Journal of Oral Rehabilitation
Journal of Oral Rehabilitation 2010 37; 663–669
Neuromuscular function in healthy occlusion
S . E . F O R R E S T E R * , S . J . A L L E N †, R . G . P R E S S W O O D ‡, A . C . T O Y § & M . T . G .
P A I N † *Wolfson School of Mechanical and Manufacturing Engineering, Loughborough University, Loughborough, †School of Sport,
Exercise and Health Sciences, Loughborough University, Loughborough, ‡Gaylord, Houston, Texas, USA and §Gorse Covert Dental Practice,
Loughborough, UK
SUMMARY This study aimed to measure neuromus-
cular function for the masticatory muscles under a
range of occlusal conditions in healthy, dentate
adults. Forty-one subjects conducted maximum
voluntary clenches under nine different occlusal
loading conditions encompassing bilateral posterior
teeth contacts with the mandible in different posi-
tions, anterior teeth contacts and unilateral poster-
ior teeth contacts. Surface electromyography was
recorded bilaterally from the anterior temporalis,
superficial masseter, sternocleidomastoid, anterior
digastric and trapezius muscles. Clench condition
had a significant effect on muscle function
(P = 0Æ0000) with the maximum function obtained
for occlusions with bilateral posterior contacts and
the mandible in a stable centric position. The
remaining contact points and moving the mandible
to a protruded position, whilst keeping posterior
contacts, resulted in significantly lower muscle
activities. Clench condition also had a significant
Introduction
The human stomatognathic system is required to fulfil
numerous disparate functions, with the primary ones
being speech and mastication. It has been shown using
invasive experimental techniques in animals that the
basic features of mastication are controlled by central
pattern generators in the brain stem (1), which are
modified using feedback from intraoral, joint and
muscle receptors to cope with the form and structure
of the food (1, 2). It is not possible to use such invasive
techniques in humans; however, studies (3) utilizing
surface electromyography (EMG) have indicated that
ª 2010 Blackwell Publishing Ltd
effect on the per cent overlap, anterior–posterior
and torque coefficients (P = 0Æ0000–0Æ0024), which
describe the degree of symmetry in these muscle
activities. Bilateral posterior contact conditions had
significantly greater symmetry in muscle activities
than anterior contact conditions. Activity in the
sternocleidomastoid, anterior digastric and trape-
zius was consistently low for all clench conditions,
i.e. <20% of the maximum voluntary contraction
level. In conclusion, during maximum voluntary
clenches in a healthy population, maximum masti-
catory muscle activity requires bilateral posterior
contacts and the mandible to be in a stable centric
position, whilst with anterior teeth contacts, both
the muscle activity and the degree of symmetry in
muscle activity are significantly reduced.
KEYWORDS: electromyography, masseter, temporalis,
maximum voluntary clench, jaw
Accepted for publication 28 March 2010
mechanoreception, in addition to neuromuscular pro-
prioception, influences the control of human jaw
function (4–7). Mechanoreception from the periodontal
ligament and stretch of the jaw closing muscles can
both elicit substantial short-term inhibitory reflexes and
long-term excitatory reflexes (3–5).
A knowledge of muscle function in simple jaw
movements such as clenching is an important step in
understanding more complex activities such as chewing
and swallowing (8). A number of studies have been
performed in an attempt to explain the differences in
muscle recruitment between clenches on different bite
points using EMG (8–13) and mathematical models
doi: 10.1111/j.1365-2842.2010.02097.x
664 S . E . F O R R E S T E R et al.
(14–16). In studies employing mathematical models of
the jaw (14–16), it was assumed that the activity levels
of muscles during clenches at different bite points were
determined purely by the mechanics of the jaw-muscle
system. The use of this approach leads to the prediction
that activation levels in all jaw adductor muscles will
reach maximum for a maximal clench at any bite point
(14, 15). However, in EMG studies that required
subjects to clench maximally at different bite points,
this result has not been reported. This indicates that
feedback from involved structures leads to a modera-
tion of the action of these muscles.
Surface EMG has been used to measure neuromus-
cular function during occlusion, most commonly for
maximum voluntary clenches as a basis for investigat-
ing temporomandibular disorders. For maximum vol-
untary clenches onto natural dentition, significant
differences have been reported in anterior temporalis
and superficial masseter activity for healthy versus
temporomandibular disorder versus neck pain subject
groups (17). However, no significant differences have
been found for healthy subjects with different angle
class occlusions (18). It has been shown that healthy
individuals have a prevalent side, on which they display
higher relative levels of muscle activity during bilateral
clenches (19). However, whilst it has been shown that
asymmetry in muscle activity exists (19), this was only
for clenches on posterior contacts, and little is known
about how this asymmetry is affected by changes in bite
point. Such information, which describes the co-ordi-
nation of muscle activity rather than simply individual
muscle activities, has the potential to add useful
knowledge to our understanding of occlusal muscle
function in healthy adults and in patients with tempo-
romandibular disorders. This study aimed to measure
neuromuscular function, including co-ordination, in
healthy dentate adults over a wider range of normal
and perturbed static occlusal conditions than has
previously been investigated. These encompassed max-
imum voluntary clenches with: bilateral posterior teeth
contacts with the mandible in different positions;
anterior teeth contacts; and unilateral posterior teeth
contacts. The hypotheses were: clenches on posterior
teeth with varied mandible positions will result in
altered muscle activity; the degree of symmetry in
muscle activity will differ for the different clench
conditions; and that the co-ordination pattern in
muscle onset times will differ for the different clench
conditions.
Materials and methods
Written informed consent was given by 41 volunteers
(32 men, 9 women; age 24Æ7  6Æ3 years) with healthy
jaw function. Loughborough University ethical advi-
sory committee approved the study which was con-
ducted in accordance with the Declaration of Helsinki.
Healthy jaw function was defined as having no more
than four teeth removed, no audible clicking during
normal jaw function and no history of jaw pain
requiring medical attention. Subjects were seated in
the alert feeding position and completed a protocol
which included maximal clenches under nine different
occlusal conditions (Table 1). In each case, the subject
was instructed to start relaxed, clench maximally for 3 s
and relax again. The order of the clenches was
randomized although always started and finished with
a natural dentition maximum voluntary clench to test
for measurement reliability. Two trials were completed
for each condition, and the most maximal was selected
for analysis. The full protocol included a number of sub-
maximal familiarization trials and trials to elicit a
maximum voluntary activation for each of the recorded
muscles (Table 1).
Electromyography signals were recorded at 2000 Hz
bilaterally from the anterior temporalis (TA), superficial
masseter (MS), sternocleidomastoid, anterior digastric
and trapezius using an active bipolar surface electrode
system*. The skin was cleaned using isopropyl wipes,
and rectangular pre-gelled disposable electrodes
(28 mm · 18 mm; Ag–AgCl†) were applied to the
muscle belly, determined through palpation, parallel
to the fibre direction with an interelectrode distance of
18 mm and secured using surgical tape. The reference
electrode was placed on the forehead. The EMG signals
were amplified at source using small custom-built
amplifiers taped firmly to the skin (gain setting, 3000;
input impedance, 1012 W; bandwidth, 10–1000 Hz; and
CMRR better than 100 dB).
The signals were bandpass filtered at 10–600 Hz using
a fourth-order zero-lag Butterworth filter. Electromy-
ography amplitude was evaluated as a 50-ms root mean
square average and normalized to the global maximum
(root mean square average) value obtained for that
muscle in all trials on a per subject basis. Muscle
activities were then expressed as a fraction of maximal,
*Biovision, Wehrheim, Germany.
†
Bio-logic, Mundelein, IL, USA.
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 NEUROMUSCULAR FUNCTION IN HEALTHY OCCLUSION 665

