Journal of Applied Biomechanics, 2008, 24, 333-339

© 2008 Human Kinetics, Inc.

Lower Extremity Muscle Functions During Full Squats

D.G.E. Robertson, Jean-Marie J. Wilson, and Taunya A. St. Pierre
University of Ottawa

The purpose of this research was to determine eccentric if it lengthens, and isometric when it is active
the functions of the gluteus maximus, biceps but there is no change in muscle length. The recruitment
femoris, semitendinosus, rectus femoris, vastus level of a muscle is usually identified by the relative
lateralis, soleus, gastrocnemius, and tibialis magnitude of its electromyogram (EMG) compared
anterior muscles about their associated joints with its maximal magnitude. More difficult to define is
during full (deep-knee) squats. Muscle func- the work done by a specific muscle and where the energy
tion was determined from joint kinematics, it produces is used within the musculoskeletal system.
inverse dynamics, electromyography, and To overcome this currently unsolvable problem, the
muscle length changes. The subjects were six work done by the moments of force across each joint are
experienced, male weight lifters. Analyses used to estimate the net work done by all the structures
revealed that the prime movers during ascent that cross the joint. Since the main contributors to the
were the monoarticular gluteus maximus and work done across a joint are muscles, especially when
vasti muscles (as exemplified by vastus latera- the joint does not reach its anatomical limits, biomecha-
lis) and to a lesser extent the soleus muscles. nists have a partial way of determining the roles of mus-
The biarticular muscles functioned mainly as cles during motion.
stabilizers of the ankle, knee, and hip joints by One can, for example, analyze a simple flexor
working eccentrically to control descent or movement and observe an increasing moment of force
transferring energy among the segments during and a simultaneous increase in the EMG of the flexor
ascent. During the ascent phase, the hip exten- muscle. To terminate the period of flexion, an antago-
sor moments of force produced the largest nistic extensor muscle may turn on to cause an extensor
powers followed by the ankle plantar flexors moment of force. The flexor muscle would be observed
and then the knee extensors. The hip and knee to be shortening during its period of contraction while
extensors provided the initial bursts of power the extensor muscle would be shown to be lengthening
during ascent with the ankle extensors and or eccentrically contracting. The situation becomes
especially a second burst from the hip exten- more complex when multiple joints are involved and
sors adding power during the latter half of the some of the muscles cross more than one joint. For
ascent. example, researchers have come to opposite conclusions
when analyzing the role of biarticular muscles during
Keywords: electromyography, inverse dynam- the vertical jump. Bobbert and Van Ingen Schenau
ics, kinesiology (1988) stated that energy was transferred distally by
biarticular muscles, whereas Pandy and Zajac (1991)
showed a distal-to-proximal transfer of energy. To reach
Traditionally, kinesiologists and physical educators
their conclusion, Pandy and Zajac determined the con-
classify muscles in several ways: by anatomical func-
tributions of muscles based on a musculoskeletal model
tion, by type of contraction, by level of recruitment, and
of the lower extremity. They pointed out that muscles
by work done by the muscle during a particular motion.
crossing a particular joint can deliver power to segments
Anatomically, muscles are defined by whether they are
remote from the joint(s) that they cross. If only it were
flexors, extensors, abductors, adductors, and so on based
possible to attach power meters to the muscles to watch
on their lines of action across the joints that they cross.
the flows of mechanical energy as we do to measure the
Muscle contractions are categorized by how a muscle’s
flow of electrical energy to a house.
length changes—concentric if the muscle shortens,
Elftman (1939a, 1939b) suggested a partial solution
to this problem and applied it to walking. His approach
The authors are with the School of Human Kinetics, Univer- was to calculate the powers due to the net forces and
sity of Ottawa, Ottawa, ON, Canada. moments at each joint as well as the instantaneous

