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J Neurophysiol (January 1, 2003). 10.1152/jn.00600.2002
Submitted on Submitted 24 July 2002; accepted in final form 24 September 2002
1Department of Physics and 2Neurosciences Graduate Program, University of California at San Diego, La Jolla, California 92093
Berg, Rune W. and
David Kleinfeld.
Rhythmic Whisking by Rat: Retraction as Well as Protraction of
the Vibrissae Is Under Active Muscular Control. J. Neurophysiol. 89: 104-117, 2003. The
rhythmic motor activity of the vibrissae that rodents use for the
tactile localization of objects provides a model system for
understanding patterned motor activity in mammals. The muscles that
drive this whisking are only partially fixed relative to bony
attachments and thus shift their position along with the movement. As a
means to characterize the pattern of muscular dynamics during different
patterns of whisking, we recorded electromyogram (EMG) activity from
the muscles that propel individual follicles, as well as EMG activity
from a muscle group that moves the mystacial pad. The dominant pattern
of whisking in our behavioral paradigm, referred to as exploratory
whisking, consisted of large amplitude sweeps in the frequency range of
5-15 Hz. The frequency remained remarkably constant within a bout of
whisking but changed values between bouts. The extrinsic musculature,
which shifts the surface of the pad backwards, was found to be
activated in approximate antiphase to that of the intrinsic muscles,
which rotate individual vibrissae forward. Thus retraction of the
vibrissae was driven by a backward shift in the attachment point of the
follicles to the mystacial pad. In a less frequent pattern of whisking,
referred to as foveal whisking, the vibrissae are thrust forward and
palpate objects with low-amplitude movements that are in the higher
frequency range of 15-25 Hz. Protraction of the vibrissae remains
driven by the intrinsic muscles, while retraction in this pattern is largely passive. Interestingly, a mechanical argument suggests that
activation of the extrinsic muscles during foveal whisking is not
expected to affect the angle of the vibrissae. As a means to establish
if the phasic control of the intrinsic versus extrinsic muscles
depended on sensory feedback, we characterized whisking before and
after bilateral transections of the infraorbital branch of the
trigeminal sensory nerve. The loss of sensory feedback had no net
effect on the antiphase relation between activation of the intrinsic
versus extrinsic muscles over the full frequency range for exploratory
whisking. These data point to the existence of a dual-phase central
pattern generator that drives the vibrissae.
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