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1  than to suppress responses to the principal whisker.
2 ralateral thalamus and cortex represent each whisker.
3 e responses to the deflection of surrounding whiskers.
4 uitable for visualization of one or more rat whiskers.
5 nd contralateral stimulation of the specific whiskers.
6 us, salt-and-pepper tuning to many different whiskers.
7  by a late overt cross-modal reactivation by whiskers.
8 rates and low sensitivity to the movement of whiskers.
9 continuous map of the space swept out by the whiskers.
10 the trigeminal nerve (PrV) correspond to the whiskers.
11    In the column corresponding to the spared whisker, 300 layer (L) 2/3 pyramidal neurons (17%) each
12 tryptamine (5-HT) receptors are expressed on whisker Abeta-afferent endings and that their activation
13 , primary among them the low contrast of the whisker against its background.
14 ploratory behavior, rats brush and tap their whiskers against objects, and the mechanical signals so
15 e of sensory input were driven by volitional whisker and body movements.
16 o bending moment (torque) at the base of the whisker and its rate of change and largely explained by
17 t and postnatal survival, but had defects in whisker and pelage hair formation.
18                           We showed that the whiskers and activity in the primary somatosensory corte
19 without whisking and that this involves both whiskers and barrel cortex activity.
20 formation available to rodents through their whiskers and how rodents direct their attention.
21 -response relationships involving individual whiskers and likely emerges within cortical circuits.
22 rafast "ping" (>50,000 degrees /s) to single whiskers and sampled primary afferent activity at 500 kH
23 tained a brief vibration stimulus applied to whiskers and the remaining 20% of trials contained a bri
24 ad to learn to localize an object with their whiskers, and based upon this location they were trained
25 ixed mice as they explored a pole with their whiskers, and simultaneously measured both whisker motio
26 n within a whisk cycle (phase), not absolute whisker angle, and arose from stresses reflecting whiske
27 ry neuron responses were poorly predicted by whisker angle, but well-predicted by rotational forces a
28 igh-frequency bursting activity at preferred whisker angles.
29      For example, titanium trisulfide (TiS3) whiskers are made out of weakly interacting TiS3 layers,
30       Movable tactile sensors in the form of whiskers are present in most mammals, but sensory coding
31 al microstimulation compared with the caudal whisker area (CWA) in animals under deep anesthesia.
32 he caudal forelimb area (CFA) and the caudal whisker area (CWA) of M1.
33 nd whisker area, which we termed the rostral whisker area (RWA), based on its differential response t
34  rostral forelimb area (RFA) and the rostral whisker area (RWA).
35 ve previously proposed existence of a second whisker area, which we termed the rostral whisker area (
36 urces of thalamic inputs to the two proposed whisker areas.
37     We used a three-dimensional model of the whisker array to construct mappings between the horizont
38                        Systems consisting of whisker arrays are fabricated, and as a proof of concept
39                                The shapes of whiskers, as well as their movements, are exquisitely ad
40 We also carried out stimulation of different whiskers at different times.
41  layers in regions surrounding the activated whisker barrel cortex.
42 al plasticity, we find reduced somatosensory whisker barrel plasticity.
43 c depolarization arose preferentially in the whisker barrel region of parietal sensory cortex.
44 chanical signals (forces and moments) at the whisker base while simultaneously monitoring whisker kin
45                   Here, we report electronic whiskers based on highly tunable composite films of carb
46                                              Whisker-based object localization requires activation an
47  in particular, can contribute to learning a whisker-based object localization task when timing is re
48 y across motor cortex while mice performed a whisker-based object localization task with a delayed, d
49 ved and studied in rodents in the context of whisker-based tactile sensation.
50                            Here we present a whisker-based, tactile virtual reality system for head-f
51 ariables, respectively) associated with this whiskers-based perceptual process.
52 daptations in the monocular cortex, in which whiskers become a dominant nonvisual input source to att
53                            We used models of whisker bending to quantify mechanical signals (forces a
54  discriminate different textures using their whiskers, both spike-rate and spike-timing information i
55 predicted by rotational forces acting on the whisker: both during touch and free-air whisker motion.
