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    Journal of the Neurological Sciences 174 (2000) 9299www.elsevier.com/locate/jns

    Neuromuscular disturbance outlasts other symptoms of exercise-inducedmuscle damage

    a , a b a c*Michael R. Deschenes , Rhonda E. Brewer , Jill A. Bush , Raymond W. McCoy , Jeff S. Volek ,c

    William J. Kraemera

    Department of Kinesiology, The College of William & Mary, Williamsburg, VA 23187-8795, U SAb

    Department of Kinesiology, The Pennsylvania State University, University Park, PA 16802, U SAc

    The Human Performance Laboratory, Ball State University, Muncie, IN 47306, U SA

    Received 26 July 1999; received in revised form 13 October 1999; accepted 22 December 1999

    Abstract

    This study examined the biochemical, immunological, functional, and neuromuscular responses associated with exercise-induced

    muscle damage in the quadriceps of untrained men. Muscle damage and soreness was elicited with maximal concentric/eccentric muscle21

    actions at 0.53 rads s . Significant (P,0.05) soreness was evident 1, 2, and 3 days following muscle insult, while plasma creatine

    kinase, a marker of muscle damage, was elevated 3 and 5 days post-insult. Plasma interleukin-Ib was significantly increased within 5 min,

    and remained elevated 1, 2, 5, and 7 days post-insult. Maximal isometric quadriceps function was impaired ( P,0.05) for 5 days following21

    muscle challenge. Maximal isokinetic performance at 1.09 rads s was diminished (P,0.05) for 2 days post-insult; no significant21

    decrements at 3.14 rads s were noted. Average electrical activation (iEMG) of the quadriceps was unaltered, but iEMG activity of the

    rectus femoris where soreness was focused was significantly increased. Neuromuscular efficiency (torque/iEMG) was compromisedthroughout the 10-day post-insult period investigated. While other symptoms of exercise-induced muscle damage dissipate within 7 days,

    neuromuscular perturbation persists for at least 10 days. 2000 Elsevier Science B.V. All rights reserved.

    Keywords: Creatine kinase; IL-Ib; EMG; Isometric; Isokinetic; Soreness

    1. Introduction and is generally indicated by elevated concentrations of

    circulating macrophages, monocytes, and interleukins

    Participation in unaccustomed physical exertion, par- [11,12].

    ticularly that involving eccentric, or muscle lengthening, Such muscle insult is also associated with a diminution

    actions typically results in muscle damage [14]. This of maximal force production and localized discomfortdamage is evidenced by ultrastructural damage, i.e. Z-line [13,14], yet these manifestations of exercise-induced mus-

    streaming [5,6], and due to sarcolemmal disruption, in- cle damage exhibit disparate temporal characteristics. For

    creased blood-borne levels of intramuscular proteins such example, while impaired force production is most pro-

    as creatine kinase (CK), lactate dehydrogenase (LDH), and nounced within 24 h [2,13,15], peak soreness is not

    myoglobin [710]. Consequent to this damage, an in- reported until 23 days following eccentric work [13].

    flammatory response is mounted by the immune system Accordingly, the discomfort accompanying unfamiliar

    muscular exertion is referred to as delayed onset muscle

    soreness or DOMS. The precise mechanism for the

    postponement of discomfort following muscle insult is*Corresponding author. Tel.: 11-757-221-2778; fax: 11-757-221-

    unknown, but it has been suggested that it is related to a2761.E-mail address:[email protected] (M.R. Deschenes) local inflammatory stimulation [12,16]. That is, the release

    0022-510X/00/$ see front matter 2000 Elsevier Science B.V. All rights reserved.P I I : S 0 0 2 2 - 51 0 X ( 0 0 ) 0 0 2 58 - 6

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    M.R. Deschenes et al. /Journal of the Neurological Sciences 174 (2000) 9299 93

    of pain substances such as bradykinin and histamine is tube via venipuncture of an antecubital vein. Whole blood

    delayed after the onset of muscle injury. was centrifuged at 30003g for 15 min at 48C; plasma was

