Biomedical subjects

Ford F Ebner

Publications and source records attributed to Ford F Ebner.

18 recordsLinked to original sources

PubMed2007

Cortical modulation of spatial and angular tuning maps in the rat thalamus.

The massive feedback projections from cortex to the thalamus modulate sensory information transmission in many ways. We investigated the role of corticothalamic feedback projections on the directional selectivity (angular tuning) of neurons in the rat ventral posterior medial (VPM) nucleus to stimulation of their principal whisker. The angular tuning properties of single VPM neurons were compared before and after epochs of electrical stimulation of layer VI feedback neurons in the ipsilateral cortex under urethane anesthesia. Microstimulation of layer VI in "matched" (homologous) barrel columns sharpens the angular tuning curves of single VPM neurons that are tuned to the same direction as the stimulation site in the cortex. Further, microstimulation rotates the angular preference of VPM neurons initially tuned to a different direction toward the direction that cortical neurons prefer. Stimulation in "mismatched" (nonhomologous) barrel columns suppresses responses without consistent effects on angular tuning. We conclude that the primary sensory cortex exerts a significant influence on both spatial and angular tuning maps in the relay nuclei that project to it. The results suggest that the tuning properties of VPM cells in the behaving animal are continually modified to optimize perception of the most salient incoming messages.

Adaptation, Physiological↗

PubMed2006

Stimulus frequency processing in awake rat barrel cortex.

In awake rats, we examined the relationship between neural spiking activity in primary somatic sensory cortex and the frequency of whisker stimulation. Neural responses were recorded extracellularly in barrel cortex while single whiskers were deflected with 0.5-18 air puffs per second (apps), a range that includes the whisk rates observed when rats explore their environment and discriminate surfaces with their whiskers. Twenty-nine neurons in layers III and IV were isolated in three rats (23 in barrel columns and 6 in septum columns). At < or = 9 apps, cortical neurons responded with one to two spikes per stimulus, whereas at > 9 apps, the response efficacy was reduced to only 0.2-0.4 spikes per stimulus. Several mechanisms are discussed that could account for the decrement in responsiveness. Despite this adaptation, neural spike rates increased in direct proportion with stimulus frequency when cast on logarithmic scales. At > 9 apps, however, this relationship deteriorated in barrel columns in which the response approximately halved. In contrast, septum column cells continued to increase their spike rates linearly up to 18 apps, although they responded at lower magnitude than the barrel column cells. Our findings suggest that septum column neurons are potential candidates to encode stimulus frequency using spike rate across the entire frequency range relevant to rats' whisking behavior.

Action Potentials↗

PubMed2006

Rapid plasticity follows whisker pairing in barrel cortex of the awake rat.

Synaptic plasticity can be induced easily throughout life in the rodent somatic sensory cortex. Trimming all but two whiskers on one side of an adult rat's face, called 'whisker pairing', causes the active (intact) whiskers to develop a stronger drive on cortical cells in their respective barrel columns, while inactive (trimmed) whisker efficacy is down-regulated. To date, this type of activity-dependent plasticity has been induced by trimming all but two whiskers, letting the rats explore their environment from 1 day to 1 month, after which cortical responses were analyzed physiologically under anesthesia. Such studies have enhanced our understanding of cortical plasticity, but the anesthesia complicates the examination of changes that occur in the first few hours after whisker trimming. Here we assayed the short-term changes that occur in alert, active animals over a period of hours after whisker trimming. The magnitude of barrel cortex evoked responses was measured in response to stimulation of the cut and paired whiskers of rats under several conditions: (a) whisking in air (control), (b) active whisking of an object by the rat, and (c) epochs of passive whisker stimulation to identify the onset of whisker pairing plasticity changes in cortex. The main difference between whisking in air without contact and passive whisker stimulation is that the former condition induces an increased response to stimulation of inactive cut whiskers, while the latter condition increases the responses to the stimulated whiskers. The results support the conclusion that whisker pairing plasticity in barrel cortex occurs within 4 h after whisker trimming in an awake, alert animal.

Animals↗

PubMed2006

Gestational 2,3,7,8-tetrachlorodibenzo-p-dioxin exposure effects on sensory cortex function.

