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Lora J. Pittman and E.

Fiona Bailey
J Neurophysiol 101:276-282, 2009. First published Nov 5, 2008; doi:10.1152/jn.91065.2008

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J Neurophysiol 101: 276 –282, 2009.
First published November 5, 2008; doi:10.1152/jn.91065.2008.

Genioglossus and Intrinsic Electromyographic Activities in Impeded


and Unimpeded Protrusion Tasks

Lora J. Pittman and E. Fiona Bailey


Department of Physiology, College of Medicine, The University of Arizona, Tucson, Arizona
Submitted 19 September 2008; accepted in final form 28 October 2008

Pittman LJ, Bailey EF. Genioglossus and intrinsic electromyo- constitute a hydrostatic “skeleton” (Kier and Smith 1985;
graphic activities in impeded and unimpeded protrusion tasks. J Skierczynski et al. 1996; Smith and Kier 1989) and provide the
Neurophysiol 101: 276 –282, 2009. First published November 5, reaction forces needed for tongue movement. In this respect,
2008; doi:10.1152/jn.91065.2008. Eight muscles invest the human
tongue: four extrinsic muscles have external origins and insert into the tongues, trunks, and tentacles offer a novel framework within
tongue body and four intrinsic muscles originate and terminate within which to consider the nervous system control of movement
the tongue. Previously, we noted minimal activation of the genioglos- because cooperative contraction of muscles underpins move-
sus tongue muscle during impeded protrusion tasks (i.e., having ment and provides the structural platform necessary for deliv-
subjects push the tongue against a force transducer), suggesting that ery of external force.

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other muscles play a role in the production of tongue force. Accord- The human tongue comprises orthogonally related intrinsic
ingly, we sought to characterize genioglossus tongue muscle activities
muscles (verticalis, transversus, superior and inferior longitu-
during impeded and unimpeded protrusion tasks (i.e., having subjects
slowly and smoothly move the tongue out of their mouth). Electro- dinalis) that originate and terminate within the tongue and
myographic (EMG) and single motor-unit potentials of the extrinsic so-called extrinsic muscles (genioglossus, hyoglossus, stylo-
genioglossus muscle were recorded with tungsten microelectrodes and glossus, and palatoglossus) that have external bony origins and
EMG activities of intrinsic tongue muscles were recorded with hook- insert into the tongue body. The muscular hydrostat theory
wire electrodes inserted into the anterior tongue body. Tongue posi- (Kier and Smith 1985) posits a constant interaction of extrinsic
tion was detected by an isotonic transducer coupled to the tongue tip. and intrinsic muscles in all tongue functions.
Protrusive force was detected by a force transducer attached to a rigid
bar. Genioglossus and intrinsic tongue muscles were simultaneously
To begin studying the highly complex muscular interactions
active in both impeded and unimpeded protrusion tasks. Genioglossus that underlie tongue movements, relatively simple movements
whole muscle EMG and single motor-unit activities changed faith- must be studied first. Accordingly, this study focuses on the
fully as a function of tongue position, with increased discharge muscular actions leading to tongue protrusion under conditions
associated with protrusion and decreased discharge associated with where protrusion is either impeded (i.e., quasi-isometric con-
retraction back to the rest position. In contrast, during the impeded tractions) or unimpeded (simple displacement). According to
protrusion task drive the genioglossus muscle remained constant as the muscular hydrostat theory (Smith and Kier 1989), protru-
protrusion force increased. Conversely, intrinsic tongue muscle activ-
ities appropriately followed changes in both tongue position and force. sion does not result from the activation of a single muscle but
Importantly, we observed significantly higher levels of intrinsic mus- rather is a consequence of expansion in the long axis (via
cle activity in the impeded protrusion task. These observations suggest contraction of intrinsic tongue muscles verticalis and transver-
that protrusion of the human tongue requires activation of the genio- sus) and forward movement in space (via contraction of ge-
glossus and intrinsic protrudor muscles, with the former more impor- nioglossus). Importantly, the multivectorial actions of the in-
tant for establishing anterior–posterior tongue location and the latter trinsic tongue muscles are thought to stiffen the tongue, mak-
playing a greater role in the generation of protrusive force. A biome- ing the development of protrusive force possible (Kier and
chanical model of these actions is provided and discussed. Smith 1985). Although this theory appears to hold in various
animal species (Abd-El-Malek 1938; Bennett and Hutchinson
INTRODUCTION 1946; Smith 1984), to date there is no evidence that protrusion
of the human tongue requires activation of intrinsic and extrin-
Our understanding of the principles governing the nervous sic tongue muscles. Accordingly, in the present study we
system control of movement in large part comes from studies recorded electromyographic (EMG) and single motor-unit ac-
conducted in muscles of the limbs that have clearly defined tivities from the extrinsic protrudor muscle, the genioglossus,
origins and insertions and that produce force across a joint. In as well as the EMG of intrinsic protrudor muscles in healthy
some instances, however, animals use structures composed human subjects that performed impeded or unimpeded protru-
primarily of muscle both to move and to apply force, examples sion tasks. Our main hypothesis is that tongue protrusion in
of which include the elephant trunk, the tentacles of a squid, human subjects requires activation of intrinsic and extrinsic
and the mammalian tongue. These structures, known as “hy- tongue muscles. Our secondary hypothesis is that impeded and
drostats,” are capable of remarkably complex movements ac- unimpeded protrusion tasks are associated with distinct muscle
complished without the aid of a solid skeleton. That is, muscles activation patterns, attributed to the presence or absence of a
modulate the internal pressure and stiffness of the structure to reaction force.