    Similar to delayed soreness, the exact cause(s) of stored at2758C until analysis.

    attenuated muscle strength following unaccustomed exer- The subject then performed a light (40 W), nonspecific

    tion is not yet fully understood. It has been suggested that warm-up protocol on an electrically braked cycle ergome-

    the decrement in force production may be due to damage ter (Excalibur Unit, Lode, Groningen, The Netherlands) for

    to the contractile elements, impaired electrical activation, 5 min. One min into the warm-up, the subject was asked to

    and/ or disruption of the excitationcontraction (EC) rate the level of soreness of the non-dominant thigh. If anycoupling process [17]. soreness was indicated, the subject was asked whether the

    To date, the investigation of the effects of, and recovery discomfort was more pronounced in the inner, middle, or

    from, exercise-induced muscle damage has mainly em- outer region of the quadriceps.

    ployed an animal model [7,17,18], or in humans, a small, The subject was then situated on the isokinetic device

    seldom used muscle, i.e. biceps brachii [1,2,14,19], or the with the knee aligned with its axis, to test muscle function

    quadriceps following submaximal exertion [8,9,11,20]. The of the non-dominant leg. In preparation for concurrent

    aim of the present study was twofold: (1) to examine the isometric testing and electromyographic (EMG) analysis,

    effects of exercise-induced muscle trauma in a large, the involved knee was positioned at a 958 angle, and a one

    commonly used muscle mass (quadriceps) in humans square inch area of skin over each of the three surface

    following a maximal, voluntary effort, and (2) to assess the quadriceps muscles (vastus medialis, rectus femoris, vastus

    impact of muscle insult on neuromuscular function over an lateralis) was shaved, abraded, and cleansed with an

    extended period of time. alcohol wipe. Following the longitudinal contour of the

    muscle, 2-mm-diameter electrodes filled with electrolyte

    gel were secured on the skin with adhesive collars at an

    interelectrode distance of 2 cm, and traced with ink. These2. Materials and methods

    tracings enabled electrode placement at the same sites for

    each subsequent test. EMG signals were amplified by a2.1. Subjects

    factor of 1000 and passed through a bandwidth filter set at

    30 and 500 Hz, along with a 60-Hz notch filter. SignalsNine healthy, untrained men (20.961.0 years,

    were digitized at a sampling frequency of 1000 Hz, and174.862.0 cm, 72.366.1 kg; mean6S.E.) agreed to par-

    recorded by an on-line computer system during the first 5 sticipate in the study. After receiving a verbal description of

    of a 15-s maximal isometric contraction of the kneethe investigation, its potential risks, and the experimental

    extensors. The EMG signal was then full wave rectified,procedures employed, the subjects provided written in-

    and integrated (iEMG in mV/s). Subjects were given a 1-s

    formed consent. Each subject also completed a medical ramp-up phase to achieve full muscle activation beforequestionnaire that was examined by a physician to ensureEMG and force measurements were begun.

    that no contraindications to participation existed. AllFollowing completion of the 15-s isometric contraction,

    experimental procedures were approved by the Committeethe electrodes were removed from the subjects leg. In

    for the Protection of Human Subjects at The College ofpreparation for isokinetic testing, the subjects limb weight

    William & Mary.was measured by the dynamometer so that performance

    variables could be corrected for that resistance. A full

    2.2. Experimental protocol range of motion was then selected by the subject that was

    adhered to for isokinetic testing at each subsequent ses-

    Subjects initially performed a familiarization trial on an sion.

    isokinetic dynamometer (model 900-350, Biodex Co., Maximal isokinetic muscle function (peak torque, and

    Shirley, NY). Since an extended prophylactic effect for total work) was then determined with five repetition sets of

    muscle damage occurs following a single exertional effort alternating, concentric actions of the knee extensors andthat is specific to the musculature involved [15,19,21], flexors of the non-dominant leg at velocities of 1.05 and21

    only the subjects dominant leg was exercised during the 3.14 rads s . A rest interval of 5 min occurred between

    familiarization trial; the nondominant leg would be used sets, and verbal encouragement was provided during all

    during actual data collection sessions. To ensure that any isokinetic, as well as isometric, testing.