Gestational exposure to environmental contaminants such as 2,3,7,8-tetrachlorodibenzo-p-dioxin (TCDD) poses a significant threat to normal growth and differentiation of the developing brain. To characterize the impact of gestational TCDD exposure on subsequent cortical function, pregnant Long Evans rats were exposed to a single acute dose (100 or 700ng/kg b.w. via gavage) on gestational day 15. This dosing regimen had no significant effect on birth index. After the TCDD-exposed animals were born and reached maturity, neural activity was recorded under urethane anesthesia from neurons in primary somatic sensory cortex. Spontaneous activity was reduced by approximately 50% in barrel cortex compared to corn oil vehicle controls. The magnitude of neuronal response to sensory (whisker) stimuli also was significantly reduced, and responses did not achieve control levels at any stimulus intensity. The greatest deficit was in the short latency component of the cortical responses. These decrements in cortical responsiveness were present in young F1 generation TCDD-exposed animals and persisted for up to 180 days. Because glutamate receptors are crucial to the evoked responses and show developmental regulation, selected iontotropic glutamate receptor subunits (NMDA NR2A+NR2B and GluR1) were profiled for RNA levels in the cortex of F1 generation rats. The expression of NR2B (NMDA receptor) and GluR1 (AMPA receptor) subunits was significantly reduced in the TCDD-exposed F1 generation animals compared to vehicle controls. The results indicate that gestational TCDD exposure results in cortical deficits that are paralled by diminished expression of certain NMDA and AMPA receptor subunits at a time when synapses are being formed for the first time in cortex.

Action Potentials↗

PubMed2006

Rate code and temporal code for frequency of whisker stimulation in rat primary and secondary somatic sensory cortex.

We recorded responses to frequencies of whisker stimulation from 479 neurons in primary (S1) and secondary (S2) somatic sensory cortex of 26 urethane-anesthetized rats. Five whiskers on the right side of the snout were deflected with air puffs at seven frequencies between 1 and 18/s. In left S1 (barrels and septa) and S2, subsets of neurons (5%) responded to whisker stimulation across the entire range of frequencies with > or = 1 electrical discharges/ten stimuli (full responders). In contrast, 60% of the recorded cells responded above threshold only at stimulus frequencies below 6/s and 35% remained subthreshold at all frequencies tested. Thus, the full responders are unique in that they were always responsive and appeared particularly suited to facilitate a dynamic, broadband processing of stimulus frequency. Full responders were most responsive at 1 stimulus/s, and showed greatest synchrony with whisker motion at 18 stimuli/s. The barrel cells responded with the greatest temporal accuracy between 3 and 15 stimuli/s. The septum cells responded less accurately, but maintained their accuracy at all frequencies. Only septum cells continued to increase their discharge rate with increasing stimulus frequency. The S2 cells discharged with lowest temporal accuracy modulated only by stimulus frequencies < or = 6/s and exhibited the steepest decrease in discharge/stimulus with increasing stimulus frequency. Our observations suggest that full responders in the septa are well suited to encode high frequencies of whisker stimulation in timing and rate of discharge. The barrel cells, in contrast, showed the strongest temporal coding at stimulus frequencies in the middle range, and S2 cells were most sensitive to differences in low frequencies. The ubiquitous decline in discharge/stimulus in S1 and S2 may explain the decrease in blood flow observed at increasing stimulus frequency with functional imaging.

Action Potentials↗

PubMed2006

Balancing bilateral sensory activity: callosal processing modulates sensory transmission through the contralateral thalamus by altering the response threshold.

Rats tactually explore a nearly spherical space field around their heads with their whiskers. The information sampled by the two sets of whiskers is integrated bilaterally at the cortical level in an activity dependent manner via the corpus callosum. We have recently shown that sensory activity in one barrel field cortex (BFC) modulates the processing of incoming sensory information to the other BFC. Whether interhemispheric integration is dynamically linked with corticothalamic modulation of incoming sensory activity is an important hypothesis to test, since subcortical relay neurons are directly modulated by cortical neurons through top-down processes. In the present study, we compared the direct sensory responses of single thalamic relay neurons under urethane anesthesia before and after inactivating the BFC contralateral to a thalamic neuron. The data show that silencing one BFC reduces response magnitude in contralateral thalamic relay neurons, significantly and reversibly, in response to test stimuli applied to the principal whisker at two times response threshold (2T) intensity for each unit. Neurons in the ventral posterior medial (VPM) nucleus and the medial division of the posterior nucleus (POm) react in a similar manner, although POm neurons are more profoundly depressed by inactivation of the contralateral BFC than VPM neurons. The results support the novel idea that the subcortical relay of sensory information to one hemisphere is strongly modulated by activity levels in the contralateral as well as in the ipsilateral SI cortex. The mechanism of the modulation appears to be based on shifting the stimulus-response curves of thalamic neurons, thereby rendering them more or less sensitive to sensory stimuli. We conclude that global sensory processing is created by combining activity in each cerebral hemisphere and continually balancing the flow of information to cortex by adjusting the responsiveness of ascending sensory pathways.

