Journal of Electromyography and Kinesiology 20 (2010) 298–304 Contents lists available at ScienceDirect Journal of Electromyography and Kinesiology journal homepage: www.elsevier.com/locate/jelekin Which type of repetitive muscle contractions induces a greater acute impairment of position sense? Sylvie Fortier a, Fabien A. Basset a,*, François Billaut b, David Behm a, Normand Teasdale c a School of Human Kinetics and Recreation, Memorial University of Newfoundland, St. John’s, NL, Canada A1C 5S7 The Department of Kinesiology and Physical Education, University of Lethbridge, Lethbridge, AB, Canada T1K 3M4 c Groupe de Recherche en Analyse du Mouvement et Ergonomie, Division de Kinésiologie, Département de Médecine Sociale et Préventive, Faculté de Médecine, Université Laval, Québec (Qc), Canada G1K 7P4 b a r t i c l e i n f o Article history: Received 4 August 2008 Received in revised form 1 April 2009 Accepted 1 April 2009 Keywords: Elbow Muscle contraction Position sense Force Matching a b s t r a c t The objective of this study was to determine which type of repetitive muscle contractions induces a greater acute impairment of elbow position sense. Eleven male subjects participating in the study underwent (i) an exercise task (ET) consisting of 9 sets of 10 voluntary isometric, concentric, or eccentric contractions randomly performed on three separate sessions, and (ii) a pre- and post-exercise maximal voluntary isometric contraction (iMVC). Prior to and between sets of ET, a proprioception task (PT) consisting of matching the right arm to the left reference arm was performed at three different target angular positions (70°, 110° and 150°). Each ET was immediately followed by 3 PT and 1 min rest. The statistical analysis revealed that post-exercise iMVCs were significantly decreased compared to pre-exercise iMVC in all conditions with a greater drop following the eccentric task. Despite this greater drop, position sense was significantly affected by the concentric exercise task. In addition, the spectral EMG signals significantly shifted towards lower frequencies from the initial values, regardless of exercise task. The results showed that concentric muscle contractions impaired position sense to a greater extent compared to isometric and eccentric contractions. Crown Copyright Ó 2009 Published by Elsevier Ltd. All rights reserved. 1. Introduction A complex combination of many different signal sources, such as those from the tendon organs, muscle spindles, joint receptors, and cutaneous receptors, results in very sensitive and unambiguous sensations about our movement and limb position in space (McCloskey, 1978). In fact, in the absence of visual input, we have an accurate sense of position implying that we know the position of our limbs at any time during a movement. Attaining the aimed final position of either a limb or a particular end-point of the locomotor apparatus is an important motor task of our everyday behaviour; it can be crucial for success in various professional or sporting activities. Indeed, it has been suggested that muscle fatigue could predispose a joint to injury and thus an eventual decrease in athletic performance (Skinner et al., 1986). Proprioception has also been a topic of interest in sport rehabilitation because injuries have been found to have a detrimental effect on proprioception through the damage of mechanoreceptors in ligaments. * Corresponding author. Tel.: +1 709 737 6132; fax: +1 709 737 3979. E-mail addresses: [email protected] (S. Fortier), [email protected] (F.A. Basset), [email protected] (F. Billaut), [email protected] (D. Behm), normand.teasdale@kin. msp.ulaval.ca (N. Teasdale). Fatigue studies have demonstrated that changes induced by muscle fatigue may well be influenced by whether the contraction is static (Bigland-Ritchie et al., 1983) or dynamic (Colliander et al., 1988). Likewise, it is also well known that strenuous eccentric exercise induces greater acute damage to muscle fibres than concentric and isometric exercises (Clarkson and Newham, 1995; Faulkner et al., 1993; Komi and Rusko, 1974). The main difference between the three muscle contractions is that in an isometric contraction the muscle length remains constant while the contracting muscle is forcibly lengthened during an eccentric contraction and shortens during a concentric contraction. Therefore, comparing the three types of muscle contractions on a matching task becomes appealing to find out whether the type of contraction could have an effect on proprioception. As a matter of fact, it has been recently shown that proprioception is weakened following repetitive muscle contractions. The size of the matching errors was correlated with the drop in force (Saxton et al., 1995; Walsh et al., 2006). It was then concluded that the fall in force led to an increase in the effort required to maintain position of the limb against the force of gravity and that this increase in effort led to the matching errors (Walsh et al., 