56 er receptive fields, including a single best whisker (BW) and lower magnitude responses to the deflec
57 ditional representations of the forelimb and whiskers, called the rostral forelimb area (RFA) and the
58            Compared to the long reaches that whiskers can make to the side and below the rat, the rea
59 ve (IoN), or selectively altered by repeated whisker clipping on the right side.
60 tivity in rat models of sensory deprivation (whisker clipping, tail suspension, casting).
61        During two of these phases, the rats' whiskers coded object position by first temporal and the
62       Sighted animals also showed changes in whisker control strategy with increased familiarity, but
63 uits for food, we tested the hypothesis that whisker control, as measured by high-speed videography,
64 rimary (S1) and secondary (S2) somatosensory whisker cortex during texture discrimination behavior, s
65 onization during evoked responses induced by whisker deflection did not differ between the two groups
66                                       Single whisker deflection elicited low-probability spikes in hi
67                              We found that a whisker deflection evoked abnormal sensory responses in
68 es, S1 neuronal responses to BW and surround whisker deflection showed comparable latencies in short-
69 ferent amounts of information about columnar whisker deflection.
70 isual stimuli, the magnitude of responses to whisker deflections is highest in the presence of optic
71  system begins as external forces that cause whisker deformations, which in turn excite mechanorecept
72                  In mice learning an active, whisker-dependent object localization task, layer 2/3 ne
73 hey showed noticeable deficits in all of the whisker-dependent or -related tests, including Y-maze ex
74 r performance in specific tests that require whisker-dependent tactile discrimination.
75 ssociates with deteriorated performance in a whisker-dependent texture discrimination task.
76                                   Unilateral whisker deprivation decreased the strength and spatial r
77  of rodent somatosensory cortex, where D-row whisker deprivation drives Hebbian weakening of whisker-
78 berrant structural plasticity in response to whisker deprivation, impaired texture novel object recog
79                                A majority of whisker discrimination tasks in rodents are performed on
80 tes and high-resolution videography of rats' whiskers during tactile exploration to study how texture
81   We show that in Mecp2-deficient male mice, whisker-evoked activity is roughly topographic but weak
82 etween wild-type and Fmr1 KO mice in overall whisker-evoked activity, though 45% fewer neurons in you
83 n contrast, chronic ACh deprivation hindered whisker-evoked CBF responses and the amplitude and power
84  enhanced ACh tone significantly potentiated whisker-evoked CBF responses through muscarinic ACh rece
85 neurons in vivo, brief deprivation decreased whisker-evoked inhibition more than excitation and incre
86                                 We show that whisker-evoked membrane depolarization in L2 PNs arises
87 al surface) allows for convenient imaging of whisker-evoked neural activity in vivo.
88               Cerebrovascular reactivity and whisker-evoked neurovascular coupling responses were mea
89  assessed the effects of varying ACh tone on whisker-evoked NVC responses in rat barrel cortex, measu
90  were compared across development and during whisker-evoked plasticity.
91                                              Whisker-evoked response probability correlated strongly
92  INs increases trial-to-trial variability of whisker-evoked responses in L2 PNs.
93 cal layer 1 (L1) of rat barrel cortex affect whisker-evoked responses of L2 PNs.
94 asing spontaneous firing and, in some cells, whisker-evoked responses.
95 we report new cortical regions downstream of whisker-evoked sensory processing during active explorat
96 atosensory cortex that transiently maintains whisker-evoked spiking in L2/3, despite the onset of Heb
97 sker deprivation drives Hebbian weakening of whisker-evoked spiking responses after an unexplained in
98 that deprivation (3 d) transiently increased whisker-evoked spiking responses in L2/3 single units be
99 lassical Hebbian weakening (>/=5 d), whereas whisker-evoked synaptic input was reduced during both pe
100                      Stimulation of a single whisker evokes neural activity sequentially in its corre
101  regions with high tactile acuity, including whisker follicles, fingertips and touch domes.