Table 1. Summary of the maximum voluntary clench conditions

Teeth
contacts Code Summary Description

Bilateral ND Natural dentition Intercuspal position (19)

posterior NDr Repeat of Natural dentition As above
CR Cotton rolls between molars Two 10-mm-diameter cotton rolls were positioned on the mandibular second
pre-molar and molars (19)
CRp Cotton rolls between molars Two 10-mm-diameter cotton rolls were positioned on the mandibular second
with the mandible protruded pre-molar and molars with the mandible protruded
Anterior LJ Lucia jig on maxillary incisors A Lucia jig (anterior bite stop) was attached to the maxillary incisors using impression
compound. Subjects closed onto the bite stop with the mandible positioned such
that the mandibular incisors contacted the bite stop surface at an angle of 90 and
directly beneath the maxillary incisors (21)
TF Tongue blade angled upwards A single wood tongue blade, thickness 1Æ6 mm, was positioned at an angle of 45 below
at 45 on maxillary incisors the horizontal on the maxillary incisors, and the subjects closed onto the tongue
blade such that the mandibular incisors contacted the tongue blade directly beneath
the maxillary incisors
Unilateral TR Single tongue blade between A single wood tongue blade, thickness 1Æ6 mm, was positioned on the right
posterior right molars mandibular second pre-molar and molars
TL Single tongue blade between As above on the left
left molars
T3R Three tongue blades between Three wood tongue blades in a layered configuration, combined thickness 4Æ8 mm,
right molars were positioned on the right mandibular second pre-molar and molars
T3L Three tongue blades between As above on the left
left molars