2004). Moreover, the direction of the matching errors suggested that we make use of signals of both peripheral and central origin in determining the position of our limbs in space by reliance, in part, on the amount of 1050-6411/$ - see front matter Crown Copyright Ó 2009 Published by Elsevier Ltd. All rights reserved. doi:10.1016/j.jelekin.2009.04.002 S. Fortier et al. / Journal of Electromyography and Kinesiology 20 (2010) 298–304 299 effort required to maintain limb position against gravity (Walsh et al., 2006). Muscle fatigue and proprioception has been the focus of many studies in which the majority of them have sought the effect of either isometric, concentric, or eccentric exercise on position sense (Allen and Proske, 2006; Forestier et al., 2002; Walsh et al., 2006; Winter et al., 2005). Other authors studied the effect of concentric and eccentric exercises on position sense (Allen et al., 2007; Brockett et al., 1997; Walsh et al., 2004). However, because different protocols were used for the concentric and eccentric experiments, a direct comparison between them was not possible. Therefore, the aim of this study was to directly compare the effect of the three types of muscle contraction within the same study and consequently determine which one would express the greater acute impairment of elbow position sense. Knowing that eccentric exercise exhibits a large acute drop in force in the exercised muscles, the purpose of the present study was to test the hypothesis that eccentric contractions would lead to the greater decrease of force and would therefore impair proprioception to a greater degree than concentric and isometric contractions. 2. Methods 2.1. Subjects This study was conducted on 11 right handed male subjects (mean age 28.9 ± 10.6 years; mean height 173.8 ± 6.2 cm; mean weight 80.4 ± 11 kg), without any previous history of upper limb musculoskeletal problems (see Table 1). They were all aerobically active at least three times a week and 9 subjects also included weight training in their fitness program. All subjects gave their written inform consent in compliance with Memorial University of Newfoundland Human Investigation Committee regulations, were all instructed regarding the procedure of the experiment, and filled in a Physical Activity Readiness Questionnaire (PAR-Q) (Canadian Society of Exercise Physiology, 2003). 2.2. Apparatus As displayed in Fig. 1, a supporting frame with two handles fully adjustable in the horizontal and vertical planes was instrumented with linear potentiometers (Model K/RV4, Precision Electronic Components Ltd, Weston, Ont.) and a strain gauge (Omega Engineering Inc. LCCA-250, Laval, Qc). The signals from potentiometers and strain gauge were amplified and sampled (Biopac Systems Inc., Santa Barbara, CA) at a rate of 1 kHz (12-bit A/D). In addition, the EMG activity was sampled at 1 kHz, with a Blackman 61 dB band-pass filter between 10 and 500 Hz, amplified (bi-polar differential amplifier, input impedance = 2 kX, common mode rejection Table 1 Participants’ physical characteristics. Subjects Age (years) Height (cm) Body mass (kg) 1 2 3 4 5 6 7 8 9 10 11 23 49 43 43 24 25 23 21 22 20 25 172 175 163 163 173 171 185 175 175 180 180 80 99 71 67 67 86 82 76 84 74 98 Mean (±SD) 29 (±10) 174 (±7) 80.4 (±11.0) Fig. 1. Position sense apparatus consisting of a supporting frame with two handles fully adjustable in the horizontal and vertical planes instrumented with linear potentiometers. The axis of rotation of the subject’s elbow was aligned with the axis of rotation of the handle where the potentiometer lain. Shoulders and waist were tightly strapped with safety belts. ratio P 110 dB min (50/60 Hz), gain 2000, noise P 5 lV). A voltage pulse (0–5 V) was also triggered to synchronize kinetics, kinematics and EMG data. Accuracy of angular displacements and force output were 60.05% (0–5 V) and 60.03% with a full scale ranged from 0° to 314° and from 0 to 1000 N, respectively. The axis of rotation of the subject’s elbow (lateral epicondyle) was aligned with the axis of rotation of the handle where the potentiometer was positioned. Shoulders and waist were tightly strapped with safety belts to limit trunk motion. The subjects’ arms were kept next to the body with their hands holding the handles at different angles. For the exercise tasks (ET), the resistance was attached to the right handle by a strain gauge. Surface EMG recording electrodes (MediTrace, Tyco Healthcare Group, Mansfield, MA) were placed over the belly of the biceps brachii and triceps brachii muscles in a bipolar fashion and approximately 3 cm apart. Electrodes were placed along the estimated direction of the muscle fibres. The ground electrode was placed on the clavical shaft. Skin preparation for all electrodes included removal of body hair with a razor and cleaning dead epithelial cells using an isopropyl alcohol swab. 2.3. Experimental procedure Subjects first attended a familiarization session in which anthropometric measurements