Address for reprint requests and other correspondence: E. F. Bailey, De- The costs of publication of this article were defrayed in part by the payment
partment of Physiology, College of Medicine, The University of Arizona, of page charges. The article must therefore be hereby marked “advertisement”
Tucson, AZ 85721-0093 (E-mail: ebailey@u.arizona.edu). in accordance with 18 U.S.C. Section 1734 solely to indicate this fact.

276 0022-3077/09 $8.00 Copyright © 2009 The American Physiological Society www.jn.org
EMG ACTIVITY IN IMPEDED AND UNIMPEDED PROTRUSION TASKS 277

METHODS cannot state with certainty which of the muscles we recorded from.
Thus in the remainder of this study we refer to this EMG recording as
We performed experiments in 10 healthy human volunteers (6 women
the “intrinsic muscle” EMG, although most of the activity likely
and 4 men; ages 20 –54 yr). All experimental procedures were approved
originates from the verticalis and transversus muscles.
by the Human Subjects Committee at The University of Arizona. Sub-
Surface electrodes (4-mm-diameter Ag/AgCl) attached to the skin
jects gave their informed consent before participation in the study.
overlying each mastoid process served as indifferent electrodes for
tungsten and fine-wire EMG recordings. A ground strap was placed
EMG recordings around the upper arm. EMG signals were amplified (⫻1,000, Model
EMG activities were recorded from the genioglossus (GG) muscle 15; Grass Instruments, West Warwick, RI) and band-pass filtered
and from the intrinsic tongue muscles. The GG muscle arises from the (0.3–3 kHz). Intrinsic and extrinsic whole muscle EMG activities
medial aspect of the mandibular symphysis and fans out dorsoven- were sampled at 10 kHz and GG SMUAPs were sampled at 20 kHz
trally to insert into the central mass of the tongue (Takemoto 2001). and recorded on the Spike2 data acquisition and analysis system
The protocol for electrode insertion has been reported previously (CED, Cambridge, UK). The amplified and filtered whole muscle
(Bailey et al. 2007a,b). Briefly, we determined the distance from the EMG signals were subsequently rectified and moving-time averaged
skin surface to the inferior border of the GG muscle via ultrasonog- with a time constant of 200 ms.
raphy (Pro Sound 3500; Aloka, Tokyo, Japan) (Eastwood et al. 2003).
For whole muscle GG EMG activities, a tungsten needle electrode (1- Experimental protocol
to 5-␮m tip diameter, 250-␮m shaft diameter; FHC, Bowdoinham,
ME) with insulation removed from the terminal tip (⬃2 mm) was Subjects were seated upright in a dental chair with their head
inserted transcutaneously into the floor of the mouth, with the entry supported. Respiratory movements of the chest were monitored by a
point about 1.5 cm from the midline and about 2– 4 cm posterior to the respiratory effort transducer (Pro-Tech, Mukilteo, WA) positioned