Action Potentials↗

PubMed2006

Two-photon imaging of cortical surface microvessels reveals a robust redistribution in blood flow after vascular occlusion.

A highly interconnected network of arterioles overlies mammalian cortex to route blood to the cortical mantle. Here we test if this angioarchitecture can ensure that the supply of blood is redistributed after vascular occlusion. We use rodent parietal cortex as a model system and image the flow of red blood cells in individual microvessels. Changes in flow are quantified in response to photothrombotic occlusions to individual pial arterioles as well as to physical occlusions of the middle cerebral artery (MCA), the primary source of blood to this network. We observe that perfusion is rapidly reestablished at the first branch downstream from a photothrombotic occlusion through a reversal in flow in one vessel. More distal downstream arterioles also show reversals in flow. Further, occlusion of the MCA leads to reversals in flow through approximately half of the downstream but distant arterioles. Thus the cortical arteriolar network supports collateral flow that may mitigate the effects of vessel obstruction, as may occur secondary to neurovascular pathology.

Animals↗

PubMed2006

A combined deficiency of vitamins E and C causes severe central nervous system damage in guinea pigs.

A short period of combined deficiency of vitamins E and C causes profound central nervous system (CNS) dysfunction in guinea pigs. For this report, CNS histopathology was studied to define the nature and extent of injury caused by this double deficiency. Weanling guinea pigs were fed a vitamin E-deficient or -replete diet for 14 d. Then vitamin C was withdrawn from the diet of some guinea pigs. Four diet groups were thus formed: replete, vitamin E deficient, vitamin C deficient, and both vitamin E and C deficient. From 5 to 11 d after institution of the doubly deficient diet, 9 of 12 guinea pigs developed paralysis, and 2 more were found dead. The remaining guinea pig in the doubly deficient group and all animals in the other 3 groups survived without clinical impairment until the experiment was terminated at 13-15 d. Brains and spinal cords were serially sectioned and stained for examination. Only the combined deficiency produced damage in the CNS. The damage consisted mainly of nerve cell death, axonal degeneration, vascular injury, and associated glial cell responses. The spinal cord and the ventral pons in the brainstem were most severely affected, often exhibiting asymmetric cystic lesions. Several features of the lesions suggest that the primary damage was to blood vessels. These results indicate that the paralysis and death caused by combined deficiency of vitamins E and C in guinea pigs is caused by severe damage in the brainstem and spinal cord.

Animals↗

PubMed2005

Chronic suppression of activity in barrel field cortex downregulates sensory responses in contralateral barrel field cortex.

Numerous lines of evidence indicate that neural information is exchanged between the cerebral hemispheres via the corpus callosum. Unilateral ablation lesions of barrel field cortex (BFC) in adult rats induce strong suppression of background and evoked activity in the contralateral barrel cortex and significantly delay the onset of experience-dependent plasticity. The present experiments were designed to clarify the basis for these interhemispheric effects. One possibility is that degenerative events, triggered by the lesion, degrade contralateral cortical function. Another hypothesis, alone or in combination with degeneration, is that the absence of interhemispheric activity after the lesion suppresses contralateral responsiveness. The latter hypothesis was tested by placing an Alzet minipump subcutaneously and connecting it via a delivery tube to a cannula implanted over BFC. The minipump released muscimol, a GABA(A) receptor agonist at a rate of 1 mul/h, onto one barrel field cortex for 7 days. Then with the pump still in place, single cells were recorded in the contralateral BFC under urethan anesthesia. The data show a approximately 50% reduction in principal whisker responses (D2) compared with controls, with similar reductions in responses to the D1 and D3 surround whiskers. Despite these reductions, spontaneous firing is unaffected. Fast spiking units are more sensitive to muscimol application than regular spiking units in both the response magnitude and the center/surround ratio. Effects of muscimol are also layer specific. Layer II/III and layer IV neurons decrease their responses significantly, unlike layer V neurons that fail to show significant deficits. The results indicate that reduced activity in one hemisphere alters cortical excitability in the other hemisphere in a complex manner. Surprisingly, a prominent response decrement occurs in the short-latency (3-10 ms) component of principal whisker responses, suggesting that suppression may spread to neurons dominated by thalamocortical inputs after interhemispheric connections are inactivated. Bilateral neurological impairments have been described after unilateral stroke lesions in the clinical literature.

