent high-frequency tactile stimulation (iHFS) to a group of 15 healthy volunteers ("rTMS+iHFS" group). In a second group ("rTMS w/o iHFS"), rTMS was applied without iHFS, with a third assessment performed after a similar wait period. In the rTMS w/o iHFS group, the 5-Hz rTMS induced an increase in cortical excitability that continued to build for at least 25min after stimulation, with the effect on excitability after the wait period being inversely correlated to the baseline state. In the rTMS+iHFS group, the second intervention prevented the continued increase in excitability after rTMS. In contrast to the effect on cortical excitability, rTMS produced an improvement in tactile acuity that remained stable until the last assessment, independent of the presence or absence of iHFS. Our results show that these methods can interact homeostatically when used consecutively, and suggest that different measures of cortical plasticity are differentially susceptible to homeostatic interactions.

Occipital cortex activation by long-term repetitive tactile stimulation is necessary for object recognition in blinds: A case report

Tactile vision has been approached from a variety of angles using different techniques. So far, a certain kind of object (and text) recognition has been shown, though seeing as such has not been achieved yet, and it remains unclear. Trough repetitive passive tactile stimulation perceptual processing is transferred from temporo-parietal to occipital areas, which affects object recognition. We report the results of passive tactile stimulation, as well as rTMS, applied to a 50 year old left handed blind male with over 97% loss of vision, who suffers from Peter’s anomaly and microphthalmia. After 15 weeks of passive tactile stimulation, the subject showed increased activity in occipital areas associated with the development of visual-like perception which remained unchanged after three months without passive tactile stimulation. Inhibitory rTMS over the visual cortex led to noticeable reduction of spatial recognition performance and visual sensations in this subject. Stable changes in occipital cortical activity can be associated with subjective sensations of seeing. Once occipital activation has been achieved, it is necessary for spatial object recognition. Both facts highlight the implication of occipital areas in tactile vision and the cortical plasticity of passive tactile long-term stimulation in people with blindness.

Spatiotemporal Segregation of Neural Response to Auditory Stimulation: An fMRI Study Using Independent Component Analysis and Frequency-Domain Analysis

Although auditory processing has been widely studied with conventional parametric methods, there have been a limited number of independent component analysis (ICA) applications in this area. The purpose of this study was to examine spatiotemporal behavior of brain networks in response to passive auditory stimulation using ICA. Continuous broadband noise was presented binaurally to 19 subjects with normal hearing. ICA was performed to segregate spatial networks, which were subsequently classified according to their temporal relation to the stimulus using power spectrum analysis. Classification of separated networks resulted in 3 stimulus-activated, 9 stimulus-deactivated, 2 stimulus-neutral (stimulus-dependent but not correlated with the stimulation timing), and 2 stimulus-unrelated (fluctuations that did not follow the stimulus cycles) components. As a result of such classification, spatiotemporal subdivisions were observed in a number of cortical structures, namely auditory, cingulate, and sensorimotor cortices, where parts of the same cortical network responded to the stimulus with different temporal patterns. The majority of the classified networks seemed to comprise subparts of the known resting-state networks (RSNs); however, they displayed different temporal behavior in response to the auditory stimulus, indicating stimulus-dependent temporal segregation of RSNs. Only one of nine deactivated networks coincided with the “classic” default-mode network, suggesting the existence of a stimulus-dependent default-mode network, different from that commonly accepted.

Le toucher chez les enfants prématurés

Touch being the first sense to develop in utero, it plays an essential role in the perceptual-cognitive development in infancy. The study of tactile skills in premature infants is a relevant approach to better understand the early development of touch. After briefly summarizing some elements of neurophysiological substrates of passive and active touch, namely the somatosensory system and its early development, this article describes, first, a review of experimental work on early skills of premature infants in both passive touch and active touch, and secondly, a summary of studies about two programs of developmental care in which touch is an essential component: the massage therapy and the kangaroo mother care. Overall, studies show in premature infants the occurrence of 1. discrimination subsequent to passive tactile stimulation from 30 weeks of gestation; 2. active manual discrimination between two different object shapes from 28 weeks and finally 3. beneficial effects on their neurodevelopmental future after receiving a massage therapy or the kangaroo mother care.