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mandible. A second high-impedance tungsten microelectrode (10 M⍀ around the rib cage at the level of the axilla and around the abdomen
at 1 kHz, 1- to 5-␮m tip diameter, 250-␮m shaft diameter; FHC) was at the level of the umbilicus.
inserted to the same depth for purposes of recording GG muscle single MAXIMUM MANEUVERS. Subjects first performed a series of maxi-
motor-unit action potentials (SMUAPs). mum maneuvers for purposes of normalizing EMG activities, allow-
We recorded intrinsic whole muscle EMG activities from the ing us to compare drive to the muscle across subjects. These maneu-
anterior tongue body with an intramuscular fine-wire electrode (50 vers comprised maximum protrusions of the tongue out of the mouth
␮m; California Finewire, Grover Beach, CA) with insulation removed and maximum forces exerted against the hard palate, against the lower
from the terminal tip (⬃2 mm). The wire was threaded through a incisors and against a tongue depressor that an investigator held at the
27-gauge needle and inserted through the dorsum of the tongue, subject’s lips. Interestingly, in all subjects, maximum GG EMG
posterior to the tongue tip at the first premolar, at a point equidistant activities were recorded during unimpeded protrusions.
from the medial sulcus and the lateral border and at depths of about
5–7 mm from the mucosal surface. After proper location was docu- UNIMPEDED PROTRUSIONS. Tongue position was detected by a bat-
mented, the needle was removed and the wire remained in the muscle tery-powered isotonic transducer (Harvard Apparatus, Kent, UK)
belly. The target muscles in this case were the verticalis and trans- (Bailey et al. 2007b). The transducer was connected to the tongue by
versus that, as their names suggest, are vertically and transversely a customized lever arm that engaged a thermoplastic housing affixed
oriented muscles of the tongue core; these muscles are thought to play to the dorsum of the tongue (Fig. 1A). This coupling posed minimal
a major role in tongue protrusion. Note, however, that due to the impediment to movement and subjects were able to swallow and
complex arrangement of muscle fibers within the tongue body we speak during the experiment. The position signal was amplified

FIG. 1. Schematic of the experimental


setup. A: tongue position was detected by a
battery-powered isotonic transducer coupled
to the tongue via a lever arm that engaged a
thermoplastic housing affixed to the tongue
surface. B: impeded protrusion trials were
conducted with the aid of a rigid force trans-
ducer that was brought into place to engage
the lever arm, thereby preventing protrusion.

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278 L. J. PITTMAN AND E. F. BAILEY