Action Potentials↗

PubMed2005

Comparison of bilateral whisker movement in freely exploring and head-fixed adult rats.

Rats move their whiskers actively during tactile exploration of their environment. The whiskers emanate from densely innervated whisker follicles that are moved individually by intrinsic facial muscles and as a group by extrinsic muscles. Several descriptions of whisker movements in normal adult rats during unrestrained exploration indicate that rats move their whiskers in the 6-9 Hz range when exploring a new environment. The rate can be elevated to nearly 20 Hz for brief episodes just prior to making a behavioural decision. The present studies were undertaken to compare whisker dynamics in head-restrained and freely moving rats with symmetrical or asymmetrical numbers of whiskers on the two sides of their face and to provide a description of differences in whisker use in exploring rats after trimming all but two whiskers on one side of the face, a condition that has been shown to induce robust cortical plasticity. Head-fixed rats were trained to protract their whiskers against a contact detector with sufficient force to trigger a chocolate milk reward. Whisker movements were analyzed, and the results from head-fixed animals were compared with free-running animals using trials taken during their initial exploration of novel objects that blocked the rat's progress down an elevated runway. The results show that symmetrical whisker movements are modulated both by the nature of the task and the number of whiskers available for exploration. Rats can change their whisker movements when the sensitivity (threshold) of a contact detector is raised or lowered, or when the nature of the task requires bilateral input from the whiskers. We show that trimming some, but not all whiskers on one side of the face modifies the synchrony of whisker movement compared to untrimmed or symmetrically trimmed whiskers.

Animals↗

PubMed2004

Effect of subthreshold up and down states on the whisker-evoked response in somatosensory cortex.

Changes in spontaneous activity within the cortex recognized by subthreshold fluctuations of the membrane potential of cortical neurons modified the response of cortical neurons to sensory stimuli. Sensory stimuli occurring in the hyperpolarized "down" state evoked a larger depolarization and were more effective in evoking action potentials than stimuli occurring in the depolarized "up" state. Direct electrical stimulation of the thalamus showed the same dependence on the cell's state at the time of the stimulus, ruling out a strictly thalamic mechanism. Stimuli were more effective at triggering action potentials in the down state even during moderate de- or hyperpolarization of the somatic membrane potential. The postsynaptic potential (PSP) evoked from the down state was larger than the up state PSP but achieved about the same peak membrane potential, which was also near the reversal potential of the PSP (about -51 mV). Chloride loading shifted the reversal potentials of both the up state and the whisker-evoked PSP toward a more depolarized membrane potential. In addition, the threshold for action potentials evoked from the down state was lower than for spikes evoked in the up state. Thus the larger PSP from the down state may be caused by its larger driving force, and the state dependence of action potential generation in response to whisker stimulation may in part be related to a shift in threshold. Different mechanisms are therefore responsible for the state-dependence of PSP amplitude and the spike frequency response to the whisker stimulus.

Action Potentials↗

PubMed2004

The contribution of the principal and spinal trigeminal nuclei to the receptive field properties of thalamic VPM neurons in the rat.