were obtained to fit the subjects to the manipulandum. In addition, maximal isometric force output was determined by producing three maximal voluntary isometric contractions (iMVC) with the dominant arm flexed at 90° and a 5 min rest interval. According to Schmidtbleicher (1985), maximal isometric force output is approximately 20% higher and 20% lower 300 S. Fortier et al. / Journal of Electromyography and Kinesiology 20 (2010) 298–304 than maximal concentric and eccentric force outputs, respectively. Based on this assertion, maximal isometric force output was used to predict maximal concentric and eccentric force outputs. The exercise workload was, then, calculated by multiplying maximal force outputs for each muscle contraction by a factor of 0.75. For instance, for an individual reaching an iMVC of 200 N, the exercise workload for the exercise task would be set at 150 N, 120 N, and 180 N for isometric, concentric, and eccentric contractions, respectively. All subjects participated in three different sessions separated by a minimum interval of 7 days. For each session, testing was conducted at the same time of day to nullify possible differences attributed to diurnal rhythms. The three experimental sessions (randomly presented) were differentiated by the nature of the exercise tasks (ET) which consisted of isometric contractions, concentric contractions–concentric phase only, or eccentric contractions–eccentric phase only. All experimental sessions started by collecting the resting blood lactate level (AccutrendÒ Lactate Analyzer, Mannheim, Germany) followed by four tasks: (i) preexercise (control) proprioception task at each target angular position (TAP), (ii) exercise task (isometric, concentric or eccentric), (iii) proprioception task (PT) at each TAP, and (iv) iMVC (see Fig. 2). The PT consisted of matching angular position of the right indicator arm to that of the left reference arm. Target angular positions were set at 70°, 110° and 150° of elbow flexion. During the PT the subjects voluntarily positioned their reference arm at the TAP using visual feedback provided through the computer screen. They were asked to maintain, unsupported, this reference position, to close their eyes, and then, to match with their indicator arm. No feedback was given about their performance. When the subject indicated that both arms matched, a voltage pulse was triggered by the experimenter to identify the matching (Fig. 3). Between each TAP, subjects returned both arms to 90°. In each experimental session, subjects performed 9 sets of the exercise task. Each set consisted of 10 voluntary contractions of 4 s with a 2 s rest between muscle contractions. For the isometric condition, the elbow was flexed at 90° and the force output was provided on the computer screen. For the concentric condition, subjects were asked to extend their elbow starting from an elbow angle of 180° and to flex the arm until the arm was fully flexed, keeping their shoulder stable. At this point, they were required to relax their indicator arm while the experimenter returned the weight to its initial position. Fig. 3. Typical set of kinematic data. The solid line represents the angular displacement of the reference arm and the dash line represents angular displacement of the indicator arm. The starting position of both arms was always set at 90° of elbow flexion and when both arms were matched, a voltage pulse was triggered and corresponds to by the dotted vertical line in this figure. Similar procedures were adopted for the eccentric condition; subjects were asked to do series of lifts from full flexion to full extension. Each set of 10 contractions was separated by approximately 2 min (about 1-min to complete the 3 PT at the same TAP and 1min of complete rest). This procedure was completed three times for each TAP for a total of 9 sets per session. The TAP were semirandomized and differed from set to set. Once subjects could no longer maintain the imposed rhythm of the exercise task for two consecutive muscle contractions, the ongoing set was stopped. Finally, immediately after the last set, a post-exercise iMVC and blood lactate concentration were collected for each experimental session. 2.4. Data analysis All kinetic and kinematic parameters were analysed using MATLAB software (MathWorks Inc., Natick, MA). From these parame- 3X iMVC 3x iMVC Pre-exercise PT ET 3x PT70 ET 3x PT110 ET 3x PT150 1 min 3x PT150 Set 3 1 min A) Familiarization session 3x PT110 1 min 5 min 5 min 3x PT70 Set 2 Set 1 B) Experimental session Blood lactate sample Blood lactate sample Fig. 2. (A) Subjects first attended a familiarization session where they produced 3 iMVCs. (B) All subjects then participated to 3 different experimental sessions (randomly presented) differentiated by the nature of the exercise tasks (isometric, concentric, or eccentric contractions). Each session included three tasks: an exercise task (ET), a proprioception task (PT), and an iMVC. All sessions started by collecting the resting blood lactate level followed by 3 pre-exercise PT at each target angular position (70°, 110°, and 150°). Thereafter, subjects performed the first of nine sets of the exercise task. Each set consisted of 10 voluntary contractions and was immediately followed by 3 PT at the same angular position. Each ET was separated by approximately 2 min (about 1-min to complete the 3 PT and 1-min of complete rest). This procedure was completed nine times for a total of 3 sets of PT at each angular position. Finally, immediately after the ninth set, post-exercise iMVC and blood lactate level were collected. 