(Grass) and displayed using the Spike2 data acquisition and analysis position). When a significant F value was obtained, differences were
program (CED). tested using post hoc contrasts with significance levels adjusted
Tongue tip position (mm) was referenced to neutral or rest position according to the Bonferroni procedure. A one-way ANOVA was used
(i.e., relaxed jaw, mouth slightly open, and tongue tip visible just to test the effect of task (i.e., force vs. position) on the average initial
behind the lower incisors). Positive position values indicate protrusion and peak firing frequencies (Hz) of GG single motor units for each
from the rest position, whereas negative values indicate retraction trial. When a main effect was found, post hoc comparisons were
from this position. Subjects started from their rest position and first performed with significance levels adjusted according to the Bonfer-
protruded and then retracted the tongue to the rest position. Subjects roni procedure. A significance level of P ⬍ 0.05 was used for all
were provided with visual feedback via an ongoing display of tongue comparisons.
position on a slowed timescale but were not instructed to attain a
particular point/location in space. RESULTS
IMPEDED PROTRUSIONS. In this task, an isometric force transducer
(World Precision Instruments, Sarasota, FL) was attached to the lever Within-subject performance on successive unimpeded and
arm described earlier (Fig. 1B). Because forward movement of the impeded protrusion tasks revealed excellent reproducibility
tongue was impeded in this configuration (see Fig. 1B) the task (Fig. 2, A and B). Although the absolute tongue excursion
approximated an isometric contraction. Subjects were required to varied between subjects, the within-subject SE in performance
attain force targets of 10 and 30 g using visual feedback of the target on successive trials was minimal for both unimpeded (⫾2.7
force and their task was to match the target force. mm) and impeded protrusion (⫾3.2 g) trials.
Subjects completed three trials comprising each of the tasks in Representative raw recordings of GG and intrinsic muscle
sequence as follows: unimpeded protrusions; impeded protrusions; EMG activities, tongue position (mm), and force (gm) in

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unimpeded protrusions (see Fig. 3). unimpeded protrusion (left- and rightmost panels) and impeded
SINGLE MOTOR-UNIT ACTIVITIES. In addition to recording whole protrusion (middle panel) tasks are presented in Fig. 3. Al-
muscle EMG activities we also characterized the activities of 33 GG though both the GG and intrinsic muscles were consistently
single motor units in a subset of individuals (n ⫽ 7). Each trial activated in both tasks, the level of activity differed. Thus the
comprised three unimpeded protrusions and three impeded protru- magnitude of whole muscle GG EMG activities (% max) in
sions. Subjects were provided with auditory and visual feedback (i.e., impeded protrusion tasks was two- or threefold lower relative
via an ongoing display of the spike record on a slowed timescale) of to activities in the unimpeded protrusion task (Fig. 4; P ⬍
motor-unit discharge. For the unimpeded protrusion task, subjects
0.05). In contrast, intrinsic muscle EMG activities tended to be
were instructed to maintain the tongue in the least-protruded position
that was also associated with stable motor-unit discharge (i.e., firing higher in impeded protrusion than in the unimpeded protrusion
without interruption) for 10 s. Subjects were then instructed to retract
the tongue and then slowly advance the tongue to recruit the target
motor unit. The force transducer was subsequently coupled to the
lever arm and subjects were instructed to increase force exerted on the
transducer to recruit the target motor unit, to match the force produced
to the target force (i.e., 10 or 30 g), and then to slowly reduce force
output back to the baseline level.

Data analysis
All data were acquired and analyzed using Spike2 and custom-
designed software (CED). Quantitative comparisons of EMG activi-
ties were made as follows. Average whole muscle GG and intrinsic
muscle activities were expressed as a percentage of the maximal
response (% max), as defined earlier. Single motor-unit action poten-
tials were discriminated using a template-matching algorithm based
on waveform shape and amplitude as discussed previously (Bailey
et al. 2007b). For analysis of average discharge rate and variability,
only those motor units whose activities could be followed throughout
each series of unimpeded protrusion and impeded protrusion tasks
were included. The initial firing rate at recruitment was calculated
from the first 10 interspike intervals (ISIs) in both the unimpeded and
impeded protrusions. Peak firing rate was determined as the highest
rate for 10 consecutive ISIs during the task. For each recorded motor
unit mean values for initial and peak firing rates were calculated from
the average of three trials recorded in each condition. The estimate of
the population variance in discharge frequency (needed to calculate
sample size) is based on the only available estimate of the SD in
discharge frequency measured for GG motor units in human subjects
during volitional tasks (3.3 Hz) (Bailey et al. 2007). For this SD and
to detect differences of 4 Hz, with a power of 0.8 at a significance of
0.01 requires about 25 samples.
Task-related differences in whole muscle average EMG activities FIG. 2. A: distance (mm) relative to rest position attained by each subject
were assessed initially with a single within-subjects repeated-mea- (1–10) on successive unimpeded protrusion trials. B: forces attained by each
sures ANOVA testing main effects (and interactions) for muscle (% subject (1–10) on successive impeded protrusion trials for 10-g (black trian-
max GG vs. % max intrinsic) as a function of task (i.e., force vs. gles) and 30-g (gray triangles) force target trials.