   333
 334   Robertson, Wilson, and St. Pierre

powers of each segment. The segmental powers could 1988), vertical lifting (Molbech, 1965; Wilson & Rob-
then be accounted for by the flows of energy to or from ertson, 1988), sprinting (Simonsen et al., 1985), and
the segment at each end. Winter and Robertson (1978) jumping (Bobbert & Van Ingen Schenau, 1988; Pandy
used his methods to show that energy could be tracked & Zajac, 1991; Zajac, 1993; Prilutsky & Zatsiorsky,
during gait and proved that during the push-off phase of 1994; Jacobs et al., 1996).
walking, work generated by the ankle plantar flexor The purpose of this study was to determine the
moment was used to supply energy to the foot, leg, functions of the major lower limb muscles, particularly
thigh, and even the trunk by the transfer of energy the biarticular muscles, during full squats (descent and
though passive joint structures. However, no effort was ascent) based on EMG activity, inverse dynamics,
made to identify which muscles were responsible for moment powers and estimated muscle length changes.
the various bursts of positive and negative work during Past research (Molbech, 1965; Wilson & Robertson,
the complete gait cycle. 1988) examining biarticular muscles during full squat-
In this study, we will apply inverse dynamics and ting has provided some evidence for the existence of
moment power analysis to determine the work done at paradoxical muscle activity. A study by Andrews (1985)
the joints coupled with information about various major attempted to define paradoxical activity through the
muscles of the lower extremity to determine how the examination of a first-order differential relationship
motions of the full squat are achieved. In particular, the between muscle length and joint angle that yields the
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roles of the biarticular muscles will be elucidated based moment arm length. Unfortunately, his method did not
on their levels of contraction (EMGs) and their states of account for changes in muscle recruitment.
contraction (lengthening or shortening). For example, a
curious paradox can occur when two opposing biarticu- Methods
lar muscles contract simultaneously to produce motion
at both joints instead of stiffening the joints they cross. The subjects were six male experienced weight lifters.
Such a situation, first described by Duchenne (in 1885, The subjects varied in height from 1.78 to 1.90 m and in
see Kuo, 2001) and Lombard (1903) and subsequently mass from 71.8 to 95.5 kg. The body was modeled as
named Lombard’s paradox, is observed when rectus four rigid segments connected by frictionless pin joints
femoris and biceps femoris contract concurrently during at the hip, knee, and ankle. Segmental masses, radii of
the motion of rising from a chair. The extension seen at gyration, and centers of gravity were calculated from
both the hip and knee is the result of the differential proportions described by Dempster (1955) and Plagen-
moment arms of the two muscles at each joint. Since the hoef (1971). The length of the muscles of interest
rectus femoris has a greater moment arm across the (soleus, tibialis anterior, gastrocnemius, vastus lateralis,
knee, due to the patella, it creates an extensor moment at semitendinosus, biceps femoris, rectus femoris, and
the knee. Biceps femoris has the longer moment arm at gluteus maximus) were calculated using Frigo and
the hip so it creates an extensor moment there. Thus, Pedotti’s (1978) model with modifications by Hubley
simultaneous contractions of these muscles from the (1981) to allow scaling for different sized persons. The
seated position causes extension of the both the knee bar was treated as a particle with its center of gravity
and hip. acting at its geometric center. Angular motion of the bar
Experimentally determining how two-joint muscles about its center of gravity was considered negligible
contribute during a full squat requires information about (McLaughlin et al., 1978).
muscle lengths, joint kinematics, and net moments of Before data collection, pairs of silver–silver chlo-
force. Molbech (1965) suggested that biarticular mus- ride electrodes were placed on the muscles at locations
cles of the lower extremity act in a “paradoxical” fash- specified by Delagi et al. (1975). Skin impedance was
ion when the movement is constrained or controlled. confirmed to be below 20 kΩ and the interelectrode dis-
For his example, the movement consisted of having the tance set to 2.5 cm to reduce cross talk (Winter et al.,
feet motionless on the ground and the hips following a 1994). High-input-impedance (10 MΩ) differential
vertical track. He showed that a paradoxical situation amplifiers (>110 dB CMRR, 10–500 Hz band-pass)
occurred because when a biarticular muscle, such as the were used to obtain reliable EMG signals. The full-
gastrocnemius contracted, it caused knee extension wave-rectified EMG signals were filtered through 2nd-
when normally it was a knee flexor. In a subsequent order Butterworth filters with a 6-Hz cutoff frequency
paper (Carlsöö & Molbech, 1966), he and Carlsöö pro- (Winter, 1990), yielding their linear envelopes.
posed that a similar situation existed for seated cycling Following a warm-up and rest period, the subjects
where knee flexors, such as the hamstrings, acted as were required to perform 12 full squat (knees maximally
knee extensors. They considered cycling a controlled flexed) trials with 3 min of rest between trials. Six of the
motion since the pelvis was fixed to the seat and the feet trials were performed unloaded as a warm-up, and the
must travel a circular path. Other studies have docu- other half were performed with a load representing 80%
mented that paradoxical activity may occur during of each subject’s previously recorded maximum. The
movements when several joints have reduced degrees of squat consisted of a 2-s descent during which the ankle,
freedom, for example, during seated bicycling (Gregor hip, and knee became flexed into a full squat followed
et al., 1985; Andrews, 1987), rowing (Robertson et al., immediately by a 2-s ascent during which the three
 Muscle Activity During Full Squats   335