301 S. Fortier et al. / Journal of Electromyography and Kinesiology 20 (2010) 298–304 ters, the following variables were determined: (a) maximal voluntary isometric contraction (iMVC): the force curve was filtered (second-order low-pass Butterworth filter with a 7 Hz cutoff frequency) and the iMVC was determined by the maximal force output of the curve; (b) position matching errors: the angular position signal was filtered with a fourth-order Butterworth filter (10 Hz lowpass cutoff frequency with dual pass to remove phase shift) and position matching error was calculated as: where 90° = horizontal forearm and 180° = vertical forearm. For each trial, matching accuracy was determined using the constant error (CE). Constant error is the mathematical operation difference between the position of the arms. By convention, a negative value refers to a more extended indicator arm whereas a positive value refers to a more flexed indicator arm; (c) total matching time: defined as the time between the first movement of the right arm from the starting position and the onset of the voltage pulse; (d) muscle contraction time: defined as the time between the start and the end of a muscle contraction performed during the exercise task; and (e) integrated EMG (iEMG): the EMG signals from the biceps brachii were full-wave rectified, integrated between onsets and offsets of bursts, and normalized by time. The onset and offset of initial and terminal bursts of first, fifth and last sets were determined from EMG signal using a threshold value of 10% of peak value. The frequency spectrum of each epoch of EMG data was analysed by a fast Fourier transformation algorithm. The frequency spectrum was restricted to frequencies in the range 5–500 Hz and was defined by 512 points in amplitude and phase. The median frequency (MF) was then computed. 2.5. Statistical analysis All variables are presented as mean (±SE) and 95% confidence intervals unless otherwise mentioned. In all instances, Greenhouse-Geisser corrections were applied if violation of sphericity was detected (Mauchly procedures). First, a one-way analysis of variance [4 experimental conditions; 1 pre-exercise (control) and 3 exercise tasks (isometric, concentric, and eccentric)] with repeated measures was performed on the iMVC parameters to look at the effect of types of contraction on post iMVC performance. Second, a two-way analysis of variance [2 data acquisition time (pre– post) 3 exercise tasks (isometric, concentric, and eccentric)] with repeated measures was computed on blood lactate. Third, a threeway analysis of variance [2 bursts (initial terminal bursts) 3 sets (first, fifth and last) 3 exercise tasks] was performed on iEMG and MF values. Fourth, a one-way analysis of variance with repeated measure (3 exercise tasks) was performed on muscle contraction time to determine any variation in EMG burst duration among exercise tasks. Finally, a two-way analysis of variance [4 experimental conditions 3 PT (70°, 110°, and 150°)] with repeated measures was computed on CE. A post-hoc HDS Tukey test was used for statistical comparisons of means. The level of significance was set at a p-value of 0.05. For all statistical tests, SPSS 14.0 for Windows was used (SPSS inc., Chicago, IL). Table 2 Mean values (±SE) in Newtons for iMVC. * Isometric Concentric Eccentric Mean Post-exercise Mean value ±SE [95% CI] Mean value ±SE [95% CI] 1.8 1.8 1.8 1.8 ±0.2 ±0.1 ±0.3 ±0.1 [1.4–2.2] [1.5–2.2] [1.2–2.5] [1.6–2.1] 3.6 3.7 3.6 3.6* ±0.3 ±0.2 ±0.5 ±0.2 [2.8–4.2] [3.2–4.3] [2.5–4.8] [3.1–4.1] Significantly different from pre-exercise at p 6 0.05. 3. Results The pre-and post-exercise mean value (±SE) for iMVCs, and blood lactate concentration are presented in Tables 2 and 3, respectively. The statistical analysis revealed a significant effect of exercise tasks on iMVCs (p < 0.05); Post hoc analysis showed a significant decrease in post-exercise iMVCs compared to pre-exercise with the greater drop in force seen with the eccentric contractions. Indeed, subjects achieved 92.6%, 84.0%, and 75.4% of their initial iMVC values during post-exercise iMVC for isometric, concentric, and eccentric muscle contractions, respectively. In parallel with force output decrement, there was a main significant effect of exercise tasks on blood lactate concentrations (p < 0.05). Yet it is worth noticing that blood lactate