102           Rodents use their mechanosensitive whiskers for a diverse range of tactile behaviors such a
103 octurnal insects and mammals use antennae or whiskers for near-range orientation.
104 r, they moved more rapidly, protracted their whiskers further, and showed decreased whisking amplitud
105 hly abundant in fingertips, touch domes, and whisker hair follicles of mammals.
106                                  Using mouse whisker hair follicles, we show herein that tactile stim
107                                    Using rat whisker hair follicles, we show that Merkel cells rather
108              We trained rats to discriminate whisker impulse sequences that varied in single-impulse
109 r, these units did not effectively integrate whisker impulses, but instead combined weak impulse resp
110 hout substantial temporal integration across whisker impulses.
111       Discrete neural modules represent each whisker in the somatosensory cortex ("barrels"), thalamu
112 ker system of mice by deflection of a single whisker in vivo.
113 y of the tag to measure vibration in excised whiskers in a flume in response to laminar flow and dist
114                     These animals move their whiskers in a purposive manner to locations of interest.
115 ovel manual stimulation technique to deflect whiskers in a way that decouples kinematics from mechani
116  rats run at high speed, they protract their whiskers in front of the snout without large movements.
117 ead-fixed mice that tracked walls with their whiskers in tactile virtual reality.
118  in the thalamic barreloids by deflection of whiskers in vivo.
119 er angle, and arose from stresses reflecting whisker inertia and activity of specific muscles.
120                                       Rodent whisker input consists of dense microvibration sequences
121 it somewhat modest synaptic plasticity after whisker input deprivation.
122                                   Lesions to whisker input pathways had similar effects.
123             Thus, S1 encoded fast time scale whisker input without substantial temporal integration a
124 ary somatosensory cortex, generated by multi-whisker interactions during active touch.
125  directly encode mechanical signals when the whisker is deflected in this decoupled stimulus space.
126              A biomechanical modeling of the whisker is developed, which yields quantitative predicti
127                                  Each facial whisker is represented by discrete modules of neurons al
128 find that the sea lion's impressive array of whiskers is matched by a large trigeminal representation
129     Visualization and tracking of the facial whiskers is required in an increasing number of rodent s
130 x, activity related to movement of digits or whiskers is suppressed, which could facilitate detection
131 whisker base while simultaneously monitoring whisker kinematics and recording single Vg units in both
132 and temporal characteristics of the observed whisker kinetics, for any given topography.
133                    Many species that possess whiskers lack the modular "barrel" organization found in
134             Functional reorganization of the whisker map in rodent barrel cortex has long served as a
135 lly, the spatial organization of boutons and whisker map organization revealed the subdivision of the
136                                      The two whisker maps crowd in a space normally devoted to the co
137           This led to formation of bilateral whisker maps in both the thalamus and the cortex.
138 d uncrossed sensory inputs creates bilateral whisker maps in the thalamus and cortex.
139 se line, leads to the formation of bilateral whisker maps in the ventroposteromedial, as well as the
140                          Mice with bilateral whisker maps perform well in general sensorimotor tasks
141  interactions between the ipsi/contralateral whisker maps.
142  and ease of fabrication of the demonstrated whiskers may enable a wide range of applications in adva
143     Here we show that the vibrations of seal whiskers may provide information about hydrodynamic even
144  discrimination task, nor does it affect the whiskers' mechanical properties.
145 5 mice but is relaxed in adults to allow the whisker-mediated reactivation.
146 ed in two contexts in which either visual or whisker modality was more likely to occur.
147 ideography has proven adequate for measuring whisker motion and deformation during interaction with a
148 r whiskers, and simultaneously measured both whisker motion and forces with high-speed videography.
149  the whisker: both during touch and free-air whisker motion.
150 e information is mechanically encoded in the whisker motion.
151 This review focuses specifically on cortical whisker motor control.
152                                          The whisker motor neurons receive synaptic inputs from premo
153  a miniature accelerometer tag to study seal whisker movement in situ.