Head Stabilization Shows Visual and Inertial Dependence During Passive Stimulation: Implications for Virtual Rehabilitation

Sensorimotor coordination relies on the fine calibration and integration of visual, vestibular, and somatosensory input. Using virtual environments (VE) allows for the dissociation of visual and inertial inputs to manipulate human behavioral outputs. Our goal was to employ VE technology in a novel manner to investigate how head stabilization is affected by spatiotemporal properties of dynamic visual input when combined with passive motion on a linear sled. Healthy adults (n = 12) wore a head-mounted display during naso-occipital sinusoidal horizontal whole body translations while seated. Subjects were secured in a seat with a five-point harness, with the head free to move. Frequency and amplitude of sinusoidal input (i.e., inertial conditions) were set to create overlapping conditions of maximum acceleration (amax) or velocity (vmax). Four inertial conditions were combined with four visual conditions (VIS). VIS were created so that direction of optic flow either matched direction of passive motion or did not. The effect of near and far fixation distance within the VE was also tested. Head kinematics were collected with a three-axis gyro. Head stability showed a complex interaction dependent on changes in weighting of visual and inertial inputs that changed with the sled driving frequency. Inertial condition affected amplitude (p < 0.0000) and phase (p < 0.0000) of head pitch angular velocity. In the absence of visual input, head pitch velocity amplitude increased (p < 0.01). An interaction effect between inertial and VIS conditions on head yaw occurred in SW (p < 0.05). There was also a significant interaction of depth of field and inertial condition on amplitude (p < 0.001) and phase (p < 0.05) of head yaw velocity in SW, especially during high vmax conditions. We conclude visual flow can organize lateral cervical responses despite being discordant with inertial input. When using VE for rehabilitation, possible unintended, involuntary or reflexive motor responses that may not be present in traditional training environments should be taken into consideration.

Self-regulated music stimulation for persons with Alzheimer's disease: Impact assessment and social validation

Objective: To assess the impact and the social rating of an active music condition (in which 10 patients with Alzheimer's disease regulated their music input) vs. a passive music condition.Method: In the active condition, the patients used a simple hand response and a microswitch to activate music stimulation periods. In the passive condition, music stimulation was prearranged and continued through the sessions. The active and passive stimulation sessions were preceded and followed by control (non-stimulation) sessions.Results: The active condition sessions showed an increase in the patients’ indices of positive participation (e.g., singing or music-related movements, and smiles) similar to that observed in the passive condition sessions. Social raters (140 psychology students) favored the active condition on a six-item questionnaire dealing, among others, with conditions’ suitability, respect of patients’ dignity and independence, and practicality.Conclusion: An active music stimulation condition can be viable, effective, and socially preferable.

Functional repetitive transcranial magnetic stimulation increases motor cortex excitability in survivors of stroke

ObjectiveTo determine if repetitive transcranial magnetic stimulation (rTMS) applied to the motor cortex with simultaneous voluntary muscle activation, termed functional-rTMS, will promote greater neuronal excitability changes and neural plasticity than passive-rTMS in survivors of stroke. MethodsEighteen stroke survivors were randomized into functional-rTMS (EMG-triggered rTMS) or passive-rTMS (rTMS only; control) conditions. Measures of short-interval intracortical inhibition (SICI) and intracortical facilitation (ICF), force steadiness (coefficient of variation, CV) at 10% of maximum voluntary contraction, and pinch task muscle activity were assessed before and after rTMS. Functional-rTMS required subjects to exceed a muscle activation threshold to trigger each rTMS train; the passive-rTMS group received rTMS while relaxed. ResultsSignificant interactions (time×condition) were observed in abductor pollicis brevis (APB) SICI, APB ICF, CV of force, and APB muscle activity. Functional-rTMS decreased APB SICI (p<0.05) and increased ICF (p<0.05) after stimulation, whereas passive-rTMS decreased APB muscle activity (p<0.01) and decreased CV of force (p<0.05). No changes were observed in FDI measures (EMG, ICF, SICI). Conclusion(s)Functional-rTMS increased motor cortex excitability, i.e., less SICI and more ICF for the APB muscle. Passive stimulation significantly reduced APB muscle activity and improved steadiness. SignificanceFunctional-rTMS promoted greater excitability changes and selectively modulated agonist muscle activity.