Primary sensory information from neurons innervating whisker follicles on one side of a rat's face is relayed primarily through two subnuclei of the brainstem trigeminal complex to the contralateral thalamus. The present experiments were undertaken to separate the contribution of the principal trigeminal nucleus (PrV) from that of the spinal trigeminal nucleus (SpV) to whisker evoked responses in the ventral posterior medial (VPM) nucleus in the adult rat thalamus. Extracellular single-unit responses of VPM neurons to controlled stimulation of the contralateral whiskers under urethane anesthesia were quantified in terms of receptive field size, modal latency, response probability and response magnitude. The SpV contribution to VPM cell responses was isolated by making kainic acid lesions of the PrV. The PrV contribution was ascertained by cutting the trigeminothalamic axons arising from SpV just before they cross the midline. After destruction of the PrV, the SpV pathway alone produced large receptive fields (mean: 9.04 whiskers) and long latency (mean: 11.07 ms) responses from VPM neurons. In contrast, PrV input alone (SpV disconnected) generated small receptive fields (mean: 1.06 whiskers) and shorter latency (mean: 6.74 ms) responses. With both pathways intact the average receptive field size was 2.4 whiskers and peak (modal) response latency was 7.33 ms. The responses with both pathways intact were significantly different from either pathway operating in isolation. Response probability and magnitude followed the same trend. We conclude that normal responses of individual VPM neurons represent the integration of input activity transmitted through both PrV and SpV pathways.

Action Potentials↗

PubMed2003

Lesions of mature barrel field cortex interfere with sensory processing and plasticity in connected areas of the contralateral hemisphere.

Lesions of primary sensory cortex produce impairments in brain function as an outcome of the direct tissue damage. In addition, indirect lesion effects have been described that consist of functional deficits in areas sharing neural connections with the damaged area. The present study characterizes interhemispheric deficits produced as a result of unilateral lesions of the entire vibrissa representation of S-I barrel field cortex (BFC) in adult rats using single-neuron recording under urethane anesthesia. After unilateral lesions of adult BFC, responses of neurons in the contralateral homotopic BFC are severely depressed. Background (spontaneous) activity is reduced by approximately 80%, responses to test stimuli applied to the whiskers are reduced by approximately 50%, and onset of synaptic plasticity induced by trimming all but two whiskers ("whisker-pairing plasticity") is delayed over sevenfold compared with sham-lesion control animals. These deficits persist with only slight improvement for at least 4 months after lesion. Both fast-spiking and regular-spiking neuron responses are diminished contralateral to the lesion, as are cells above, below, and within the cortical barrels. Enriched environment experience increased the magnitude of responses and accelerated the rate of synaptic plasticity but did not restore response magnitude to control levels. Deficiencies in evoked responses and synaptic plasticity are primarily restricted to areas that share direct axonal connections with the lesioned cortex, because equivalently sized lesions of visual cortex produce minimal deficits in contralateral BFC function. These results indicate that interhemispheric deficits consist of remarkable and persistent decrements in sensory processing at the single-neuron level and support the idea that the deficits are somehow linked to the shared neural connections with the area of brain damage.

Action Potentials↗

PubMed2003

Experience-dependent plasticity is impaired in adult rat barrel cortex after whiskers are unused in early postnatal life.

The capacity of adult barrel cortex to show experience-dependent plasticity after early restricted neonatal sensory deprivation was analyzed in barrel field cortex neurons. Selective sensory deprivation was induced by trimming two whiskers from postnatal day 0 (P0) to P21, namely, the principal D2 whisker plus one adjacent surround whisker (D3). At maturity (P90), responses of supragranular (layer II/III) and barrel (layer IV) neurons, all located in the D2 barrel column, were analyzed for modified responses to the deprived principal whisker (D2) and the nondeprived (D1) and deprived (D3) adjacent surround whiskers. For supragranular neurons, the responses to both principal and surround whiskers were reduced at maturity, whereas the barrel neurons showed mildly elevated responses to the principal whisker but a reduced response to the deprived surround whisker. In normal adult rats, trimming all but the principal D2 whisker and an adjacent D3 whisker for 3 d (whisker pairing) produced the expected bias: elevated responses from the intact D3 compared with the cut D1 whisker in both barrel and supragranular neurons. When the neonatally deprived D2 and D3 whiskers were paired at maturity, a similar D3/D1 bias was generated in barrel neurons, but no bias occurred in supragranular neuron responses. Pairing the maintained D1 and deprived D2 whiskers produced a much greater bias toward D1 compared with the deprived D3 whisker in barrel neurons than in supragranular neurons. There were minimal effects on response latencies in layer IV under any of the experimental conditions. These findings indicate that a restricted period of sensory deprivation in early postnatal life (1) impairs intracortical relay of deprived inputs from layer IV to layer II/III in barrel cortex at maturity and (2) degrades receptive field plasticity of the supragranular layer cells but not the thalamic-recipient barrel neurons.