154 he middle portion of the rising phase of the whisker movement protraction elicited by artificial (fic
155 ion declined by half in surrounding columns; whisker movement representation was unchanged.
156     Purkinje cells (PCs) in Crus 1 represent whisker movement via linear changes in firing rate, but
157 and also show an increase in spike rate with whisker movement.
158 nucleus (VPM) fired in response to touch and whisker movement.
159  barrel cortex of neonatal rats, spontaneous whisker movements and passive stimulation by the litterm
160  barrel cortex of neonatal rats, spontaneous whisker movements and passive stimulation by the litterm
161 ar dependence on stimulus duration as evoked whisker movements and S1 activity.
162 eption, consistent with the coding of evoked whisker movements by reafferent sensory input.
163 ad in Emx1-Cre;Ai27D transgenic mice induces whisker movements due to activation of ChR2 expressed in
164                                  By studying whisker movements during tactile behaviors, we can learn
165 tes' position in the litter, and spontaneous whisker movements efficiently triggered bursts of activi
166  of a broader neural network that can decode whisker movements in air and on objects, which is a stra
167          In this primer, we focus on how the whisker movements of rats and mice are providing clues a
168 that the rats developed stereotypic head and whisker movements to solve this task, in a manner that c
169                       Under some conditions, whisker movements were phase-coupled to strides.
170 showed that tactile signals arising from the whisker movements with touch and stimulation by the litt
171                                         Yet, whisker movements with touch were more efficient than fr
172 e mice control tactile input through learned whisker movements.
173 ramidal neurons (17%) each encoded touch and whisker movements.
174          In mice locating objects with their whiskers, neurons in the ventral posteromedial nucleus (
175 s per animal, during performance of a single whisker object localization task.
176 on the transient response generated during a whisker-object collision.
177 ontrol of layer 4 neurons can substitute for whisker-object contact to guide behavior resembling wall
178                           Stimulation of the whiskers on either side activates the corresponding regi
179 inuous modulated noise sequence delivered to whiskers or fingertips, defined by its temporal patterni
180 single-impulse electrical stimulation of the whisker pad in the anesthetized rat to identify componen
181 ractions evoked by optogenetic activation of whisker pad muscles results in cortical activity and sen
182 anesthetized mice indicated that optogenetic whisker pad stimulation evokes robust yet longer latency
183 ead-fixed mice trained to report optogenetic whisker pad stimulation, psychometric curves showed simi
184 ion in glabrous skin of the paws, but in the whisker pads and body skin ectopic K8+ cells were confin
185 of the skin with additional abnormalities in whisker pads, footpads, and eyes.
186                                              Whisker plucking induces axonal growth and pruning of ho
187 atosensory cortex before and after selective whisker plucking.
188 d to the brain multiplexed information about whisker position and surface features, suggesting that p
189 ebellar Purkinje cells (PCs) linearly encode whisker position but the precise circuit mechanisms that
190                Self-motion responses encoded whisker position within a whisk cycle (phase), not absol
191                       We found that surround whiskers powerfully transform the cortical representatio
192 vates brain stem reticular nuclei containing whisker premotor neurons, which might form a central pat
193                                    Mammalian whiskers present an important class of tactile sensors t
194                                          The whisker primary motor cortex (M1) strongly innervates br
195   Here, we investigate the function of mouse whisker primary motor cortex (wM1), a frontal region def
196 tal region defined by dense innervation from whisker primary somatosensory cortex (wS1).
197 togenetic stimulation of wM1 evokes rhythmic whisker protraction (whisking), whereas optogenetic inac
198 orm a central pattern generator for rhythmic whisker protraction.
199                             We conclude that whisker protractions evoked by optogenetic activation of
200 tized mice, we characterize the amplitude of whisker protractions evoked by varying the intensity, du
201 y be 'the window to the soul' in humans, but whiskers provide a better path to the inner lives of rod
202 sion discriminates between single- and multi-whisker receptive field layer 2 pyramidal neurons.