Assessing the impact and social perception of self-regulated music stimulation with patients with Alzheimer's disease

We assessed the impact and social rating of an active and a passive music condition implemented with six patients with Alzheimer's disease. In the active condition, the patients used a simple hand response and a microswitch to self-regulate music stimulation inputs. In the passive condition, music stimulation was automatically presented throughout the sessions. Active and passive stimulation sessions were preceded and followed by control (non-stimulation) sessions. The active condition sessions showed an increase in the patients' indices of positive participation (e.g., singing or music-related movements, and smiles) greater than that observed in the passive condition sessions for five of the six patients. Positive intervention effects could also spread to the post-intervention sessions. Social raters (42 care and rehabilitation staff members working with persons with multiple disabilities) favored the active condition on a six-item questionnaire dealing with, among others, conditions' suitability, respect of patients' dignity and independence, and practicality. The implications of the findings as to the plausibility/desirability of an active stimulation condition were discussed.

Tactile exploration of virtual objects for blind and sighted people: the role of beta 1 EEG band in sensory substitution and supramodal mental mapping

The neural correlates of exploration and cognitive mapping in blindness remain elusive. The role of visuo-spatial pathways in blind vs. sighted subjects is still under debate. In this preliminary study, we investigate, as a possible estimation of the activity in the visuo-spatial pathways, the EEG patterns of blind and blindfolded-sighted subjects during the active tactile construction of cognitive maps from virtual objects compared with rest and passive tactile stimulation. Ten blind and ten matched, blindfolded-sighted subjects participated in the study. Events were defined as moments when the finger was only stimulated (passive stimulation) or the contour of a virtual object was touched (during active exploration). Event-related spectral power and coherence perturbations were evaluated within the beta 1 band (14-18 Hz). They were then related to a subjective cognitive-load estimation required by the explorations [namely, perceived levels of difficulty (PLD)]. We found complementary cues for sensory substitution and spatial processing in both groups: both blind and sighted subjects showed, while exploring, late power decreases and early power increases, potentially associated with motor programming and touch, respectively. The latter involved occipital areas only for blind subjects (long-term plasticity) and only during active exploration, thus supporting tactile-to-visual sensory substitution. In both groups, coherences emerged among the fronto-central, centro-parietal, and occipito-temporal derivations associated with visuo-spatial processing. This seems in accordance with mental map construction involving spatial processing, sensory-motor processing, and working memory. The observed involvement of the occipital regions suggests that a substitution process also occurs in sighted subjects. Only during explorations did coherence correlate positively with PLD for both groups and in derivations, which can be related to visuo-spatial processing, supporting the existence of supramodal spatial processing independently of vision capabilities.

Olfactory Processing in Children and Young Adults

Because central nervous processing of odorous stimuli in different stages of adolescence has rarely been studied, in this preliminary fMRI investigation, 20 subjects were grouped according to age (children 9–12 years old and young adults 17–20 years old) to build two equally sized samples. Patterns of cerebral activation were compared between both groups following passive nasal stimulation with three odorants. Children mainly showed activation of olfactory cortex, namely, piriform cortex and amygdala. In young adults, enhanced activation was revealed in neocortical areas, suggesting a greater involvement of areas engaged in cognitive integration of olfactory stimuli. However, it is emphasized that additional psychological and developmental confounds should be taken into account in future studies to establish more profound results.