Afferent Pathways↗

PubMed2003

Experimental model for functional magnetic resonance imaging of somatic sensory cortex in the unanesthetized rat.

Functional magnetic resonance imaging (fMRI) has evolved into a method widely used to map neural activation in the human brain. fMRI is a method for recording blood oxygen level-dependent (BOLD) signals. These signals change with local cerebral blood flow coupled to neural activity. However, the relationship between BOLD signals and neural function is poorly understood and requires the development of animal models. Here we use an unanesthetized rat preparation to study BOLD responses to whisker stimulation in somatic sensory barrel cortex. Five rats were trained to tolerate restraint in a holder and fMRI noise with positive reinforcement. For maximal immobilization, the head was fastened to the holder with nuts screwed on threaded bolts attached to the head. On scanning day, residual stress was alleviated with injections of diazepam, and the rats were restrained in the holder and transferred into the scanner. After >75 min to allow the tranquilization to abate, structural images were acquired from three coronal brain slices. Subsequently, functional images were taken utilizing 4-min epochs without stimulation alternated with equivalent epochs during which the right caudal whiskers were stimulated with three air puffs/s. After 4 weeks, fMRI could be repeated in four rats. In seven of the nine functional runs, head motion was minimal and whisker stimulation resulted in a statistically significant (P </= 0.05) increase in BOLD signal in barrel cortex predominantly on the contralateral side. The results provide encouragement that long-term fMRI studies on cerebral function in unanesthetized rats may be feasible with our procedure.

Animals↗

PubMed2003

Unilateral vibrissa contact: changes in amplitude but not timing of rhythmic whisking.

Electromyographic recordings from the mystacial pad of rats were used to assess the effect of unilateral vibrissa contact on the bilateral movement of the vibrissae. A first group of animals was trained to whisk freely in air and served to establish the baseline variability in bilateral symmetry. We observed that the electromyogram (EMG) activity across the two mystacial pads was rhythmic and synchronous to within 2 ms on a whisk-by-whisk basis; this value is small in comparison with the approximately 50 ms required for protraction during the whisk cycle. A second group of animals was trained to use their vibrissae to contact a sensor that was located on one side of the head. The average EMG activity across the two pads was synchronous at the time of vibrissa contact, albeit with higher variability than for the case of free whisking. In contrast, the average amplitude of the activity on the contact vs noncontact side of the face was transiently greater, by 25% or approximately 10 degrees, at the time of contact. These data show that the amplitude of the vibrissae on the two sides of the face can be controlled independently, while the timing of vibrissa movement is largely synchronous.

Animals↗

PubMed2002

Adaptive filtering of vibrissa input in motor cortex of rat.

We studied the transformation of sensory input as it progresses from vibrissa primary sensor (S1) to motor (M1) cortex. Single-unit activity was obtained from alert adult rats that did not to whisk upon application of punctate, rhythmic stimulation of individual vibrissae. The spike response of units in S1 cortex largely reproduced the shape of the stimulus. In contrast, the spiking output of units in M1 cortex were modulated solely as a sinusoid at the repetition rate of the stimulus for frequencies between 5 and 15 Hz; this range corresponds to that of natural whisking. Thus, the S1 to M1 transformation extracts the fundamental frequency from a spectrally rich stimulus. We discuss our results in terms of a band-pass filter with a center frequency that adapts to the change in stimulation rate.

Action Potentials↗

PubMed2002

Divergent movement of adjacent whiskers.

The current view of whisker movement is that approximately 25 whiskers on each side of the face move in synchrony. To determine whether whiskers are constrained to move together, we trained rats to use two whiskers on the same side of the face in simple behavioral tasks and videotaped the whiskers during the task. Here we report that the movement of adjacent whiskers is usually synchronous but can diverge: 1) the distance between whiskers can vary dramatically during movement; 2) one whisker can move while the second one remains stationary; 3) two whiskers can simultaneously move in opposite directions; and 4) one whisker can be maintained in contact with an object while the other is retracted and protracted. The frequency of whisker movement during the task falls within the previously reported range for rats whisking freely into air or performing roughness discrimination with their whiskers. Our data also suggest that whisker movement can be divided into three distinct phases: protraction, retraction, and a measurable delay between these movements. We conclude that, although whiskers often move in concert, adjacent caudal whiskers can be moved independently of each other.

Animals↗