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EMG ACTIVITY IN IMPEDED AND UNIMPEDED PROTRUSION TASKS 279

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FIG. 3. Genioglossus (GG) and intrinsic muscle electromyographic (EMG) activities during successive unimpeded (left) and impeded protrusion (right) trials.
Top: force recording (g). As depicted in Fig. 1A, force was not measured in unimpeded protrusion trials and is set to zero (left). Middle: position of tongue relative
to the subject’s neutral or rest position (mm). Lower: raw intrinsic muscle EMG activities. Bottom: raw GG EMG activities.

task. Relative to GG EMG, the intrinsic muscle EMG activities decreased protrusive force but rather remained stable through-
were significantly smaller in magnitude in the unimpeded out (Fig. 5, right). Thus for all impeded protrusion trials the
protrusion task and significantly larger in the impeded protru- average firing rate was 16.4 ⫾ 8.2 Hz, slightly higher than that
sion tasks (i.e., 30 g, Fig. 4). measured at the onset of the unimpeded protrusions (10.8 ⫾ 2).
Representative raw recordings showing the instantaneous The absence of any correspondence between firing rate and
firing rate of a discriminated single GG motor unit, tongue force during impeded protrusion tasks supports the hypothesis
position, and tongue protrusion force are presented in Fig. 5. that GG motor-unit activities are modulated in accordance with
Consistent with previously published findings (Bailey et al. tongue position, but not isometric force production. These
2007b), we observed systematic changes in GG muscle motor- findings have important implications for our understanding of
unit firing rate as a function of unimpeded tongue protrusion how the nervous system controls tongue muscles during simple
tasks (see Fig. 5, left). In this example, the motor unit was protrusion tasks, as explained in the following text.
recruited about 5 mm from the rest position and discharged at
an initial rate of 10.8 ⫾ 2 (SD) Hz, increasing to a peak rate of DISCUSSION
22.5 ⫾ 2 Hz. For all unimpeded protrusion trials, the firing rate
at recruitment was 11.5 ⫾ 4 Hz and increased to 25.3 ⫾ 4 Hz Summary
at peak displacement. By contrast, firing rate did not exhibit Hydrostats have been extensively studied at a kinematic
any systematic change either as a function of increased or level yet we know remarkably little about patterns of muscle

FIG. 4. Average whole muscle GG and


intrinsic EMG activities for the group (n ⫽
10) in unimpeded protrusions and 2 levels of
the isometric force (10 and 30 g) tasks.
Values are expressed as a percentage of the
appropriate maximum mean EMG activity
(% max). * Indicates a significant difference
between GG and intrinsic muscle EMG for a
given task (P ⬎ 0.05). ⫹ Indicates a signif-
icant task-related difference in GG EMG
activities.

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280 L. J. PITTMAN AND E. F. BAILEY

FIG. 5. Representative raw record showing


GG single motor-unit activities during unim-
peded and impeded protrusion trials (see text
for details). Top: force trace (as depicted in
Fig. 1A); force was not measured in unim-
peded protrusion trials and thus is set to zero.
Middle: position of tongue relative to the neu-
tral or rest position (mm). Lower: instanta-
neous firing rate (dots). Bottom: Discriminated
GG motor-unit potentials; inset shows overlay
of 35 consecutive potentials of the identified
unit from each trial.

