The relationship of vibrissal motor cortex unit activity to whisking in the awake rat
Abstract
Rats actively sweep their whiskers back and forth to locate and palpate objects within their immediate environment. Microstimulation studies in anesthetized rats have demonstrated the presence of a large vibrissal motor representation in agranular cortex. However, the manner in which motor cortex neurons contribute to whisking behavior in the awake animal is unknown. This study represents an initial investigation of the relationship between the activity of task-related neurons in vibrissal motor cortex and the electromyographic (EMG) activity of the deep vibrissal pad muscles in the awake, freely whisking rat. Each animal was gently held in an experimenter's hands while the animal whisked the air. A spring-loaded, metal microelectrode mounted in a removable, miniature microdrive provided stabile recordings of motor cortex unit activity. Fine-wire electrodes implanted in the mystacial pad simultaneously recorded facial muscle activity. Results showed that the discharge of task-related neurons was correlated with changing levels of muscle output. Unit discharge was related in a tonic or phasic-tonic fashion to EMG activity. No units were found to discharge rhythmically in a 1:1 fashion with the periodicity of the whisking pattern. These findings support a role for vibrissal motor cortex in the initiation and modulation of the overall level of mystacial pad muscular output, but not in the generation of bursts of EMG activity responsible for individual whisking sweeps.
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At the single-cell spiking level, a subset of neurons increases firing rate just prior (~1 s) to locomotion onset.
“At the level of single-cell spiking, an increase in firing rate can be observed in a subset of neurons just prior (~1 s) to the onset of locomotion”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Carvell et al., 1996
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BackgroundResultNew Modules Are Added to Vibrissal Premotor Circuitry with the Emergence of Exploratory Whisking
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“This observation led to the hypothesis that whisking-related activity in primary motor cortex [49,72] might gate PO thalamus so that it is sensitive to whisking.”
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“…neurons, whose activity significantly correlated with whisking kinematics, but- in contrast to the primate (Georgopoulos et al., 1982, 1986)- there was no relationship between population activity and whisking (Carvell et al., 1996; Hill et al., 2011; Friedman et al., 2012; Gerdjikov et al., 2013).”
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“A miniature microdrive assembly was positioned over the appropriate area (for review, see Carvell et al., 1996).”
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BackgroundCortical Control of a Whisking Central Pattern Generator
Cramer, N., & Keller, A. (2006) · Journal of Neurophysiology · doi:10.1152/jn.00071.2006 · Cited by 81 · see citing works
“Other evidence suggests that vMCx does not drive vibrissa motoneurons directly (Carvell et al. 1996), but rather that vMCx acts through its interactions with a subcortical whisking CPG (Friedman et al.”
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BackgroundMethodCortical Control of Whisker Movement
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“Whisker movements can entrain rhythmic activity in wMCx neurons (Kleinfeld et al., 1999) and activity in wMCx neurons has been recorded during periods of exploratory whisking (Carvell et al., 1996).”
BackgroundThe Whisking Rhythm Generator: A Novel Mammalian Network for the Generation of Movement
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“The activity of vMCx neurons does not co-vary with whisking frequency, suggesting that vMCx does not directly drive vMNs (Carvell et al. 1996).”
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“It remains to be determined whether the effects on M1 representations observed in the birth-trimmed group are reversible, but the observation that such animals do not regain normal discriminative capacity, even with months of renewed vibrissa growth ( Carvell et al., 1996 ), suggests that they are not (but see Keller…”
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“…from the neocortex in the rat during behavior and paved the way for more recent work on the relation between electrical activity in the neocortex and sensory-motor tasks (Taube et al., 1990; Chen et al., 1994; McNaughton et al., 1994; Nicolelis et al., 1995; Carvell et al., 1996; Fee et al., 1997).”
BackgroundVibrissa motor cortex activity suppresses contralateral whisking behavior
Ebbesen, C. L., Doron, G., Lenschow, C., & Brecht, M. (2016) · Nature Neuroscience · doi:10.1038/nn.4437 · Cited by 47 · see citing works
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“Previous studies of deeper layers have observed both increases and decreases in firing rates of somatosensory (Curtis and Kleinfeld, 2009; de Kock and Sakmann, 2009) and motor (Carvell et al., 1996; Schiemann et al., 2015; Zagha et al., 2015) cortical neurons duringmovement.”
Vibrissae motor cortex unit activity during whisking
Friedman, W. A., Zeigler, H. P., & Keller, A. (2011) · Journal of Neurophysiology · doi:10.1152/jn.01132.2010 · Cited by 32 · see citing works
“Extracellular recordings from single vMCx neurons (Carvell et al. 1996) and recordings of local field potentials (LFPs) (Friedman et al.”