Viability, Morphology and Function of Primary Endothelial Cells on Poly(n-Butyl Acrylate) Networks Having Elastic Moduli Comparable to Arteries

Soft hydrophobic poly(n-butyl acrylate) networks (cPnBA) were developed as entropy elastic substrates for passive mechanical stimulation of cells, where the elastic modulus of the cPnBAs could be systematically adjusted by variation of the cross-link density. The networks were synthesized by thermally-induced radical polymerization from n-butyl acrylate, with poly(propylene glycol) dimethacrylate (PPGDMA) acting as cross-linker, whereby the purity of the cPnBAs was confirmed by1 H-NMR spectroscopy and gas chromatography. In this work two cPnBA polymer networks with an elastic modulus around 200 kPa and 1 MPa were investigated having an elastic modulus similar to that of arteries. Both cPnBAs exhibited an almost smooth surface with a surface roughness (R q) in the wet state ranging from 17 to 37 nm and a similar zetapotential, indicating an almost identical chemical composition within the topmost surface layer in terms of functional groups. In contrast, wettability of the samples was found to be different with an advancing angle ( advancing) of 123 ± 3.8° for cPnBA0250, while for cPnBA1100 significantly lower values for advancing (111 ± 3.8°) were obtained. First in vitro tests were performed with primary endothelial cells (HUVEC) to study its effects on vascular cell functions. Within the time period of cultivation (72 h), the cells on the cPnBA samples reached subconfluence and showed a viability rate of almost 100%. Although cell density differed after 72 h with more cells on cPnBA0250 than on cPnBA1100, both materials showed no significant effect on the cell morphology, the cellular LDH-release, which was used as marker for the integrity of the cell membrane, and the organisation of the VE-cadherin. However, lower cell density and less actin stress fibre formation on cPnBA1100 might indicate that cell–material interaction was weaker on cPnBA1100 than on cPnBA0250. The secretion of the vasoactive cytokines prostacyclin (PGI2) and thromboxane A2 (TXA2) was low compared to previously reported values. However, the anti-thrombogenic ratio of PGI2/TXA2 — which is balanced under physiological conditions — with much higher PGI2 compared to TXA2 (up to 17.6-fold after 72 h for cPnBA1100) suggests that this material might be effective to preventing thrombosis.

Neuronal modulation in appetitive and aversive contexts in the primate lateral hypothalamus

Sensory experience has a profound effect on neocortical neurons. Passive stimulation of whiskers or sensory deprivation from whiskers can induce long-lasting changes in neuronal responses or modify the receptive field in adult animals. We recorded barrel cortical neurons in urethane-anesthetized rats in layers 2/3 or 5/6 to determine if repetitive stimulation would induce long-lasting response facilitation. Air-puff stimulation (20-ms duration, 40 pulses at 0.5–8 Hz) was applied to a single whisker. This repetitive stimulation increased tactile responses in layers 2/3 and 5/6 for 60 min. Moreover, the functional coupling (coherence) between the sensory stimulus and the neural response also increased after the repetitive stimulation in neurons showing response facilitation. The long-lasting response facilitation was due to activation of N-methyl-d-aspartate (NMDA) receptors because it was reduced by APV ((2R)-amino-5-phosphonovaleric acid, (2R)-amino-5-phosphonopentanoate) and MK801 application. Inactivation of layer 2/3 also blocked response facilitation in layer 5/6, suggesting that layer 2/3 may be fundamental in this synaptic plasticity processes. Moreover, i.p. injection of eserine augmented the number of layer 2/3 neurons expressing long-lasting response facilitation; this effect was blocked by atropine, suggesting that muscarinic receptor activation favors the induction of the response facilitation. Our data indicate that physiologically repetitive stimulation of a single whisker at the frequency at which rats move their whiskers during exploration of the environment induces long-lasting response facilitation improving sensory processing.

A computational method for developing hierarchical large deformation viscoelastic models of the contracting heart

In this study, a new computational method for modelling the contracting heart is described. Using this method, the cardiac wall is constructed from basic, repeating contractile units that represent individual myocardial units and collagen, each with its own set of parameters, including orientation, passive and active behaviour and stimulation propagation. The method allows individual control of each structural unit (e.g. at the level of a single myocardial unit). Feasibility of modelling dynamic heart contraction using this method is demonstrated using 2D cross-sections and simplified 3D geometries. Effects of non-contractile scar and myopathic tissue were also tested in these geometrical configurations. Results from the 2D and 3D simulations were, overall, in agreement with well-established physiological data. The present method holds promise for modelling complex heart pathologies, abnormal mechanical properties (e.g. myocardial infarcts) and electrical conduction properties (branch blocks), and their spatial distributions across the myocardial tissues.