concentrations significantly increased regardless of exercise task. The mean values (±SE) of iEMG and MF are presented in Table 4. The three-way analysis of variance did not reveal a significant effect of the sets. The data were, then, pooled in order to further perform a two-way analysis of variance [2 data acquisition time (initial terminal bursts) 3 exercise tasks] on same variables. No significant difference in iEMG between initial and terminal values in all exercise tasks was observed (p > 0.05). On the other hand, there was a main significant effect of data acquisition time on MF (p < 0.05). Indeed, the spectral EMG signal significantly shifted towards lower frequencies from the initial values, regardless of exercise task. Fig. 4 displays CE for all experimental conditions (pre-, isometric, concentric, and eccentric). Recall that negative CE values correspond to a more extended indicator arm position. The statistical analysis revealed a main significant effect of experimental conditions on CE. In fact, CE decreased significantly after the concentric exercise compared to pre-exercise (p < 0.05). The statistical analysis also revealed that CE tends to be larger at 110° than at 70° and 150° of elbow flexion (p < 0.09) as shown in Table 5. The muscle contraction time was 3.65 s (±0.03) [95% CI = 3.59– 3.70] for isometric, 3.71 s (±0.03) [95% CI = 3.66–3.78] for concentric, and 4.00 s (±0.04) [95% CI = 3.93–4.07] for eccentric contractions. The analysis reveals a significant effect of exercise tasks on muscle contraction. Further Post hoc analysis showed that the muscle contraction time was significantly higher for the eccentric contractions compared to the isometric and concentric contractions. Despite the statistical significance, the 350 ms time difference in muscle contraction between the eccentric and isometric contractions is not believed to have resulted in a greater eccentric muscle fatigue, as confirmed by the blood lactate concentrations. Hence, this parameter will not be further discussed. 4. Discussion Mean value (N) ±SE [95% CI] 244 226* 205* 184* ±10 ±14 ±14 ±15 [221–268] [195–258] [172–238] [150–218] Significantly different from pre-exercise at p 6 0.05. Pre-exercise * angle ðreference armÞ angle ðindicator armÞ; Pre-exercise Post-exercise isometric Post-exercise concentric Post-exercise eccentric Table 3 Mean values (±SE) in mmol L1 for blood lactate concentration. The objective of this study was to determine which type of muscle contractions would impair to the greater degree position sense. The hypothesis was that eccentric exercise would lead to a larger decrease in force and consequently to a greater impairment of position sense. Our results partly confirmed our hypothesis by reveal- 302 S. Fortier et al. / Journal of Electromyography and Kinesiology 20 (2010) 298–304 Table 4 Mean values (±SE) for iEMG (mV s1) and MF (Hz). Initial burst Terminal burst Mean value ±SE [95% CI] Mean value ±SE [95% CI] iEMG Isometric Concentric Eccentric 0.193 0.243 0.178 ±0.027 ±0.056 ±0.025 [0.131–0.256] [0.113–0.373] [0.120–0.236] 0.177 0.178 0.169 ±0.019 ±0.037 ±0.017 [0.133–0.220] [0.092–0.263] [0.130–0.208] Mean 0.205 ±0.027 [0.143–0.267] 0.174 ±0.018 [0.133-0.216] MF Isometric Concentric Eccentric 129.8 128.9 125.0 ±3.5 ±2.9 ±3.8 [121.7–137.8] [122.2–135.6] [116.3–133.8] 115.0 105.9 119.4 ±4.7 ±6.0 ±3.5 [104.2–125.9] [92.1–119.7] [111.3–127.5] Mean 127.9 ±2.2 [122.8–132.9] 113.4* ±3.1 [106.4–120.5] * Significantly different from initial burst at p 6 0.05. ing that eccentric exercise induced a greater force loss compared to isometric and concentric exercises. However, data of the present study do not confirm the second half of the research hypothesis. In fact, proprioception was significantly affected with the concentric exercise indicated by matching errors resulting in a more extended indicator arm position. The results of the present study showed that post-exercise iMVCs were significantly reduced in all exercise tasks indicating that fatigue occurred with all types of muscle contraction (Table 2). There was an 8%, 16%, and 25% decline in iMVC after the isometric, concentric, and eccentric exercise task, respectively. Indeed, possible causes of the muscle performance decline may be different according to the type of contraction. It is known that when a muscle lengthens (eccentric) during activation the energy requirement and mechanical response differ from those of shortening Fig. 4. Subjects (n = 11) mean constant error (CE), that is the sign difference between the reference arm and the indicator arm, for the pre-exercise matching condition, the isometric condition, the concentric condition, and the eccentric condition (*p < 0.05; Error bars + SE). (concentric) contractions (Clarkson and Newham, 1995; Enoka, 1996). Eccentric contractions used in the present study could be associated with a lower energy cost, but with a higher tension output that may damage the muscle–tendon system (Clarkson and Newham, 1995; Friden and