BackgroundMethodInfluentialShaping the Cortical Landscape: Functions and Mechanisms of Top-Down Cortical Feedback Pathways
Zagha, E. (2020) · Frontiers in Systems Neuroscience · doi:10.3389/fnsys.2020.00033 · Cited by 39 · see citing works
Response durations encode nociceptive stimulus intensity in the rat medial prefrontal cortex
Zhang, R., Tomida, M., Katayama, Y., & Kawakami, Y. (2004) · Neuroscience · doi:10.1016/j.neuroscience.2004.01.055 · Cited by 33 · see citing works
Anticipatory Activity of Motor Cortex in Relation to Rhythmic Whisking
Friedman, W. A., Jones, L. M., Cramer, N., Kwegyir-Afful, E. E., Zeigler, H. P., & Keller, A. (2005) · Journal of Neurophysiology · doi:10.1152/jn.00945.2005 · Cited by 30 · see citing works
“The longer pre-movement LFP activity, compared with that reported by Carvell et al. (1996) is likely due to the fact that LFPs represent temporally integrated synaptic activity from many neurons, whereas unit activity reports discrete spikes from individual neurons.”
InfluentialPersistence of vibrissal motor representation following vibrissal pad deafferentation in adult rats
Franchi, G. (2001) · Experimental Brain Research · doi:10.1007/s002210000652 · Cited by 30 · see citing works
“Since the motor cortex appeared involved in the high degree of whisking control (Carvel et al. 1996), the increase in electrical stimulation threshold after deafferentation could be an electrophysiological correlate of the impairment of sensorimotor coordination of whisking observed in deafferented…”
BackgroundAruljothi, K., Marrero, K., Zhang, Z., Zareian, B., & Zagha, E. (2020) · Journal of Neuroscience · doi:10.1523/jneurosci.2993-19.2020 · Cited by 27 · see citing works
Martinetti, L. E., Bonekamp, K. E., Autio, D. M., Kim, H.-H., & Crandall, S. R. (2021) · Cerebral Cortex · doi:10.1093/cercor/bhab325 · Cited by 26 · see citing works
Cellular mechanisms of motor control in the vibrissal system
Brecht, M., Grinevich, V., Jin, T.-E., Margrie, T. W., & Osten, P. (2006) · Pflügers Archiv - European Journal of Physiology · doi:10.1007/s00424-006-0101-6 · Cited by 27 · see citing works
“In line with this idea, unitrecording studies in VMC [60] did not observe a one to Pflugers Arch - Eur J Physiol (2006) 453:269–281 275”
ResultCircuit mechanisms for cortical plasticity and learning
Chéreau, R., Williams, L. E., Bawa, T., & Holtmaat, A. J. D. G. (2021) · Seminars in Cell and Developmental Biology · doi:10.1016/j.semcdb.2021.07.012 · Cited by 25 · see citing works
Gerdjikov, T. V., Haiss, F., Rodríguez-Sierra, O. E., & Schwarz, C. (2013) · Journal of Neuroscience · doi:10.1523/jneurosci.0337-13.2013 · Cited by 26 · see citing works
“…to be in RW. Friedman et al. (2006, 2012) have characterized their recording sites using prolonged microstimulation, and the pioneering study of Carvell et al. (1996) recorded at sites that showed protraction flicks in response to short microstimulation indicative of RW (Haiss and Schwarz,…”
BackgroundMethodResultInfluentialCortical barrel field ablation and unconditioned whisking kinematics
Harvey, M. A. (2001) · Somatosensory & Motor Research · doi:10.1080/01421590120072213 · Cited by 26 · see citing works
Vibrissa Myoclonus (Rhythmic Retractions) Driven by Resonance of Excitatory Networks in Motor Cortex
Castro‐Alamancos, M. A. (2006) · Journal of Neurophysiology · doi:10.1152/jn.00454.2006 · Cited by 26 · see citing works
Proceedings of the Barrels II Workshop: sensorimotor loops
Hattox, A. M. (2003) · Somatosensory & Motor Research · doi:10.1080/08990220310001623031 · Cited by 22 · see citing works
Dash, S., Autio, D. M., & Crandall, S. R. (2022) · Journal of Neuroscience · doi:10.1523/jneurosci.2219-21.2022 · Cited by 21 · see citing works
Movement, Confusion, and Orienting in Frontal Cortices
Brecht, M. (2011) · Neuron · doi:10.1016/j.neuron.2011.10.002 · Cited by 21 · see citing works
“The predominance of cells concerned with slow movement time scales is in line with an earlier recording study, which also showed that cells did not covary 1:1 with the whisking rhythm and that cells would globally turn off and on with whisking (Carvell et al., 1996).”