Behavioral training enhances cortical temporal processing in neonatally deafened juvenile cats

Deaf humans implanted with a cochlear prosthesis depend largely on temporal cues for speech recognition because spectral information processing is severely impaired. Training with a cochlear prosthesis is typically required before speech perception shows improvement, suggesting that relevant experience modifies temporal processing in the central auditory system. We tested this hypothesis in neonatally deafened cats by comparing temporal processing in the primary auditory cortex (AI) of cats that received only chronic passive intracochlear electric stimulation (ICES) with cats that were also trained with ICES to detect temporally challenging trains of electric pulses. After months of chronic passive stimulation and several weeks of detection training in behaviorally trained cats, multineuronal AI responses evoked by temporally modulated ICES were recorded in anesthetized animals. The stimulus repetition rates that produced the maximum number of phase-locked spikes (best repetition rate) and 50% cutoff rate were significantly higher in behaviorally trained cats than the corresponding rates in cats that received only chronic passive ICES. Behavioral training restored neuronal temporal following ability to levels comparable with those recorded in naïve prior normal-hearing adult deafened animals. Importantly, best repetitition rates and cutoff rates were highest for neuronal clusters activated by the electrode configuration used in behavioral training. These results suggest that neuroplasticity in the AI is induced by behavioral training and perceptual learning in animals deprived of ordinary auditory experience during development and indicate that behavioral training can ameliorate or restore temporal processing in the AI of profoundly deaf animals.

Effect of Three Different Between-Inning Recovery Methods on Baseball Pitching Performance

A decrease in blood hydrogen ions (H) may allow for the recovery of a muscle, which should allow for greater performance in subsequent activity. The purpose of this study was to determine which of 3 forms of recovery were the most effective after an inning of pitching in baseball. Three different measurements were used to determine which recovery method was most effective; the difference in blood lactate (BLa) levels was used as a biological measurement, average pitching speed was the physiological measurement, and the psychological measurement was done on how the pitchers perceived their pitching and recovery. The recovery methods that were used were passive recovery (PR), active recovery (AR), and electromuscular stimulation (EMS). Seven college men aged 21 (±2 years) who were National Collegiate Athletic Association Division II college baseball pitchers were assessed during game play simulations. Blood lactate levels decreased significantly from the premeasurement to the postmeasurement with the EMS recovery method (p < 0.0005); however, BLa did not change for PR (p = 0.017) or AR (p = 0.134). Perceived recovery was also found to be best in the EMS and PR conditions. These findings suggest that EMS is an effective recovery method between innings of pitching.

Modulation of the response to a somatosensory stimulation of the hand during the observation of manual actions

Observation of hand movements has been repeatedly demonstrated to increase the excitability of the motor cortical representation of the hand. Little attention, however, has been devoted to its effect on somatosensory processing. Movement execution is well known to decrease somatosensory cortical excitability, a phenomenon termed 'gating'. As executed and observed actions share common cortical representations, we hypothesized that action observation (hand movements) should also modulate the cortical response to sensory stimulation of the hand. Seventeen healthy subjects participated in these experiments in which electroencephalographic (EEG) recordings of the somatosensory steady-state response (SSSR) were obtained. The SSSR provides a continuous measure of somatosensory processing. Recordings were made during a baseline condition and five observation conditions in which videos showed either a: (1) hand action; (2) passive stimulation of a hand; (3) static hand; (4) foot action; or (5) static object. The method employed consisted of applying a continuous 25 Hz vibratory stimulation to the index finger during the six conditions and measuring potential gating effects in the SSSR within the 25 Hz band (corresponding to the stimulation frequency). A significant effect of condition was found over the contralateral parietal cortex. Observation of hand actions resulted in a significant gating effect when compared to baseline (average gating of 22%). Observation of passive touch of the hand also gated the response (17% decrease). In conclusion, the results show that viewing a hand performing an action or being touched interferes with the processing of somatosensory information arising from the hand.