Lieber, 1992; Lieber et al., 1996) leading to a greater acute force loss with eccentric compared to isometric and concentric contractions. Along with the force output decrement, the blood lactate increased from pre- to post-exercise regardless of exercise task suggesting that the fatigue protocol used in this study induced the same metabolic stress in all exercise tasks. In parallel, spectral EMG signal values significantly compressed towards lower frequencies from the initial burst of the first set to the terminal burst of the last set, corroborating blood lactate outcomes. Spectral analysis also provides reliable information about metabolic activity and membrane potential changes (Billaut et al., 2006; Taylor et al., 1997). Our results agree with previous studies conducted during isometric and dynamic contractions (Billaut et al., 2006; Hagg, 1992). Hence, in the present study, the significant iMVC and CE differences observed between the types of contraction should not be related to difference in metabolic stress induced by the exercise tasks. It is generally accepted that fatigue negatively affects joint proprioception (Allen and Proske, 2006; Walsh et al., 2006; Winter et al., 2005). Indeed, immediately after the concentric exercise task, subjects of the present study made significant position matching errors (2.6 degrees) by adopting a more extended position with the indicator arm (Fig. 4). According to Brockett et al. (1997) and Gandevia et al. (2002), sense of limb position is provided by signals from skin, joint and muscle receptors. The level of resting activity from muscle spindles would, therefore, signal the length of the muscle and, accordingly, the position of elbow. The present results could suggest that the signal from muscle spindles had decreased as a result of the concentric exercise task. To match the level of proprioceptive signal (i.e., spindle discharge rates), the exercised muscle would have been stretched more than the control muscle by adopting a more extended arm position. Hence, it could be argued that Table 5 Mean values (±SE) of constant error (CE) in degrees for the three angles in all experimental conditions. 70° Pre-exercise Isometric Concentric Eccentric Total mean * Trend at p 6 0.09. 110° Mean value ±SE [95% CI] 0.02 0.58 0.78 0.35 0.03 ±1.1 ±1.5 ±1.1 ±1.2 ±1.1 [2.4 [2.9 [3.3 [2.4 [2.3 to to to to to 2.4] 4.0] 1.7] 3.1] 2.4] 150° Mean value ±SE [95% CI] 1.63 3.32 4.92 3.62 3.37* ±0.9 ±1.1 ±1.4 ±1.7 ±1.1 [3.6 [5.8 [8.1 [7.5 [5.8 to to to to to 0.3] 0.8] 1.7] 0.3] 0.9] Mean value ±SE [95% CI] 0.04 0.23 2.25 2.18 1.04 ±1.2 ±1.1 ±1.2 ±1.1 ±0.8 [2.7 [2.2 [5.0 [4.8 [2.8 to to to to to 2.8] 2.6] 0.5] 0.4] 0.8] S. Fortier et al. / Journal of Electromyography and Kinesiology 20 (2010) 298–304 the pattern of positional errors observed in this study is the result of exercise-related changes in the response of muscle spindles. Others studies have also indicated that the activity of muscle spindles declines during sustained voluntary contractions (Macefield et al., 1991; Vallbo, 1970). Bongiovanni et al. (1990) presented evidence that the fusimotor mediated spindle discharge would progressively decline during iMVC, and this decline was attributed to a progressive withdrawal of spindle-mediated fusimotor support to a-motoneurons. It has also been stated that the spindle afferents’ role would be different with concentric contractions because fusimotor drive may be insufficient to overcome muscle shortening. Consequently, spindle afferents in the contracting muscle may fall silent (Gandevia, 1998). This reduced stretch sensibility of spindles after voluntary concentric contractions could be one explanation for the increased threshold for muscle spindle discharge. This would be matched by the indicator arm adopting a more extended position, where the muscle and its spindles were subjected to a greater degree of stretch, and thus leading to a decline in position sense after concentric contractions, as seen by the main finding in the present study. On the other hand, disturbance in position sense was not significant in the eccentric experimental condition probably because, as mentioned by Gregory et al. (2004), intrafusal fibres of muscle spindles are not prone to damage of the kind seen in extrafusal fibres after a series of eccentric contractions. The difference in our exercise protocols compared to other studies could be one of the explanations for the dissimilarity in results. In the present study, high-intensity exercise protocols were used. These protocols led to a lower drop in iMVC than seen in the other studies using lower-intensity exercise protocols with higher number of muscle contractions. Different mechanisms may be involved when using a high-intensity exercise protocol which could be associated with different outcomes on position sense. Another explanation of the present results could be that our subjects adopted a more extended forearm position because the latter could have been subject to a