i.e. an activity of 0Æ9 indicates that the muscle was at
90% of the global maximum value. For each maximum
voluntary clench, the following four sets of parameters
were evaluated from the normalized EMG signals:
1 maximum and mean amplitude for each muscle over
the duration of the clench
2 per cent overlap coefficients (POC) for TA and MS,
anterior–posterior coefficient (APC) and torque coef-
ficient (TC)
3 muscle onset times
4 mean EMG amplitude and the coefficients given in 2
for consecutive time windows immediately following
onset
The coefficients given by 2 are those described by
Ferrario et al. (17–19) and were used to give a measure
of muscle co-ordination. The coefficients are given as a
percentage, where 100% represents the right and left
muscles contracting with perfect symmetry and 0%
perfect asymmetry for POC. Anterior–posterior coeffi-
cient and TC give similar measures for total TA and MS,
and opposing TA and MS, respectively. The onset times
used in 3 were obtained using the algorithm of Hodges
and Bui (20), which is based on an amplitude threshold
of the mean plus three standard deviations of the quiet
time amplitude to be exceeded for 25 ms based on a 50-
Hz linear envelope of the signal. The values of the
parameters described in 1–4 were compared between
the nine occlusal conditions using a repeated measures
ANOVA with Tukey’s HSD post hoc test and a significance
level of P £ 0Æ05.
Results
Muscle activity for different bite points and mandible positions
There were no significant differences in any of the
muscle activity parameters between the initial and
repeated clenches onto natural dentition, indicating
good measurement reliability (Fig. 1).
For clenches onto natural dentition, TA dominated
over MS (0Æ83  0Æ12 versus 0Æ72  0Æ16, P = 0Æ001).
MS activity increased for clenches onto cotton rolls
(0Æ72  0Æ16 to 0Æ87  0Æ12; P < 0Æ001), leading to
balanced TA and MS activities.
There was a significant reduction (P < 0Æ001) in both
TA (55–60%) and MS (50%) activity for clenches onto
anterior contacts such that the MS activity was
relatively higher than TA activity (P £ 0Æ010). For
clenches onto unilateral posterior tooth contacts, there
was a significant reduction in contralateral TA activity

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666 S . E . F O R R E S T E R et al.
(a) (b)
(c) (d)
(15%, P < 0Æ001), whilst ipsilateral (blade-side) TA
activity and MS activity on both sides were similar to
the natural dentition condition. Consequently, ipsilat-
eral TA activity significantly dominated MS activity
(P = 0Æ001–0Æ002), whilst contralateral TA and MS
activities were approximately balanced. The trends for
the single and triple tongue blade were similar; how-
ever, for the triple layer, activities in both TA and MS
were consistently lower (by 5–10%).
For clenches onto cotton rolls with the mandible
protruded, there was a significant reduction in both TA
and MS activities, by approximately 35% for TA
(P < 0Æ001) and 30% for MS (P < 0Æ001) compared to
clenches onto cotton rolls with the mandible in a stable
centric position. This reduction was greater for the TA,
such that MS activity became relatively higher than TA
activity (P = 0Æ001).
Muscle co-ordination: per cent overlap, anterior–posterior
and torque coefficients
The POC (Fig. 2) varied with clench condition for both
TA and MS. Pooling clench conditions based on contact
(a) (b)
Fig. 1. Maximum activity for (a)
temporalis and (b) masseter, and
mean activity for (c) temporalis and
(d) masseter in each of the maximum
clench conditions (mean  standard
deviation). For a description of each
maximum clench condition, see
Table 1.
points (bilateral posterior versus anterior versus unilat-
eral posterior) indicated that POC was significantly
higher for bilateral posterior contacts compared to
anterior contacts for both TA (88Æ7  7Æ4% versus
83Æ9  8Æ3%) and MS (83Æ9  9Æ8% versus 75Æ4 
11Æ7%; Table 2). The APC (Fig. 3) showed a similar
dependency on contact points with APC significantly
higher for bilateral posterior contacts compared to
anterior contacts (87Æ5  8Æ1% versus 79Æ1  10Æ2%;
Table 2). The TC (Fig. 3) showed lower variability
between clench conditions but was still significantly
higher for bilateral posterior contacts compared to
anterior contacts (91Æ7  4Æ4% versus 89Æ7  4Æ0%;
Table 2).
Per cent overlap coefficients was consistently higher
for TA compared to MS, particularly for clenches onto
natural dentition, cotton rolls with the mandible
protruded and anterior contacts. The difference was
less marked for the clenches onto cotton rolls with the
mandible centric and the unilateral tongue blades.
The cotton rolls with the mandible centric provided the
most consistent co-ordination coefficient values, i.e.
the lowest group variances (Figs 2–3).
Fig. 2. Per cent overlap coefficients
for (a) temporalis and (b) masseter in
each of the maximum clench condi-
tions (mean  standard deviation).
For a description of each maximum
clench condition, see Table 1.
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 NEUROMUSCULAR FUNCTION IN HEALTHY OCCLUSION 667