    potential elevations in circulating markers of muscle After collection of baseline (day 0) data was completed,

    damage or inflammation were dissipated following this subjects rested for 5 min before carrying out the damage-

    exercise, a 1014-day period interspersed the familiariza- inducing event. This muscle insult was presented as four

    tion trial from the first data collection session [22,23]. At sets of 25 repetitions, interspersed by 3-min rest periods, of

    the first test session (day 0) baseline data were collected. unilateral, alternating concentric / eccentric muscle actions21

    The subjects height and weight were determined, and a of the non-dominant knee extensors at 0.53 rads s . Five

    3-ml sample of blood was collected into a heparin-treated min following completion of the concentric / eccentric work

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    protocol, another blood sample was drawn, centrifuged, contraction divided by average iEMG during that time,

    and plasma was stored at 2758C until analysis. was determined and compared among test sessions.

    With the exception of the muscle damaging protocol and

    the blood sample taken 5 min after that muscle challenge, 2.4. Statistical analysis

    data collection, i.e. blood draw, soreness assessment,

    EMG, isometric and isokinetic testing, was repeated (61 Data are reported as means6S.E. To assess changes in

    h), at 24 h (day 1), 48 h (day 2), 72 h (day 3), 120 h (day each variable of interest during the experimental period,

    5), 168 h (day 7), and 240 h (day 10) after collection of repeated measures one-way ANOVAs were utilized. Whenbaseline data. For 10 days prior to, and throughout the significant F-ratios were found, Fisher PLSD post-hoc

    study period, subjects were instructed to avoid anti-in- analyses were conducted. In this report, only statistically

    flammatory medications, maintain normal sleeping pat- significant differences from baseline values will be iden-

    terns, and to refrain from unaccustomed muscular exertion. tified since deviations from, and resumptions to, pre-insult

    For 68 h prior to arriving at the laboratory for testing, conditions were the focus of the study. Statistical signifi-

    subjects consumed only water. cance was established at P#0.05.

    2.3. Quantitation 3. Results

    Subjective ratings of muscular soreness were determined The combination of concentric/ eccentric actions of the

    with a 010 scale where 05no soreness, 55moderate knee extensors resulted in damage, and soreness of the

    soreness, and 105extreme soreness. Plasma CK activities involved muscles. Muscle damage was indicated by in-

    were measured in duplicate with a Vitros blood chemistry creased plasma CK activity at days 3 and 5. In addition,

    analyzer (model DT 60 II, Johnson and Johnson Clinical significant elevations in plasma IL-Ib were detected at 5

    Diagnostics, Rochester, NY). Plasma concentrations of min post-insult, as well as on days 1, 2, 5, and 7.

    interleukin Ib (IL-Ib) were assessed in duplicate with Responses of these blood-borne variables are presented in

    ELISA kits (Cytimmune Sciences Inc., College Park, MD). Figs. 1 and 2, respectively.

    During the 15-s maximal isometric contractions of knee There was significant overall soreness of the quadriceps

    extensors, peak and total torque, along with fatigue (% at 1, 2, and 3 days following muscle insult, with the

    difference in torque from first to last 5 s) were calculated highest ratings occurring at day 2. When subjects were

    by the Advantage software accompanying the Biodex asked to localize where pain was most acute, 74% of the

    dynamometer. During isokinetic maximal actions of knee time the central region of the thigh (rectus femoris) was

    extensors and flexors, peak torque and total work were identified. This was significantly greater than the inner

    ascertained with the Advantage software. In addition to (vastus medialis) or outer (vastus lateralis) regions at 26%average iEMG values derived from the three surface and 12%, respectively. Soreness of the quadriceps over the

    muscles of the quadriceps, iEMG data from each of those 10-day investigation is illustrated in Fig. 3.

    muscles were examined individually over the experimental Performance of the quadriceps, during both isometric

    period. In addition, neuromuscular efficiency, defined as and isokinetic actions was significantly diminished follow-

    total torque generated during the first 5 s of the isometric ing muscle insult. During the 15-s maximal isometric

    Fig. 1. Plasma activities of creatine kinase (CK). Values are means6S.E., n59. *Indicates significant difference (P,0.05) from day 0.