Inferior Olive Response to Passive Tactile and Visual Stimulation with Variable Interstimulus Intervals

The unique anatomical and electrophysiological features of the inferior olive and its importance to cerebellar function have been recognized for decades. However, understanding the exact function of the inferior olive has been limited by the general lack of correlation between its neural activity and specific behavioral states. Electrophysiological studies in animals showed that the inferior olive response to sensory stimuli is generally invariant to stimulus properties but is enhanced by unexpected stimuli. Using functional magnetic resonance imaging in humans, we have shown that the inferior olive is activated when subjects performed a task requiring perception of visual stimuli with unpredictable timing (Xu et al. J Neurosci 26(22):5990-5995, 2006, Liu et al. J Neurophysiol 100(3):1557-1561, 2008). In the current study, subjects were scanned while passively perceiving visual and tactile stimuli that were rendered unpredictable by continuously varying interstimulus intervals (ISIs). Sequences of visual stimuli and tactile stimuli to the right hand were presented separately within the same scanning session. In addition to the activation of multiple areas in the cerebellar cortex consistent with previous imaging studies, the results show that both tactile and visual stimulation with variable ISIs were effective in activating the inferior olive. Together with our previous findings, the current results are consistent with the electrophysiological studies in animals and further support the view that the inferior olive and the climbing fiber system primarily convey the temporal information of sensory input regardless of the modality.

Transmodal comparison of auditory, motor, and visual post-processing with and without intentional short-term memory maintenance

Objective To elucidate the contributions of modality-dependent post-processing in auditory, motor and visual cortical areas to short-term memory. Methods We compared late negative waves (N700) during the post-processing of single lateralized stimuli which were separated by long intertrial intervals across the auditory, motor and visual modalities. Tasks either required or competed with attention to post-processing of preceding events, i.e. active short-term memory maintenance. Results N700 indicated that cortical post-processing exceeded short movements as well as short auditory or visual stimuli for over half a second without intentional short-term memory maintenance. Modality-specific topographies pointed towards sensory (respectively motor) generators with comparable time-courses across the different modalities. Lateralization and amplitude of auditory/motor/visual N700 were enhanced by active short-term memory maintenance compared to attention to current perceptions or passive stimulation. The memory-related N700 increase followed the characteristic time-course and modality-specific topography of the N700 without intentional memory-maintenance. Conclusions Memory-maintenance-related lateralized negative potentials may be related to a less lateralised modality-dependent post-processing N700 component which occurs also without intentional memory maintenance (automatic memory trace or effortless attraction of attention). Encoding to short-term memory may involve controlled attention to modality-dependent post-processing. Significance Similar short-term memory processes may exist in the auditory, motor and visual systems.

Multiple Timescale Encoding of Slowly Varying Whisker Stimulus Envelope in Cortical and Thalamic NeuronsIn Vivo

Adaptive processes over many timescales endow neurons with sensitivity to stimulus changes over a similarly wide range of scales. Although spike timing of single neurons can precisely signal rapid fluctuations in their inputs, the mean firing rate can convey information about slower-varying properties of the stimulus. Here, we investigate the firing rate response to a slowly varying envelope of whisker motion in two processing stages of the rat vibrissa pathway. The whiskers of anesthetized rats were moved through a noise trajectory with an amplitude that was sinusoidally modulated at one of several frequencies. In thalamic neurons, we found that the rate response to the stimulus envelope was also sinusoidal, with an approximately frequency-independent phase advance with respect to the input. Responses in cortex were similar but with a phase shift that was about three times larger, consistent with a larger amount of rate adaptation. These response properties can be described as a linear transformation of the input for which a single parameter quantifies the phase shift as well as the degree of adaptation. These results are reproduced by a model of adapting neurons connected by synapses with short-term plasticity, showing that the observed linear response and phase lead can be built up from a network that includes a sequence of nonlinear adapting elements. Our study elucidates how slowly varying envelope information under passive stimulation is preserved and transformed through the vibrissa processing pathway.