Table 2. Magnitude of the per cent overlap (POC), asymmetric (AS), anterior–posterior (APC) and torque (TC) coefficients for clenches
with [1] bilateral posterior contacts, [2] anterior contacts, [3] unilateral posterior contacts and averaged over all conditions (mean  s.d.)

[1] Posterior bilateral [2] Anterior [3] Posterior unilateral All clench
Coefficient (ND, NDr, CR) (LJ, TF) (TL, T3L, TR, T3R) conditions P Post hoc

POC, TA 88Æ7  7Æ4 83Æ9  8Æ3 84Æ2  10Æ0 86Æ0  8Æ9 0Æ0000 [1] > [2]
[1] > [3]
POC, MS 83Æ9  9Æ8 75Æ4  11Æ7 83Æ8  8Æ7 82Æ0  10Æ5 0Æ0000 [1] > [2]
[3] > [2]
APC 87Æ5  8Æ1 79Æ1  10Æ2 89Æ5  5Æ0 86Æ5  8Æ4 0Æ0000 [1] > [2]
[3] > [2]
TC 91Æ7  4Æ4 89Æ7  4Æ0 89Æ9  5Æ5 90Æ5  5Æ0 0Æ0024 [1] > [2]
[1] > [3]

Significant differences between the different contact conditions are indicated. TA, anterior temporalis.

(a) (b)

Fig. 3. a) Anterior-posterior coeffi-

cient and (b) torque coefficient in
each of the maximum clench condi-
tions (mean  standard deviation).
For a description of each maximum
clench condition, see Table 1.

cotton rolls), the lowest muscle activities occurred for

Onset of muscle activity
The onset times for muscle activity did not identify any
significant differences between muscles. The muscle
activity parameters and co-ordination coefficients eval-
uated over short time periods immediately following
onset showed the same significant trends as described
earlier for maximum muscle activity, POC, APC and TC
evaluated over the whole clench.
Co-activity of the sternocleidomastoid, anterior digastric
and trapezius
clenches with anterior teeth contacts (Lucia jig and
angled tongue blade) and asymmetric TA activity
(lower on the contralateral side) but symmetric MS
activity occurred for clenches onto unilateral posterior
teeth contacts (single and triple tongue blades). Conse-
quently, for the latter, the ipsilateral (blade) side
showed similar neuromuscular function to clenches
onto natural dentition with the TA activity relatively
higher, whilst the contralateral (non-blade) side had
approximately balanced TA and MS activation levels.

Activities of sternocleidomastoid, anterior digastric and Table 3. Maximum and mean activities for the sternocleidomas-
toid, anterior digastric and trapezius averaged over all maximum
trapezius were generally low (Table 3), and there were
clench conditions and left and right sides (mean  s.d.)
no significant differences between occlusal conditions.
Maximum Mean
Muscle activity ()) activity ())
Discussion
Sternocleidomastoid 0Æ36  0Æ23 0Æ17  0Æ11
Neuromuscular function of the masticatory muscles Anterior digastric 0Æ27  0Æ18 0Æ11  0Æ07
was measured for maximum voluntary clenches over a Trapezius 0Æ15  0Æ10 0Æ066  0Æ045
wide range of occlusal conditions in a healthy popula-
Note that the values are normalized based on the maximum
tion. As has been previously observed (8–13, 21, 22), voluntary electromyography (EMG) amplitudes obtained from
the highest muscle activities occurred for clenches with independent trials aimed specifically at eliciting a maximum effort
bilateral posterior teeth contacts (natural dentition and in each of the muscle groups.