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    evidenced no performance decrements (data not shown).

    Data regarding quadriceps function are found in Table 1.

    Integrated EMG data, averaged among the measured

    knee extensor muscles, exhibited no significant differences

    during the study. In addition to this overall iEMG value,

    the electrical activation of the rectus femoris, vastus

    lateralis, and vastus medialis was examined individually.

    Results of these analyses revealed no significant variationin iEMG data for the vastus lateralis or vastus medialis

    subsequent to the DOMS-inducing event. However, electri-

    cal activation of the rectus femoris was significantly

    enhanced at days 2, 3, and 10. The results of iEMG

    analyses are presented in Fig. 4.

    Neuromuscular efficiency (torque/ iEMG) was deter-

    mined during the first 5 s of the maximal isometric

    contraction of the quadriceps. This parameter demonstratedFig. 2. Plasma concentrations of interleukin Ib (IL-Ib). Values area significant diminution at each of the 10 days studiedmeans6S.E.,n59. *Indicates significant difference (P,0.05) from day 0.following the collection of baseline data at day 0. Changes

    in neuromuscular efficiency during the investigation can becontraction, significant decrements in peak and total torqueobserved in Fig. 5.

    produced were evident at days 1, 2, 3, and 5 compared tobaseline values at day 0. Torque generated during the first

    5 s of the isometric contraction demonstrated an identical4. Discussion`pattern of disturbance. Specifically, vis-a-vis baseline data,

    total force produced during the initial 5 s was less at daysThe data presented here add to a large body of evidence1, 2, 3, and 5. Interestingly, no changes in fatigue during

    confirming that unaccustomed muscle exertion results in15-s isometric contractions were demonstrated throughoutmuscle damage, and a sequela of associated physiologicalthe experimental period.responses. The plasma CK elevations consequent to mus-Isokinetic data indicate that during five repetition sets at

    21 cle insult noted here are similar to previous studies despite1.09 and 3.14 rads s , significant impairments in quad-the use of high intensity isokinetic resistance exercise toriceps performance were displayed only at the slower

    21 elicit damage and pain in the quadriceps. While othervelocity of movement. At 1.09 rads s , peak torque wasresearchers have studied circulating levels of intramuscularsignificantly lower at day 1 compared to day 0, while total

    proteins to indicate muscle damage in the knee extensors,work was significantly decreased at days 1 and 2 relative21 they typically employed submaximal efforts such as down-to baseline values. However, at 3.14 rads s , neither peak

    hill running [7,8,20], cycle ergometry [5,6], or benchtorque nor total work produced by the quadriceps variedstepping [4,9] as the unaccustomed exertion. Maximalsignificantly during the investigation. Knee flexors, whicheccentric contractions have been used to investigate circu-were not subjected to damage-inducing eccentric actions,lating markers of sarcolemmal damage, however, those

    studies focused on the elbow flexors [1,14,19]. Despite

    differences in muscles utilized, or intensity of muscle

    exertion, the temporal responses of blood-borne CK levels

    noted here are consistent with those earlier studies. Spe-

    cifically, elevations in CK activity are most marked 35

    days following unaccustomed physical activity.

    Muscle soreness manifested itself most acutely 23 daysfollowing muscle insult. The intensity of the discomfort in

    the quadriceps reported by our subjects was similar to that

    experienced in the elbow flexors following eccentric

    contractions in previous studies [19,24]. Based upon these

    and other investigations [1,4] comparing the effects of

    different modes of muscle actions on soreness, it is

    reasonable to assume that the eccentric component of our

    exercise protocol was principally responsible for the

    muscle damage and soreness evinced in our subjects.