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and motoneuron activities and their relationship to hydrostatic simple protrusion tasks, using both impeded and unimpeded
movements. In this study, we characterized the activities of planar movements, in an effort to characterize muscle activities
extrinsic and intrinsic protrudor muscles in human subjects under each condition separately. Whereas these findings pro-
performing unimpeded and impeded protrusion tasks. We vide new insight into the nervous system modulation of tongue
show that whereas there is activation of GG and intrinsic movement, caution should be exercised in extrapolating the
tongue muscles in both tasks, the magnitude and shape of the results to more complex natural behaviors.
drive to the intrinsic muscles and to the GG are very different, The target forces adopted under the current protocol were an
at least within the framework of the simple tasks attempted order of magnitude smaller than those reported previously in
here. “submaximal” protrusions exerted against the upper incisors or
palate (i.e., 5–24 N) (Blumen et al. 2002; BuSha et al. 2002;
Critique of method Mortimore et al. 2000; Scardella et al. 1993; Solomon and
Munson 2004; Weijnen et al. 2000). The present targets were
We selected a site in the anterior tongue body for purposes established based on the force exerted by each subject on the
of recording intrinsic protrudor muscle activities (Napadow teeth and palate (alveolar ridge) in verbal counting and swal-
et al. 1999). However, because intrinsic muscle fibers interdig- lowing tasks (10 –50 g). Based on this assessment, it is our
itate within the tongue body and are not spatially distinct, it view that the force targets were within the physiological range
was not possible for us to definitively identify the specific and that the EMG activities reported here in impeded protru-
intrinsic muscle(s) that we recorded from at that location. sion tasks approximate those developed by the tongue in
Although it is most likely that we recorded the activity of speech and some gustatory functions.
verticalis and transversus, which are both tongue protrudor
muscles, we do not know the relative contributions of each
muscle to the recording. In addition, we cannot exclude the Drive to tongue protrudor muscles in impeded
possibility that at least some of the electrical activity that we and unimpeded protrusion tasks
recorded originated in tongue muscles other than verticalis and
transversus (Slaughter et al. 2005; Stal et al. 2003). Three main findings arise from the current data set. First,
The complex interdigitation of muscles confers on the mam- within the framework of these very simple tasks, both impeded
malian tongue a remarkable flexibility of movement that un- and unimpeded protrusions beyond the teeth involve activation
derpins highly coordinated behaviors such as speaking and of the mutually orthogonal intrinsic fibers in the anterior
swallowing. Based on kinematic (Gilbert et al. 2007; Hiiemae tongue and the extrinsic GG. Although respiratory-related
et al. 2002; Stone et al. 2004) and electromyographic studies activation of tongue muscles has been documented previously
(Kayalioglu et al. 2007; Liu et al. 2007) the tongue is consid- in rat (Bailey and Fregosi 2004; Bailey et al. 2005) and human
ered to comprise several regions or “segments” capable of (Mateika et al. 1999; Oliven et al. 2007), to our knowledge this
semi-independent movement. Evidence for distinct muscle is the first time activation of both extrinsic and intrinsic tongue
compartments (Mu and Sanders 1998, 2000) and regional protrudor muscles has been demonstrated in human subjects
differences in muscle fiber type (Sokoloff 2000; Stal et al. performing volitional unimpeded and impeded protrusion
2003) add some support to independent regional control, al- tasks. Thus the present findings provide valuable new support
though this has not been demonstrated in human subjects. for a fundamental tenet of the muscular hydrostat theory
Under the current protocol, subjects were required to perform originally proposed by Smith and Kier—that there is a constant
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EMG ACTIVITY IN IMPEDED AND UNIMPEDED PROTRUSION TASKS 281