smaller moment of the force of gravity. In fact, the most recent hypothesis in the literature is that accurate placement of forearms would be achieved by a combination of muscle spindle signals and effort-related signals. For instance, results from Walsh et al. (2004), Winter et al. (2005), and Allen and Proske (2006) have been interpreted as evidence in favour of an effort-related signal contributing to position sense during limb placement. They proposed that the effort required maintaining position of the arm against the force of gravity would provide us with information about its location in space. If subjects match efforts to align their arms they would place the fatigued arm more nearly vertically where less force is required to support it and where the same effort would be required to maintain its position. Further to their procedure, they concluded that subjects adopted of a more vertical arm position for two reasons. First, the moment of the force of gravity on the arm was less. Second, a more vertical position was closer to the elbow flexors’ optimum length for active tension (Walsh et al., 2004). Similarly to the aforementioned studies, after the concentric exercise our subjects placed their fatigued arm closer to the vertical, that is, a more flexed indicator arm position with the angle 70° (CE = 0.03° ± 1.04) and a more extended indicator arm position with the angles 110° (CE = 3.37° ± 1.09) and 150° (CE = 1.04° ± 0.79). These patterns could be explained by the smaller moment of the force of gravity acting on the fatigued arm. In addition, although not significant, it is worth noticing that CE tends to be larger at 110° than at 70° and 150° of elbow flexion (p < 0.09) which further supports the force of gravity effect (see Table 5). Therefore, results of the present study agree with those of Winter et al. (2005), that is, muscle spindles as well as sense of effort are responsible for the sense of position. Potential limitations to the study warrant considerations. First, the calculation of MF based on the assumption of non-stationary 303 signal may have had an impact on our EMG interpretation. However, the spectral EMG signal was analysed along with other parameters (i.e., post-iMVC and blood lactate) to confirm the occurrence of muscle fatigue during exercise tasks. Second, the use of iMVC to predict maximal concentric and eccentric muscle contractions may have led to over/underestimation of the true muscle contraction performances. Notwithstanding slight possible inaccuracies in force determinations, post-exercise tasks iMVCs were altered indicating that fatigue occurred with all types of muscle contraction. 5. Conclusion The hypothesis of this experiment was that eccentric exercise would result to a greater decrement in force leading to a larger impairment of position sense. The present results showed that a high-intensity concentric exercise protocol impaired position sense to a greater extent compared to the other types of muscle contractions despite a greater drop in iMVC after eccentric exercise task, which partly refute our assumptions. It is concluded that muscle spindle afferents and the effort required to support the limb against the force of gravity can provide information about forearm position. The nature of repetitive muscle contractions inducing fatigue should be taken into consideration to minimize a loss in movement accuracy in contexts such as research, sport performance and clinical rehabilitation. However, further research is needed to understand better muscle spindle and effort-related signal mechanisms related to muscle fatigue. Acknowledgments We thank all of the volunteers who participated in the project. We also thank Dr. Nicolas Forestier for his comments on the final version of the manuscript. The technical assistance of Lise Petrie is gracefully acknowledged and the School of Human Kinetics and Recreation for the use of laboratory facilities. References Allen TJ, Proske U. Effect of muscle fatigue on the sense of limb position and movement. Exp Brain Res 2006;170:30–8. Allen TJ, Ansems GE, Proske U. Effects of muscle conditioning on position sense at the human forearm during loading or fatigue of elbow flexors and the role of the sense of effort. J Physiol 2007;580:423–34. Bigland-Ritchie B, Johansson R, Lippold OC, Woods JJ. Contractile speed and EMG changes during fatigue of sustained maximal voluntary contractions. J Neurophysiol 1983;50:313–24. Billaut F, Basset FA, Giacomoni M, Lemaitre F, Tricot V, Falgairette G. Effect of highintensity intermittent cycling sprints on neuromuscular activity. Int J Sports Med 2006;27:25–30. Bongiovanni LG, Hagbarth KE, Stjernberg L. Prolonged muscle vibration reducing motor output in maximal voluntary contractions in man. J Physiol 1990;423:15–26. Brockett C, Warren N, Gregory JE, Morgan DL, Proske U. A comparison of the effects of concentric versus eccentric exercise on force and position sense at the human elbow joint. Brain Res 1997;771:251–8. The Canadian physical activity, fitness and lifestyle approach. 