    It has been postulated [16] that the delay that occursFig. 3. Soreness of quadriceps muscles. Values are means6S.E., n59.*Indicates significant difference (P,0.05) from day 0. between muscle insult and the onset of soreness is concor-

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    Table 1a

    Muscle function of knee extensors

    Day 0 Day 1 Day 2 Day 3 Day 5 Day 7 Day 10

    Peak torque (N)21

    0 rads s 219.0620.3* 179.7621.2 190.3621.2 188.1623.3 194.9622.8 203.6624.8 214.5625.0

    Total torque (N)21

    0 rads s 2846.96262.9* 2338.46275.7 2473.86274.0 2445.86303.4 2531.26297.6 2647.56321.2 2787.76325.2

    Torque, first 5 s (N)21

    0 rads s 942.1685.8* 775.2687.9 823.9685.5 823.6698.5 824.0686.1 869.6695.6 917.06110.0

    Peak torque (Nm)21 D

    1.09 rads s 203.3621.9 177.3614.0 191.8612.6 194.8615.0 197.6618.9 197.3618.0 200.7618.1

    Total work (J)21 [

    1.09 rads s 754.7674.5 653.1651.0 686.8639.2 723.0651.4 741.5658.3 734.0664.7 754.1666.5

    Peak torque (Nm)21

    3.14 rads s 150.8614.3 135.068.2 139.969.8 142.8612.4 148.5613.7 145.0614.1 148.0614.4

    Total work (J)21

    3.14 rads s 504.8653.6 470.7624.6 478.0625.4 494.9634.3 524.3638.6 500.9640.0 512.3647.7

    a DValues are means6S.E.;n59. *Indicates significant difference (P,0.05) from days 1, 2, 3, and 5. Indicates significant difference (P,0.05) from day

    [

    1. Indicates significant difference (P,0.05) from days 1 and 2.

    dant with the time necessary for the activation of the cle function included testing during isometric and iso-

    inflammatory response resulting from muscle damage. Our kinetic contractions. During both modes of contraction,

    plasma IL-Ib data indicate that the inflammatory response significant decreases in force production followed muscle

    of the immune system was almost immediately stimulated injury. Isometric functional capacity of the quadriceps was

    by muscle insult. After adjusting for plasma volume shifts measured during a 15-s maximal effort. A significant

    [25], a nearly threefold increase in this cytokine was decrement in peak torque, and total torque during the 15-s

    observed as early as 5 min following the series of contraction was detected 24 h following the damage-

    concentric/ eccentric muscle actions. This acute response inducing protocol, and persisted through the next 4 days.

    supports the role of IL-Ib as a pro-inflammatory cytokine By day 7, neither peak nor total torque produced were

    that is responsible for the release of other soluble agents of significantly different from baseline values.

    the immune system that more directly mediate the in- The 15-s isometric test was also used to evaluate muscle

    flammatory response [26], and suggests that it may serve fatigue. Our results indicate that while the capacity of theas a particularly sensitive marker of muscle damage. affected muscle to produce force was modulated by

    Unlike previous studies, the present evaluation of mus- damage, the ability to maintain force was not. Based upon

    Fig. 4. Average iEMG among three surface quadriceps muscles, and iEMG for rectus femoris. Values are means6S.E., n59. *Indicates significant

    difference (P,0.05) from day 0.

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    210 to day 1 occurred at 0 rads s , while the least severe

    21was at 3.14 rads s . Moreover, knee extensor function

    (peak and total torque) remained significantly compro-

    mised for 5 days post-insult during maximal contractions21 21

    at 0 rads s , for 2 days (total work) at 1.09 rads s ,21

    while it was never significantly affected at 3.14 rads s .

    In fact, it may be that these data do not conflict with

    evidence that fast-twitch fibers suffer more damage thanslow-twitch muscle. Rather, these results reasonably imply

    that a relationship exists between the degree of functional

    disturbance, and the capacity to generate force prior to

    muscle insult. In essence, the greater the ability to produce

    force, the greater the degree and duration of the loss

    suffered following injury. Alternatively, it is possible that

    since muscle damage was elicited with slow muscleFig. 5. Neuromuscular efficiency (torque/ average iEMG). Values are actions, the effects are most pronounced during move-means6S.E.,n59. *Indicates significant difference (P,0.05) from day 0. ments of similar velocities.