ResultBeak, S. K., Hong, E. Y., & Lee, H. S. (2010) · Brain Research · doi:10.1016/j.brainres.2010.03.100 · Cited by 17 · see citing works
Gao, P. P., Ploog, B. O., & Zeigler, H. P. (2003) · Somatosensory & Motor Research · doi:10.1080/08990220310001623031-411 · Cited by 17 · see citing works
Khateb, M., Schiller, J., & Schiller, Y. (2017) · eLife · doi:10.7554/elife.21843 · Cited by 15 · see citing works
“Yet previous studies have shown that the activity of both vM1 and vS1 is phased locked with exploratory rhythmic whisking movements in rats (Ahrens and Kleinfeld, 2004), and vM1 neurons are probably critical in rhythmically driving whisking (Carvell et al., 1996; Kleinfeld et al., 2002; Ahrens and Kleinfeld, 2004; Bre…”
BackgroundCortical Localization of the Sensory-Motor Transformation in a Whisker Detection Task in Mice
Zareian, B., Zhang, Z., & Zagha, E. (2021) · eNeuro · doi:10.1523/eneuro.0004-21.2021 · Cited by 15 · see citing works
“…are consistent with the choice-related signals in wMC and ALM initiating the whisking and licking response sequence, respectively. wMC is a frontal region traditionally studied in the context of whisking initiation and modulation (Carvell et al., 1996; Kleinfeld et al., 1999; Hill et al., 2011).”
BackgroundFranchi, G., & Veronesi, C. (2006) · The Journal of Physiology · doi:10.1113/jphysiol.2006.109116 · Cited by 15 · see citing works
“…rhythmic whisking is maintained by the whisking central pattern generator (CPG) at the level of medulla (Gao et al. 2001, 2003; Hattox et al. 2003), whereas voluntary initiation and modulation of the whisking pattern are mediated by cortical mechanisms (Carvel et al. 1996; Friedman et al. 2006).”
BackgroundFranchi, G., & Veronesi, C. (2004) · European Journal of Neuroscience · doi:10.1111/j.1460-9568.2004.03653.x · Cited by 13 · see citing works
“Experimental evidence supports a role for vibrissal motor cortex in the initiation and modulation of the mystacial pad muscular activity during sniffing behaviour ( Carvel et al., 1996 ) but not in the generation of the rhythmical whisking sweeps (Sharp & Evans, 1982; Isokawa-Akerson & Komiasaruk, 1987; Gao et al., 20…”
BackgroundKim, N., Bahn, S.-K., Choi, J. H., Kim, J. S., & Rah, J. (2021) · Cerebral Cortex · doi:10.1093/cercor/bhab236 · Cited by 11 · see citing works
Neuroplasticity of face sensorimotor cortex and implications for control of orofacial movements
Avivi‐Arber, L., Lee, J., Yao, D., Adachi, K., & Sessle, B. J. (2009) · Japanese Dental Science Review · doi:10.1016/j.jdsr.2009.11.007 · Cited by 11 · see citing works
“[2,6—10]) as noted below.”
BackgroundZhuang, K., Lebedev, M., & Nicolelis, M. A. L. (2014) · Journal of Neurophysiology · doi:10.1152/jn.00031.2013 · Cited by 10 · see citing works
“Additionally, STA and cross-correlation analysis have revealed task-dependent entrainment of EMG activity to oscillations of cortical field potentials in rodents (Carvell et al. 1996; Nicolelis et al. 1995), nonhuman primates (Baker et al.”
BackgroundSanger, T. D. (2002) · Journal of Neurophysiology · doi:10.1152/jn.00121.2001 · Cited by 11 · see citing works
“This problem also occurs when using the instantaneous spike rate (Carvell et al. 1996).”
BackgroundBrecht, M. (2004) · Journal of Neurophysiology · doi:10.1152/jn.00404.2004 · Cited by 7 · see citing works
Motor Control of Distinct Layer 6 Corticothalamic Feedback Circuits
Martinetti, L. E., Autio, D. M., & Crandall, S. R. (2024) · eNeuro · doi:10.1523/eneuro.0255-24.2024 · Cited by 6 · see citing works
“Since vM1 activity correlates with vibrissa movement (Carvell et al., 1996; Hill et al., 2011; Sreenivasan et al., 2016), vM1 projections might function to communicate the motor plan to vS1 (Kleinfeld et al., 2006).”