203 ging with deflection of many whiskers to map whisker receptive fields, characterize sparse coding, an
204                    Here, we show that in the whisker-related barrel cortex of neonatal rats, spontane
205                   Here, we show that, in the whisker-related barrel cortex of neonatal rats, spontane
206 te layers of the superior colliculus receive whisker-related excitatory afferents from the trigeminal
207 (S1) of mice and rats, but it is unclear how whisker-related input is represented in these species.
208 nts, possibly due to the smaller size of the whisker-related modules and interference between the ips
209            In addition, both spontaneous and whisker-related neural activities in the superior collic
210 tte neurons cross the midline and confer the whisker-related patterning to the contralateral ventropo
211 cyte differentiation from divided cells, and whisker removal decreased the survival of divided cells
212  mammals, but sensory coding in the cortical whisker representation has been studied almost exclusive
213 e report ipsilateral cortical connections of whisker representation in RMA, and compare them with con
214 tical imaging verified functional, bilateral whisker representation in the barrel cortex and activati
215                           Barrel cortex, the whisker representation of primary somatosensory cortex,
216 s are segregated resulting in duplication of whisker representations and doubling of the number of ba
217 ce between the ipsilateral and contralateral whisker representations in the same thalamus and cortex.
218 om neurons located in the surrounding intact whisker representations.
219                                 Importantly, whisker response amplitude is also modulated by presenta
220 d provide evidence for dynamic regulation of whisker responses according to visual experience.
221 nitial delay, but no homeostasis of deprived whisker responses is known.
222 timuli modulates the amplitude of concurrent whisker responses.
223  electrophysiology, we find that a subset of whisker-responsive neurons in the ventral posterior medi
224  electrophysiology, we find that a subset of whisker-responsive neurons in the ventral posterior medi
225 c whisking, whereas stimulation of S1 drives whisker retraction.
226                  Removal of all but a single whisker row for 24 h led to an apparent increase in syna
227                   The process makes the dyed whisker(s) easily visible against a dark background.
228 ponents, thereby allowing for exploration of whisker sensors with excellent performance.
229 ed voltage-sensitive dye imaging to evaluate whisker sensory evoked activity in the barrel cortex of
230                                          The whisker sensory system of rodents is an excellent model
231 alamus and cortex.SIGNIFICANCE STATEMENT The whisker sensory system plays a quintessentially importan
232                                         In a whisker session, 80% of trials contained a brief vibrati
233 ced response to vibration stimuli during the whisker session.
234  rate provide an accurate linear read-out of whisker set point.
235 ing and tests of remote memory retention for whisker-signaled trace eyeblink conditioning.
236 underwater hydrodynamic trail to measure the whisker signals available to the seal.
237 ry neurons in the juvenile (P18 to 27) mouse whisker somatosensory cortex, distinguished by expressio
238 d that mEPSC frequency nearly doubled in the whisker-spared column, a difference that was highly sign
239                       In vivo recordings and whisker-specific behavioral tests demonstrated sensory d
240 TEMENT We use a novel paradigm of repetitive whisker stimulation and in vivo calcium imaging to asses
241  mice during periods of true rest and during whisker stimulation and volitional whisking.
242  whole-cell recordings showed that principal whisker stimulation elicits similar amplitude synaptic r
243 deficit in neuronal adaptation to repetitive whisker stimulation in both young and adult Fmr1 KO mice
244 we discovered exaggerated motor responses to whisker stimulation in young Fmr1 knock-out (KO) mice (p
245 dilation and hemodynamic responses evoked by whisker stimulation involve cyclooxygenase-2 (COX-2) act
246 tosensory barrel cortex, we found that acute whisker stimulation led to a significant increase in the
247  contrast, enhancement of neural activity by whisker stimulation led to an increase in vascular densi
248 increase in spine sGluA1 intensity evoked by whisker stimulation was NMDA receptor dependent and long
249                                 Optogenetic, whisker stimulation, or cortical spreading depolarizatio
250 aptation of thalamic responses to repetitive whisker stimulation, thereby allowing thalamic neurons t
251 eptor-mediated Ca(2+) signaling, we identify whisker stimulation-evoked large responses in a subset o
252 yet longer latency responses than mechanical whisker stimulation.