    In an effort to determine potential causes of the di-

    these performance data, it appears that the bioenergetics of minished muscle function observed post-insult, iEMG

    the involved muscle, at least with respect to the phos- analyses of the three surface muscles of the quadriceps, i.e.

    phagen pathway, remained unaffected. rectus femoris, vastus lateralis, vastus medialis, were

    Isokinetic muscle performance was tested at two differ- performed. Komi and Viitasalo [30] had previously ex-21

    ent velocities. As with isometric results, at 1.09 rads s amined the electrical activation of the rectus femoris and

    isokinetic performance of the quadriceps, measured as vastus lateralis following unaccustomed exertion, but those

    peak torque and total work during the five repetition set, authors collected iEMG data during submaximal contrac-

    was significantly impaired as a result of muscle damage. tions, and for only 2 days following muscle challenge. In

    However, in contrast to isometric function of the knee the present study, iEMG data were collected on each of the

    extensors which was reduced through day 5 of the study, three surface muscles during maximal isometric contrac-21

    peak torque at 1.09 rads s demonstrated a significant tions for 10 days following muscle insult. In analyzing the

    decrement only at day 1, while total work was hindered at effects of exercise-induced damage on muscle activation,

    days 1 and 2. we examined each of the three muscles individually, as

    Interestingly, when knee extensor muscle performance well as the average iEMG value among the three muscles.21

    was assessed at 3.14 rads s , no significant decreases in This indicator of overall activation of the quadricepspeak torque, or total work were noted. Although both remained steady over the period of investigation, sug-

    slow- and fast-twitch muscle fibers are activated at this gesting that decreased muscle performance cannot be

    speed, fast-twitch fibers primarily, if not exclusively, ascribed to impaired electrical excitation of the contractile

    account for force production [27]. Our in vivo functional apparatus. When iEMG data for each of the three muscles

    results appear to be at odds with ultrastructural evidence were considered individually, no impact of injury was

    that exercise-induced muscle damage occurs mainly in apparent in either the vastus lateralis or the vastus

    fast-twitch fibers [5,28]. Presumably, if damage was medialis. However, electrical activation of the rectus

    specific to fast-twitch fibers in this study, functional femoris showed increases during the study period. These

    diminution would be most pronounced at the velocity of results are interesting in that iEMG alterations were21

    3.14 rads s , where these fibers are almost exclusively localized to the same quadriceps muscle in which soreness

    responsible for force production. Yet, significant functional was most acute. This localized soreness cannot be attribu-

    disturbances are restricted to the slower movement velocity ted to normal conditions of greater recruitment, since data21of 1.09 rads s where, during maximal exertion, both collected at day 0 indicate that the rectus femoris is

    slow- and fast-twitch muscle fibers contribute to force activated no more than the vasti muscles during maximal

    production [29]. exertion prior to muscle insult (data not shown). Unlike the

    Upon closer examination, our results reveal that the vasti muscles, the rectus femoris crosses both the hip and

    amount of force generated determines both the degree, and knee joints [31]. Perhaps this anatomical arrangement

    the duration of the functional impairment of muscle leaves the rectus femoris more susceptible to stretch-

    damaged during unaccustomed exertion. Consistent with related damage from eccentric actions, and consequently to

    the forcevelocity curve, our baseline data illustrate that greater discomfort and electrical activation. The data21

    peak torque of the quadriceps was greatest at 0 rads s presented here do not allow firm conclusions to be drawn21

    (219.0 Nm) and lowest at 3.14 rads s (150.8 Nm). on these points.