3rd ed. Ottawa: Canadian Society for Exercise Physiology; 2003. Clarkson PM, Newham DJ. Associations between muscle soreness, damage, and fatigue. Adv Exp Med Biol 1995;384:457–69. Colliander EB, Dudley GA, Tesch PA. Skeletal muscle fiber type composition and performance during repeated bouts of maximal, concentric contractions. Eur J Appl Physiol Occup Physiol 1988;58:81–6. Enoka RM. Eccentric contractions require unique activation strategies by the nervous system. J Appl Physiol 1996;81:2339–46. Faulkner JA, Brooks SV, Opiteck JA. Injury to skeletal muscle fibers during contractions: conditions of occurrence and prevention. Phys Ther 1993;73:911–21. Forestier N, Teasdale N, Nougier V. Alteration of the position sense at the ankle induced by muscular fatigue in humans. Med Sci Sports Exer 2002;34:117–22. Friden J, Lieber RL. Structural and mechanical basis of exercise-induced muscle injury. Med Sci Sports Exer 1992;24:521–30. Gandevia SC. Neural control in human muscle fatigue: changes in muscle afferents, moto neurones and motor cortical drive. Acta Physiol Scand 1998;162:275–83. 304 S. Fortier et al. / Journal of Electromyography and Kinesiology 20 (2010) 298–304 Gandevia SC, Refshauge K, Collins DF. Proprioception: peripheral inputs and perceptual interactions. In: Gandevia SC, Proske U, Stuart DG, editors. Sensorimotor control of movement and posture. Advances in experimental medicine and biology 2002;vol. 508. New York: Kluwer Academic/Plenum Publishers; 2002. p. 61–8. Gregory JE, Morgan DL, Proske U. Responses of muscle spindles following a series of eccentric contractions. Exp Brain Res 2004;157:234–40. Hagg GM. Interpretation of EMG spectral alterations and alteration indexes at sustained contraction. J Appl Physiol 1992;73:1211–7. Komi PV, Rusko H. Quantitative evaluation of mechanical and electrical changes during fatigue loading of eccentric and concentric work. Scand J Rehabil Med 1974;3:121–6. Lieber RL, Thornell LE, Friden J. Muscle cytoskeletal disruption occurs within the first 15 min of cyclic eccentric contraction. J Appl Physiol 1996;80:278–84. Macefield G, Hagbarth KE, Gorman R, Gandevia SC, Burke D. Decline in spindle support to alpha-motoneurones during sustained voluntary contractions. J Physiol 1991;440:497–512. McCloskey DI. Kinesthetic sensibility. Physiol Rev 1978;58:763–820. Saxton JM, Clarkson PM, James R, Miles M, Westerfer M, Clark S, et al. Neuromuscular dysfunction following eccentric exercise. Med Sci Sports Exer 1995;27:1185–93. Schmidtbleicher D. Strength training (part two): structural analysis of motor strength qualities and its application to training. Science Periodical on Research and Technology in Sport (Physical Training/Strength) 1985;W4:1–10. Skinner HB, Wyatt MP, Hodgdon JA, Conard DW, Barrack RL. Effect of fatigue on joint position sense of the knee. J Orthop Res 1986;4:112–8. Taylor AD, Bronks R, Smith P, Humphries B. Myoelectric evidence of peripheral muscle fatigue during exercise in severe hypoxia: some references to m. Vastus lateralis myosin heavy chain composition. Eur J Appl Physiol Occup Physiol 1997;75:151–9. Vallbo AB. Slowly adapting muscle receptors in man. Acta Physiol Scand 1970;78:315–33. Walsh LD, Hesse CW, Morgan DL, Proske U. Human forearm position sense after fatigue of elbow flexor muscles. J Physiol 2004;558:705–15. Walsh LD, Allen TJ, Gandevia SC, Proske U. Effect of eccentric exercise on position sense at the human forearm in different postures. J Appl Physiol 2006;100:1109–16. Winter JA, Allen TJ, Proske U. Muscle spindle signals combine with the sense of effort to indicate limb position. J Physiol 2005;568:1035–46. Sylvie Fortier is a M.Sc. degree holder and currently working as a Research Assistant in School of Human Kinetics and Recreation at Memorial University of Newfoundland, Canada. Her main research interests are Sports Performance and Motor Learning. Fabien A. Basset is a Ph.D. Degree holder and currently employed as an Assistant Professor in School of Human Kinetics and Recreation at Memorial University of Newfoundland, Canada. His main research interests are on Exercise Physiology and Sports Performance. François Billaut is a Ph.D. Degree holder and currently employed as an Assistant Professor in the Department of Kinesiology and Physical Education at University of Lethbridge, Canada. His main research interests are Muscle Fatigue, Sport Performance, and Sex Differences. David Behm is a Ph.D. Degree holder and currently employed as a Professor in School of Human Kinetics and Recreation at Memorial University of Newfoundland, Canada. His main research interests are Exercise Physiology, Muscle Fatigue, and Sports Performance. Normand Teasdale is a Ph.D. Degree holder and currently employed as a Professor in Groupe de Recherche en Analyse du Mouvement et Ergonomie at Université Laval, Canada. His research focus is on Motor Control.
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