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668 S . E . F O R R E S T E R et al.
A notable finding was that bilateral posterior contacts
only resulted in the highest muscle activities when the
mandible was in a muscularly supported and stable
centric position. For clenches onto cotton rolls with the
mandible protruded, both TA and MS activities were
significantly reduced and, in contrast to all other
bilateral posterior teeth contact conditions, MS was
more active than TA. This suggests geometrical feed-
back on the position of the mandible may also influence
masticatory muscle activity. Given that masticatory
muscles have previously been shown to have a sensi-
tive stretch reflex mechanism (5), this is unsurprising.
Values of the per cent overlap, anterior–posterior and
torque coefficients for clenches onto natural dentition
and cotton rolls were similar to those reported by
Ferrario et al. (18, 19) for healthy young adults under
similar conditions. Notably, the coefficients were sig-
nificantly higher for clenches on bilateral posterior
contacts compared to anterior contacts, indicating that
subjects were not able to recruit muscles as symmetri-
cally during clenches with anterior contacts. Similar
higher levels of asymmetry have been observed in
individuals with temporomandibular disorders (17),
and thus, it might be beneficial to consider changes in
muscular symmetry with differing bite points when
designing occlusal devices. Interestingly, variances in
the co-ordination coefficients tended to be lowest for
the clenches onto cotton rolls, i.e. onto a softer occlusal
surface than provided by natural dentition, indicating
that this clench condition produced the most consistent
muscle activity co-ordination patterns across the group.
To a lesser extent, this was also seen in the unilateral
tongue blade conditions. This may indicate that clench-
ing onto a softer substance leads to activation patterns
more representative of those employed by people in
more functional activities and support the use of
clenching onto cotton rolls as the most effective
condition to investigate differences in neuromuscular
function between different populations.
Low levels of co-activity were observed in the
sternocleidomastoid, anterior digastric and trapezius
(0Æ17, 0Æ11 and 0Æ07, respectively) that were unaffected
by occlusal condition. Although it should be noted that
as these activities were low, the signal-to-noise ratios
may not have been sufficiently high to allow statistical
differences between conditions to be identified. The
co-activity values from this study are in good agree-
ment with the literature: Ferrario et al. (18) reported
mean sternocleidomastoid activity for maximum
voluntary clenches onto natural dentition of 0Æ14–
0Æ24; Clark et al. (23) reported values of 0Æ12–0Æ14 for
the sternocleidomastoid under similar conditions; and
Ciuffolo et al. (24) measured sternocleidomastoid, ante-
rior digastric and trapezius for maximum voluntary
clenches onto natural dentition and found significant
co-activation in the sternocleidomastoid and anterior
digastric only. It is likely that jaw depressor muscles are
active during mastication and clenching to protect the
teeth in the event of fracture of brittle foods (8). Their
activation is lower in clenches than mastication (25),
probably because of the reduced likelihood of distur-
bance if biting in intercuspation or onto robust occlusal
appliances.
The failure of the results to distinguish differences in
onset time between the individual muscles may not
support no difference in onset time but rather the
limitation in the accuracy with which onset time could
be estimated from the EMG signals. Because of this
issue, muscle activity parameters and co-ordination
coefficients immediately following onset were also
evaluated and compared between occlusal conditions
in an attempt to gain a more reliable assessment of
neuromuscular function around onset. This analysis
produced results very similar to those obtained by
considering the whole clench, which may support the
reliability of the data presented. However, it should be
noted that such an analysis includes the effects of both
muscle onset time and rate of increase in muscle
activity following onset, and thus, differences in muscle
activity onset times cannot strictly be inferred.
In summary, this study has supported the existence
of a number of common neuromuscular patterns for
occlusion in a healthy population. Maximum mastica-
tory muscle function was obtained for occlusion with
bilateral posterior teeth contacts. However, maximum
masticatory muscle function also required the mandible
to be in a muscularly supported and stable centric
position, indicating that geometrical factors, as well as
periodontal ligament feedback, may influence muscle
activity. The co-ordination coefficients were signifi-
cantly higher for bilateral posterior contacts compared
to anterior contacts indicating clenches onto posterior
contacts resulted in more symmetric, or balanced,
muscle activity. Group variance in the co-ordination
coefficients were lowest for clenches onto cotton rolls,
indicating that this may be the most effective condition
to investigate differences in neuromuscular function
between different populations.
ª 2010 Blackwell Publishing Ltd
 NEUROMUSCULAR FUNCTION IN HEALTHY OCCLUSION
Acknowledgments
Work funded by grants from the British Society of
Occlusal Studies and Mrs Marion E Mundy, Houston,
TX.
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Correspondence: Stephanie Forrester, Wolfson School of Mechanical
and Manufacturing Engineering, Loughborough University, Lough-
borough LE11 3TU, UK. E-mail: s.forrester@lboro.ac.uk