    Interestingly, the most severe drop in peak torque from day During the initial 5 s of the maximal isometric muscle

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    action, measurements on muscle force production and significant alterations in electrical activation. Yet overall

    electrical activation were concurrently performed. This iEMG analyses indicate that the reduction in force pro-

    enabled us to examine neuromuscular efficiency, or the duction was unrelated to muscle activation which remained

    capacity of the contractile elements to respond to electrical unchanged for 10 days following muscle insult. These data

    stimulation of the sarcolemma. Komi and Viitasalo [30] contributed to the most striking finding of our inves-

    had found that 2 days following unaccustomed exertion, a tigation: neuromuscular efficiency was significantly im-

    disturbance in this relationship existed such that, relative to paired the first day following insult, and remained compro-

    original efforts, a greater degree of electrical activation mised for 10 days. This prolonged disturbance is probablywas necessary to perform the same submaximal muscle related to dysfunction within the EC coupling mecha-

    task. Our data extend those findings. During maximal nism; further research is needed to determine at what point

    effort, significant decrements in neuromuscular efficiency subsequent to unaccustomed exertion this disorder is

    were detected throughout the 10-day post-insult interval. resolved.

    Initially, this finding may be explained by the significant

    reduction in torque produced during the 5-s interval

    documented on days 1, 2, 3, and 5, without concomitant Acknowledgements

    changes in overall iEMG values. This implies that early

    neuromuscular dysfunction is mainly due to damage to The authors would like to express their gratitude to the

    contractile elements [5,6]. Yet, torque returned to normal subjects who generously gave their time and effort to

    values by day 7, while neuromuscular alterations persisted enable the successful completion of this study. This project

    through day 10. The continuation of this disturbance may was supported, in part, by a grant from the Faculty

    be related to the gradual trend (P50.07) toward an Research Committee of The College of William & Mary.

    increase in average iEMG activity, mainly accounted for

    by the rectus femoris, during the study period. Still, an

    impaired capacity of the myofilaments to respond to References

    increased electrical stimulation beyond day 5 points to the

    existence of some sort of muscular inefficiency. Several [1] Clarkson PM, Byrnes WC, McCormick KM, Turcotte LP, White JS.Muscle soreness and creatine kinase activity following isometric,recent studies utilizing isolated muscle preparations ineccentric, and concentric exercise. Int J Sports Med 1986;7:1525.mice have demonstrated a dissociation of EMG activity

    [2] Ebbeling CB, Clarkson PM. Exercise-induced muscle damage andfrom force production during electrically stimulated iso-

    adaptation. Sports Med 1989;7:20734.metric muscle actions subsequent to a series of eccentric [3] Newham DJ. The consequences of eccentric contractions and theircontractions [17,18,32]. In one of those investigations [17], relationship to delayed onset muscle pain. Eur J Appl Physiol

    1988;57:3539.it was found that although electrically stimulated eccentric

    [4] Newham DJ, Mills KR, Quigley BM, Edwards RHT. Pain andcontractions resulted in severely impaired peak isometric fatigue after concentric and eccentric muscle contractions. Clin Scitorque, the ability of the sarcolemma of the damaged

    1983;64:5562.muscle to conduct action potentials was only marginally [5] Friden J, Sjostrom M, Ekblom B. Myofibrillar damage followingcompromised. However, it was established that force intense eccentric exercise in man. Int J Sports Med 1983;4:1706.

    [6] Manfredi TG, Fielding RA, OReilly KP, Meredith CN, Lee HY,production of the afflicted muscle was restored to normalEvans WJ. Plasma creatine kinase activity and exercise-inducedwhen myofilaments were exposed to adequate amounts ofmuscle damage in older men. Med Sci Sports Exerc 1991;23:1028

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    [8] Byrnes WC, Clarkson PM, White JS, Hsieh SS, Frykman PN,nism. This also appears to be the most plausible explana-Maughan RJ. Delayed onset muscle soreness following repeatedtion for the prolonged (10 days) disturbance in neuro-bouts of downhill running. J Appl Physiol 1985;59:7105.

    muscular efficiency demonstrated in our investigation,[9] Newham DJ, Jones DA, Edwards RHT. Plasma creatine kinase

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