joints extended to return to an upright posture. The load joints extended concurrently. The minimum flexion
lifted during the full squats varied from 600 to 1226 N. angles for all three joints occurred at the end of descent.
The subjects executed full squats with their right Because there were no significant differences in the
feet on a force platform (Kistler 9261A). Cinemato- ranges of motion of the joints between the unloaded and
graphic, electromyographic, and kinetic data were col- loaded conditions, based on a dependent groups t test (p
lected simultaneously. A cine camera was positioned = .055), and since it was the loaded conditions that were
perpendicular to the plane of motion, and all data were of most interest, only results from the loaded conditions
sampled at 50 Hz. Following the 12 trials, the subjects are presented.
were required to perform three maximal isometric con- At all three joints the peak angular velocities
tractions for each muscle. To elicit the MVCs, each sub- occurred simultaneously. The peak negative angular
ject adopted a partial squat position with the heels not velocity occurred at just after 10% of the cycle whereas
touching the ground and performed a maximal isomet- the peak positive angular velocity occurred at 90% of
ric contraction against a chain that was connected with the cycle. The peak angular velocities at the hip and
the ground. In pretesting, this procedure was shown to knee joint (approximately 2 rad/s), however, were sub-
produce MVCs in all of the major muscle groups that stantially higher than that of the ankle joint (<1.0 rad/s).
were used in this study. The EMG results were averaged The hip, knee, and ankle joints had very distinct
and the maximum contraction values were used to detect patterns of the net moments of force (Figures 2, 3, and 4).
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the relative EMG activation states of the muscles during The hip and ankle joints showed extensor moments
the full squats. throughout ascent and descent, whereas the knee joint
Within each of the two testing conditions (with and started and ended with a flexor moment, with an exten-
without load), linear-envelope EMG signals for each sor moment in between. Peak moments varied between
muscle were normalized over time and to maximum subjects but, on average, the highest peak moments were
EMG values. These data were then averaged for each produced at the hips (maxima around –300 N·m) fol-
subject over six trials to yield an ensemble average lowed by the ankles (approx. –200 N·m) and then the
EMG pattern for each muscle. All within-subject ensem- knees (between ±100 N·m).
ble averages were then averaged to produce the across- The moment powers (Figures 2, 3, and 4) at the
subject grand ensemble average (Yang & Winter, 1984) knee differed notably from the patterns seen at the other
for each muscle and testing condition. two joints. The power patterns for both the hip and the
The film data were digitized to an accuracy of less ankle extensors were eccentric during the descent and
than 0.5 mm and then processed with the Biomech concentric during the ascent. In contrast, power values
Motion Analysis System (http://www.health.uottawa. at the knee started with the flexors acting concentrically
ca/biomech/csb/biomech.htm). The motion data were and then with the extensors acting eccentrically for the
smoothed with a zero-lag, 4th-order, Butterworth filter rest of the descent. The ascent began with the extensors
set to a cutoff frequency of 6 Hz. Joint angular displace- switching to concentric contractions and ended with the
ments and velocities were derived from the motion data flexors working eccentrically. Again, the peak powers
and combined with the force plate data for inverse were highest for the hip extensors followed by the ankle
dynamics analysis that resulted in the calculation of net plantar flexors and then the knee extensors.
internal moments of force and their associated powers The functions of the moments of force at the hip
(Robertson & Winter, 1980) at the ankle, knee, and hip. and ankle divide into two phases (Figures 2 and 4). In
These kinematic and kinetic data were then time nor- the descending phase, extensor moments dissipated
malized and averaged across subjects. energy in order to control the rate and amount of the
Muscle-tendon-unit lengths for each subject were descent, whereas, in the ascending portion, work was
calculated from relative angle changes, (Frigo & Pedotti, generated by extensor moments about these joints. In
1978; Hubley, 1981) normalized to their length during contrast, the functions of the knee moments of force
standing and then ensemble averaged. These averages divided into four phases (Figure 3)—two during descent
were time normalized and then averaged across sub- and two during ascent. Initially during descent, positive
jects, resulting in grand ensemble, muscle length work was performed by the knee flexors to unlock the
histories. knee and initiate downward motion (0–15% of the
cycle). During the rest of the descent, energy was dissi-
pated by the knee extensors in an effort to control knee
Results flexion (15–53% of the cycle). Positive work performed
by the knee extensors signaled the start of the ascent
A slightly greater portion of the squat was required for (53–85% of the cycle). Near the end of the ascent, the
the descent phase (0–53%) than for the ascent phase knee flexors started to perform negative work (acted
(53–100%) based on the change from flexion to exten- eccentrically) to reduce the rate of knee extension and
sion of the three joints. The changes that occurred in the presumably prevent hyperextension (85–100% of the
angles of the joints are illustrated in Figure 1. During cycle).
the descent, all three joints flexed, simultaneously; the The ensemble-averaged lengths of the muscles as
reverse was seen during the ascent, in which all three proportions of their standing lengths during the full
 336   Robertson, Wilson, and St. Pierre
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Figure 1 — Joint angles (in degrees ± 1 SD) during loaded Figure 3 — Knee angular velocity, net moment of force, and
and unloaded full squats. moment power (± 1 SD) during loaded full squats.