interaction of intrinsic and extrinsic tongue muscles in all versus) generate most of the tongue protrusion forces, but only
tongue movements (Smith and Kier 1989). when stabilized by contraction of the GG muscle, as discussed
Second, we have shown previously that GG MU activities in the following text.
are tightly modulated in accordance with tongue position
(Bailey et al. 2007b) but not force. Importantly, because the Working model for human tongue protrusion and force
sustained firing rates evident in the force task approximate
those previously recorded in static holding maneuvers (Bailey On the basis of current observations we have devised a
et al. 2007b) we suggest that the GG muscle fulfills a support- working model to clarify the roles fulfilled by the GG and the
ive but not a primary role in the generation of protrusive force intrinsic muscles in each task (see Fig. 6). Consistent with the
(see following text). In contrast, the GG plays a prominent role traditional view of tongue protrusion, GG contraction advances
in unimpeded displacement tasks. Intrinsic protrudor muscles, the tongue base, whereas activation of the intrinsic protrudor
on the other hand, appear to play a major role in both tasks. The muscles in the tongue tip reduces the diameter of the anterior
data suggest that the multivectorial intrinsic muscles that com- body of the tongue. Because the tongue is a constant volume
prise the core of the anterior tongue (i.e., verticalis and trans- structure, the reduction in tongue diameter secondary to intrin-

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FIG. 6. A working model of tongue muscle activation in unimpeded and impeded protrusions. Left panels: schematic lateral view of the tongue showing the
extrinsic tongue muscles (genioglossus [GG], hyoglossus [HG], and styloglossus [SG]) that have external bony origins on the mandible (m), hyoid bone, and
styloid process, respectively. The intrinsic muscles transversus (t) and verticalis (v) interdigitate extensively and have both origins and insertions in the central
tongue body. Right panels: force and position traces, with corresponding integrated GG and intrinsic muscle activities. A, left: tongue configuration at rest (dashed
outline) and in unimpeded protrusions. In this task there is activation of both the GG and intrinsic muscles (see EMG activities at right), with corresponding
changes in position but not force. GG activation causes anterior movement of the tongue base (denoted by arrow) and activation of the intrinsic muscles verticalis
and transversus compresses the tongue body, contributing to protrusion beyond the mandible, as described in the text. B, left: schematic of the tongue
configuration at rest (dashed outline) and during an impeded protrusion against an external structure (black bar, representing the force transducer). As in A,
impeded protrusions also involve activation of the GG and intrinsic muscles. Contraction of the intrinsic muscle fibers compresses the tongue body, resulting
in expansion in the long axis, which registers as protrusive force. However, force will not be registered unless GG activation fulfills a platform function that
stabilizes the tongue base (denoted by double-headed arrow), thereby resisting rearward movement that would otherwise occur in this soft-bodied structure. In
this task the magnitude of the intrinsic muscle activities exceeds that of the GG (see right panel). The magnitude and direction of the forces contributed by the
2 extrinsic retractor muscles, hyoglossus (HG) and styloglossus (SG) (arrows), are also incorporated into this working model. Although the activities of these
muscles and resultant forces were not recorded in our experiments, based on respiratory-related tongue muscle activities in human subjects (Mateika et al. 1999;
Oliven et al. 2007), we postulate that the retractor muscles also are coactive with the GG and intrinsic muscles in these tasks. Activation of the HG may be of
particular importance in protrusive force tasks as a secondary stabilizer of the tongue base (Oliven et al. 2007).

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282 L. J. PITTMAN AND E. F. BAILEY

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This work was supported by National Institute on Deafness and Other
longitudinalis muscle in the human tongue. 1. Motor endplate morphology
Communication Disorders (NIDCD) Grant K23-DC-007597 to E. F. Bailey.
and muscle fiber architecture. Cells Tissues Organs 181: 51– 64, 2005.
The content is solely the responsibility of the authors and does not necessarily
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represent the official views of the NIDCD or the National Institutes of Health.
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