Kim, H.-H., Bonekamp, K. E., Gillie, G. R., Autio, D. M., Keller, T., & Crandall, S. R. (2024) · eNeuro · doi:10.1523/eneuro.0154-24.2024 · Cited by 5 · see citing works
“Neural activity in M1 codes for various whisking parameters (Carvell et al., 1996; Brecht et al., 2004; Hill et al., 2011; Friedman et al., 2012; Huber et al., 2012; Petreanu et al., 2012; Sreenivasan et al., 2016), and the projections from M1 are generally thought to communicate these motor…”
Hong, E. Y., Beak, S. K., & Lee, H. S. (2010) · Brain Research · doi:10.1016/j.brainres.2010.07.093 · Cited by 5 · see citing works
Avivi‐Arber, L. (2009) · TSpace (University of Toronto) · Cited by 4 · see citing works
Spectral analysis of whisking output via optogenetic modulation of vibrissa cortex in rat
Pashaie, R., & Falk, R. (2012) · Biomedical Optics Express · doi:10.1364/boe.4.000122 · Cited by 3 · see citing works
Castro-Alamancos, M. (2007) · Journal of Neurophysiology · doi:10.1152/jn.01358.2006 · Cited by 1 · see citing works
“Because active whisking does not require sensory feedback (Gao et al. 2001; Welker 1964) or the motor cortex (Carvell et al. 1996; Gao et al., 2003; Semba and Komisaruk 1984), a circuit located in the brain stem is assumed to generate whisking (Hattox et al. 2003).”
BackgroundCELL TYPE SPECIFIC CONNECTIONS FROM PRIMARY MOTOR TO PRIMARY SOMATOSENSORY CORTEX
Kinnischtzke, A. K. (2013) · D-Scholarship@Pitt (University of Pittsburgh) · Cited by 2 · see citing works
Kim, N., Bahn, S., Choi, J. H., Kim, J. S., & Rah, J. (2020) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2020.11.08.363200 · Cited by 1 · see citing works
“On the other hand, vM1 appears to deliver signals related 53 to muscle output change, such as initiation and delicate control of whisking 54 movement (Carvell et al., 1996; Diamond et al., 2008; Ferezou et al., 2007; 55 Sreenivasan et al., 2016), to the barrel field of S1 (S1BF).”
BackgroundPrete, Z. D., Baker, S. P., Grigg, P., Kim, B., Basso, M. A., Lee, J., Kim, H. R., & Lee, C. (2015)
Kleinfeld, D., Friedman, W. A., Zeigler, H. P., Keller, A., Méndez, J. M., Mindlin, G. B., Goller, F., Lottem, E., Gugig, E., & Azouz, R. (2015)
Sood, M. (2014) · TSpace (University of Toronto)
Faculty Opinions recommendation of Motor modulation of afferent somatosensory circuits.
Sherman, S. M., & Lee, C. C. (2009) · Faculty Opinions – Post-Publication Peer Review of the Biomedical Literature · doi:10.3410/f.1138869.617243
Ansorge, J., & Manahan‐Vaughan, D. (2025) · iScience · doi:10.1016/j.isci.2025.114427
“59 This finding also suggests that the S1L6 Ntsr1-pathway to VPM does not play an important role in response tuning to whisker deflection onset and thus, to deviant detection.”
Motor cortex regulation of thalamic -cortical activity in the somatosensory system
Lee, S. (2007) · D-Scholarship@Pitt (University of Pittsburgh)
Behavioral Context Shapes Sensory Responses in Vibrissal Motor CortexPreprint
Freitag, F., Vries, J. D., Harder, L.-G., Larkum, M. E., & Sachdev, R. N. S. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.04.28.720072
- Authors
- Carvell, G. E., Miller, S. A., & Simons, D. J.
- Affiliations
- University of Pittsburgh (US)
- Corresponding author
- Carvell, George E.
- Venue
- Somatosensory & Motor Research
- Publisher
- Informa UK Limited
- Published
- 1996-01
- Volume
- 13
- Issue
- 2
- Pages
- 115-127
- ISSN
- 0899-0220 (ISSN-L)
- 1369-1651
- OpenAlex
- W2165970664
- Cited by
- 82
- References
- 49
- Citations by year
- 2026: 1 · 2025: 1 · 2024: 2 · 2022: 1 · 2021: 4 · 2020: 3 · 2018: 1 · 2017: 2 · 2016: 3 · 2015: 5 · 2014: 3 · 2013: 6 · 2012: 3
- FWCI
- 1.18
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Somatosensory & Motor Research (publishedVersion)
- PubMed (publishedVersion)
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Tactile and Sensory Interactions
- Life Sciences › Neuroscience › Cognitive Neuroscience › Motor Control and Adaptation
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neural dynamics and brain function
- Keywords
- whisking, rats
- MeSH terms
- Animals, Brain Mapping, Electromyography, Female, Motor Cortex, Motor Neurons, Orientation, Vibrissae, Wakefulness, Rats
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