Figure 2 — Ankle angular velocity, net moment of force, and

moment power (± 1 SD) during loaded full squats.

squats are displayed in Figure 5. Three of the monoar-

ticular muscles acted at lengths beyond their standing
lengths. Soleus, gluteus maximus (GM), and vastus lat- Figure 4 — Hip angular velocity, net moment of force, and
moment power (± 1 SD) during loaded full squats.
eralis (VL) all lengthened beyond their standing lengths
during descent, by 7%, 29%, and 18%, respectively,
before returning to their standing lengths at the end of descent and lengthening on ascent. Gastrocnemius,
ascent. Only the monoarticular tibialis anterior muscles semitendinosus, and biceps femoris each shortened by
shortened 5% during descent before returning to their 8%, 20%, and 6%, respectively, during descent.
standing lengths during ascent. Linear-envelope electromyograms averaged across
Of the biarticular muscles, the rectus femoris mus- subjects and normalized to each subject’s maximum
cles stayed close to their standing lengths with changes voluntary contractions (MVCs) are displayed in Figure
of less than 2%. They were also unusual by having a 6. These curves show the onset and recruitment levels of
bimodal pattern—lengthening then shortening during each muscle during the squatting motion. The descent
both descent and ascent. All other biarticular muscles phase of the full squat was characterized by tibialis
exhibited unimodal contraction patterns—shortening on anterior, vastus lateralis, and rectus femoris activity,
 Muscle Activity During Full Squats   337

of the antagonistic muscles was observed at the knee

and hip joints as the rectus femoris, gluteus maximus,
vastus lateralis, semitendinosus, biceps femoris, and
gastrocnemius contracted simultaneously during the
extension (ascent) phase. EMG activity of the semiten-
dinosus paralleled that of the biceps femoris since both
are part of the hamstrings group, as did vastus lateralis
and rectus femoris, both part of the quadriceps group.
Thus, the hamstrings and gastrocnemius acted antago-
nistically to the actions of the quadriceps group at the
knee while the gluteus maximus acted antagonistically
to the rectus femoris at the hip during the early portion
of the ascent.