253 d dendritic shaft sGluA1 intensity following whisker stimulation.
254 n and blood volume in response to mechanical whisker stimulation.
255 ne properties biased spine changes following whisker stimulation.
256 r gene, would be preferentially activated by whisker stimulation.
257 neuronal adaptation in barrel cortex, during whisker stimulation.
258 ding in rats with a multi-directional, multi-whisker stimulator system to estimate receptive fields b
259 on conveyed by thalamic neurons about paired whisker stimuli in male rat.
260         In contrast with conventional single-whisker stimuli, complex stimuli revealed markedly sharp
261 on; VPM) can be excited by visual as well as whisker stimuli.
262  microelectrode arrays to record ongoing and whisker stimulus-evoked population spiking activity in s
263 ats moved at slow speeds and performed broad whisker sweeps.
264                                          The whisker system is an important sensory organ with extens
265 the function of the cerebellum in the rodent whisker system is unknown.
266 nidentified slowly adapting afferents in the whisker system of behaving mice respond to both self-mot
267 to record 3D neural activities evoked in the whisker system of mice by deflection of a single whisker
268  without whisking.SIGNIFICANCE STATEMENT The whisker system of rodents is a widely used model to stud
269                                          The whisker system of rodents is an excellent model to study
270 tion, we probed spatial coding in the rodent whisker system using a combination of two-photon imaging
271 the remarkable abilities of their trigeminal whisker system.
272 hey do so by using two functionally distinct whisker systems: the macrovibrissae and microvibrissae.
273                     Neurons tuned to ventral whiskers tended to show broad tuning along the rostrocau
274 dination when motor behavior was paired with whisker-texture touches, suggesting that direct S1-S2 in
275 revealed an asymmetry in the position of the whiskers: they oriented toward the rewarded stimulus dur
276  to the network architecture made by the TiB whiskers (TiBw), and a decrease of the steady-state cree
277                            Rodents use their whiskers to detect a variety of tactile features of thei
278 oton calcium imaging with deflection of many whiskers to map whisker receptive fields, characterize s
279   To perform the task, the rat positions its whiskers to receive two such stimuli, "base" and "compar
280                              Mice used their whiskers to track the walls of the winding corridor with
281 ed to differences in the architecture of the whisker-to-cortex pathway.
282                        We stimulated several whiskers together to determine the sensitivity of our ap
283 ation was delivered in spatial register with whisker topography learned the task more quickly.
284 ference between the number of right and left whiskers touching the surface.
285                                              Whisker-tracking analysis revealed an asymmetry in the p
286 king sensory responsiveness before and after whisker trimming has uncovered diverse effects in indivi
287 t neither reduction of sensory input through whisker trimming nor moderately increased activity by en
288 f the vibrissal (barrel) S1 after unilateral whisker trimming.
289  dendritic spine loss, acutely (4-8 d) after whisker trimming.
290 d secondary somatosensory cortex differed in whisker tuning and responsiveness, and carried different
291   High-speed video of behaving mice revealed whisker velocities of at least 17,000 degrees /s, so we
292 relevant variables, such as head azimuth and whisker velocity.
293                      The results showed that whiskers vibrated at frequencies of 100-300 Hz, with a d
294 single units responded differentially to the whisker vibration stimulus when presented with higher pr
295                  Rats were trained to detect whisker vibrations or visual flickers.
296 ent surface coarseness or controlled passive whisker vibrations simulating different coarseness, we s
297 lthough it is known that seals can use their whiskers (vibrissae) to extract relevant information fro
298                                     However, whisker visualization and tracking is challenging for mu
299 itivity of up to approximately 8%/Pa for the whiskers, which is >10x higher than all previously repor
300 riminating surfaces by actively moving their whiskers (whisking) against stimuli, typically sampling

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