Discussion
The tibialis anterior contracted concentrically about the
ankle during descent to assist dorsiflexion. Soleus, as
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expected, functioned eccentrically but contributed little

based on its relatively low level of EMG activity. In con-
Figure 5 — Lengths of the muscles (± 1 SD) as proportions trast, gastrocnemius, which also had a relatively low
of their standing lengths during loaded full squats. level of recruitment, contracted concentrically. As a
two-joint muscle, the role of the gastrocnemius may
have been to assist knee flexion, whereas the soleus
acted to limit the amount of ankle dorsiflexion during
descent. This observation is supported by the fact that
soleus level of recruitment increased to almost 50%
MVC as the subjects reached maximum descent.
Muscle activity about the knee during descent was
characterized by two periods—one of concentric work
and one of eccentric work. During the initial brief period
of concentric work gastrocnemius, semitendinosus, and
biceps femoris acted together to initiate flexion of the
knee. These muscles concentrically contracted and were
active at around 25% of MVC. In particular, gastrocne-
mius exhibited a brief burst that combined with the two
hamstring muscles was enough to unlock the knee and
permit descent. Afterward, knee flexion continued with
the knee extensors acting eccentrically to control and
eventually terminate the descent. During the eccentric
part of descent, continuous activity of the vastus latera-
lis contracting eccentrically was evident. Rectus femo-
ris (RF), another knee extensor, exhibited increasing
Figure 6 — Linear-envelope EMGs (± 1 SD) during loaded EMG activity as the descent reached maximum depth.
full squats as percentages of maximum voluntary contractions
During the second half of decent, however, RF was
(MVCs).
shortening slightly so it may have acted more as a hip
flexor or as a stabilizer at both hip and knee. Since RF’s
whereas the ascent phase showed increasing levels of length changed less than 2%, one could characterize its
activity from all the muscles, excepting tibialis anterior, role as being isometric and therefore was responsible
which decreased. for transmitting energy across the hip and knee joints
At the ankle joint, little evidence of coactivation of for dissipation by the knee extensors and ankle plantar
antagonistic muscles occurred except at the deepest part flexors. This type of mechanism has been shown for
of the descent and during the early part of the ascent jump landings by Prilutsky and Zatsiorsky (1994), but
phase (50–70% of cycle duration). Gastrocnemius and the exact nature of RF’s contributions are difficult to
soleus EMG patterns followed each other synergisti- gauge using only inverse dynamics (Zajac, 1993).
cally, and both these muscles were relatively inactive The hip moment of force was consistently extensor
(<25% MVC) during periods of significant tibialis ante- through the squat. During descent, the extensor moment
rior activity. In contrast, coactivation (i.e., >25% MVC) of force did negative work to control the rate and amount
 338   Robertson, Wilson, and St. Pierre

of hip flexion. Gluteus maximus contracted eccentrically not vary much (<2%) so they may have acted to prevent
throughout the descent with EMG levels starting at 10% excessive hip extension, which might result in the sub-
MVC and increasing to about 25% MVC around mid- ject falling backward.
descent. The gluteals curiously reduced their activity The antagonistic hamstrings, biceps femoris, and
level at maximum squat depth. Presumably, they were semitendinosus, were also recruited during this period
not needed to maintain stability or perhaps they permit- especially at mid-ascent, but contracted eccentrically.
ted an extra degree of hip flexion that would have created Gastrocnemius, another biarticular knee flexor, was also
a deeper counter-movement immediately before ascent. actively recruited during ascent and like the hamstrings
Counter-movements enable stronger contraction forces contracted eccentrically. These antagonists would
when they are done rapidly and immediately before a appear to be working against knee extension but their
powerful movement (Van Ingen Schenau, 1984). combined actions may in fact assist extension. If we
Not surprisingly, during the ascent phase all three consider that their lines of action are both partly directed
moments of force were extensor and performed concen- rearward, their vector sum would produce a force that
tric work. The hip extensors dominated earliest and pulls the knee joint backward and into extension. Such a
throughout the ascent, having the largest peak moments mechanism has been suggested for biarticular muscles
(>300 N·m) and the largest peak powers (>200 W at when the segments are not constrained, as in jumping
60% of cycle time with a second peak >300 N·m at 85% (Zajac, 1993), but may also be true for lifting activities,
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of cycle time). The knee extensors produced relatively in which the distal ends of the kinematic chain (i.e., the
the lowest powers of the three moments and only con- feet) are constrained by the ground. This paradoxical
tributed positive work for the first two-thirds of the ability of the biarticular muscles was suggested by Mol-
ascent. The ankle plantar flexors produced larger powers bech (1965) but was not investigated for a situation
than the knee extensors but did not contribute their max- where two sets of biarticular muscles (gastrocnemius
imum power until near the end of the lift (at 85% of and hamstrings) acted simultaneously. The difficulty
cycle time). This order of power production, from prox- with Molbech’s principle is that he assumed that the
imal (hip) to distal (ankle), is similar to that of counter- biarticular muscles shortened whereas for full squats the
movement jumping (Nagano et al., 1998). Despite this biarticular muscles lengthened. Thus, his paradoxical
ordering of the powers, examination of the EMGs show muscle activity cannot be said to have occurred during
that all flexors and extensors (excepting tibialis anterior) full squats.
were recruited almost simultaneously as compared with At the hip, the gluteus maximus was clearly a major
vertical jumping (Bobbert & Van Ingen Schenau, 1988), contributor to the work done. The gluteus maximus
for which the order of recruitment was hip, knee, and muscles contracted concentrically throughout the ascent
then ankle muscles. This may be due to the static start and were actively recruited to near maximum levels
and finish required for the full squat. One obvious dif- especially during the first two-thirds of the ascent. Its
ference with the squat was that all muscles relax at the antagonist, RF, was also heavily recruited but only
end of the movement whereas in vertical jumping many during the earliest third of the ascent when it was eccen-
muscles continue to contract until and after the end of trically contracting. As mentioned previously, RF
ground contact. offered resistance to hip extension but may also have
The soleus was the major contributor to ankle been fulfilling its role as a knee extensor by transmitting
extension since it concentrically contracted and was energy from the torso to the leg (tibia). Rectus femoris
recruited almost maximally. Gastrocnemius was also could have transmitted energy because of its relatively
heavily recruited but did no positive work during this stiff behavior as shown but its minimal length change
period because it was contracting eccentrically and so (Prilutsky & Zatsiorsky, 1994). At the end of ascent, RF
could have been involved with transferring energy prox- shortened but its recruitment level was nearly zero and
imally due to its biarticular nature (Van Soest et al., therefore did not contribute energy to the motion.
1983, Zajac, 1993; Prilutsky & Zatsiorsky, 1994). As In conclusion, analysis of the EMG, inverse dynam-
expected, the antagonistic tibialis anterior reduced its ics, moment powers, and muscle length changes during
activity level during ascent but was partly activated, pre- squatting revealed the complex relationships among the
sumably to stabilize the ankle against unexpected contributions of the various major muscles of the lower
perturbations. extremity. Not surprisingly, there was a close agreement
During the first two-thirds of ascent, the knee exten- between the results of the six participants, which sup-
sor moment did positive work. The vastus lateralis, and ports the notion that they all performed the squat in a
presumably the other vasti, contracted concentrically similar fashion. The results support Pandy and Zajac’s
and were recruited near maximally. The RF, another (1991) notion that energy is provided in a proximal-to-
member of the quadriceps group, was also recruited distal sequence with the hip extensors providing the
based on its high levels of EMG but did not contribute greatest contributions during ascent, followed by the
positive work to the body as it acted eccentrically ankle plantar flexors, and then the knee extensors. The
through the first half of the ascent, during which the important role of the knee flexors as the initiators of the
majority of the external work by the knee extensors was squatting motion by unlocking the knee and permitting
done. In fact, as mentioned previously, RF’s lengths did the start of descent was also identified.
 Muscle Activity During Full Squats   339

Acknowledgments McLaughlin, T.M., Lardner, T.J., & Dillman, C.J. (1978).

Kinetics of the parallel squat. Research Quarterly, 49,
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Council for financial support of this project and to Graham Molbech, S. (1965). On the paradoxical effect of some two-
Caldwell, University of Massachusetts at Amherst, for a joint muscles. Acta Morphologica Neerlando-Scandi-
constructive critique. navica, 6, 171–176.
Nagano, A., Ishige, Y., & Fukashiro, S. (1998). Comparison
of new approaches to estimate mechanical output of indi-
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