Journal of Applied Biomechanics, 2015, 31, 69 -78 Human Kinetics http://dx.doi.org/10.1123/JAB.2013-0284 © 2015 Human Kinetics, Inc. ORIGINAL RESEARCH Eccentric and Concentric Exercise of the Triceps Surae: An In Vivo Study of Dynamic Muscle and Tendon Biomechanical Parameters Saira Chaudhry, Dylan Morrissey, Roger C. Woledge, Dan L. Bader, and Hazel R.C. Screen Queen Mary University of London Triceps surae eccentric exercise is more effective than concentric exercise for treating Achilles tendinopathy, however the mechanisms underpinning these effects are unclear. This study compared the biomechanical characteristics of eccentric and concentric exercises to identify differences in the tendon load response. Eleven healthy volunteers performed eccentric and concentric exercises on a force plate, with ultrasonography, motion tracking, and EMG applied to measure Achilles tendon force, lower limb movement, and leg muscle activation. Tendon length was ultrasonographically tracked and quantified using a novel algorithm. The Fourier transform of the ground reaction force was also calculated to investigate for tremor, or perturbations. Tendon stiffness and extension did not vary between exercise types (P = .43). However, tendon perturbations were significantly higher during eccentric than concentric exercises (25%–40% higher, P = .02). Furthermore, perturbations during eccentric exercises were found to be negatively correlated with the tendon stiffness (R2 = .59). The particular efficacy of eccentric exercise does not appear to result from variation in tendon stiffness or extension within a given session. However, varied perturbation magnitude may have a role in mediating the observed clinical effects. This property is subject-specific, with the source and clinical timecourse of such perturbations requiring further research. Keywords: Achilles tendon, force, extension, biomechanics, human, in vivo Achilles tendinopathy is difficult to treat1 and prone to recurrence,2–5 with current treatments achieving only partial success. Available data indicate that one of the most effective treatments for Achilles tendinopathy is eccentric exercise of the triceps surae, with a success rate of 50%–60% in sedentary patients5 and 60%–80% in athletes.6–8 Controlled trials of this method report superior outcomes with eccentric exercise (eccentric) when compared with surgical interventions6 or concentric exercise (concentric) regimes.7 However, the mechanisms underpinning the observed effects remain unknown. Initially it was believed the tendon experienced higher forces during eccentric than concentric, resulting in more tendon remodeling in response to higher strain.9 However, in vivo studies in healthy humans have reported no difference in peak Achilles tendon force during eccentric and concentric,10 instead reporting that the primary difference between eccentric and concentric is 8–12 Hz tendon perturbations during eccentric.11 However, these studies only considered tendon force or length changes in isolation, and to date no one has investigated stiffness across the complete dynamic force-length curve of the Achilles tendon during eccentric and concentric, nor how stiffness may relate to tendon perturbation behavior. Establishing relationships between tendon stiffness and perturbations is Saira Chaudhry is with the School of Engineering and Material Science and the Centre for Sports and Exercise Medicine, both at Queen Mary University of London, UK. Dylan Morrissey and Roger C. Woledge are with the Centre for Sports and Exercise Medicine, Queen Mary University of London, UK. Dan L. Bader and Hazel R.C. Screen are with the School of Engineering and Material Science, Queen Mary University of London, UK. Address author correspondence to Hazel R.C. Screen at h.r.c.screen@qmul.ac.uk. an essential step toward understanding the mechanical effects of exercise within tendon. The frequency of these perturbations and their effects may vary depending on the stiffness of the tissue.12,13 To measure tendon length changes during exercise, typically, manual marking of the muscle-tendon junction (MTJ) in successive ultrasound frames has been adopted.14,15 However, manual analysis of MTJ location is labor intensive; a 15 second video at 17 Hz consists of 255 frames, so tracking is generally limited to feature monitoring in 1–2 frames during an exercise cycle. In addition, while the human eye appears very good at image correlation, the accuracy and reproducibility of this approach cannot be guaranteed, and manual methods fail if movements are large and key features move out of view. There have been a number of attempts to develop algorithms to analyze MTJ movement automatically,16–20 typically based on cross-correlation or Lucas-Kanade feature tracking, and successfully tested on relatively small movements. However, the inherent limitation of these methods, that the same feature of interest must be present in similar form in adjacent frames of interest, persists and limits their application in the current study, as the MTJ will sometimes move off the screen. An automatic method suitable for large movements with intermittent key feature visibility was developed to address these requirements. The aim of the current study was to investigate how tendon mechanical properties and tendon perturbations compared between eccentric and concentric exercise cycles. We hypothesized that tendon mechanical properties are the same during eccentric and concentric throughout the exercise cycle. Further, we hypothesized that the perturbations seen during eccentric correlate negatively with tendon stiffness. To test these hypotheses, this study additionally describes the development and testing of a novel semiautomated high-strain tracking analysis procedure for the MTJ. 69 70 Chaudhry et al. Methods Subjects The study was approved by the ethics and research committee of Queen Mary University of London. Eleven healthy volunteers (6 male, 5 female; mean [SD] age 26.5 years [1.9]; body mass 65.92 kg [10.5]; height 1.73 m [0.08]) were recruited, giving written consent. Subjects were moderately active, defined as 2–3 hours of physical activity a week, but had not participated in any organized program of competitive sport over the last year and had no history of tendon injury or any systemic disease. Subjects with current or previous Achilles tendon pain, pathology, or surgery were excluded. Exercise Protocol For performing exercises, each subject was asked to stand on a step placed on a force plate (Kistler force platform 9281B, Kistler Instruments, Winterthur, Switzerland). For eccentric, subjects started on the ball of the foot of the right leg with the heel raised, lowering the heel in a controlled manner (Figure 1, right to left). The exercise was performed off the edge of the step to allow full dorsiflexion to be reached. For concentric, subjects started with the heel below the toes and raised the heel in a similar controlled manner (Figure 1, left to right). Both exercises were performed at 0.5 rad∙s–1 (~3 s exercise phase), guided by a metronome. After completing either exercise, subjects used their other leg and a second step on an adjacent force plate to assist in returning to the starting position before repeating the exercise. A single data collection consisted of two cycles of either concentric or eccentric performed consecutively. Three sets of data were recorded for each exercise paradigm, for each subject, in a randomized order. Time was allowed to perform familiarization exercises before data collection, with these exercises also providing preconditioning of the triceps surae muscle-tendon unit to ensure minimal variation in the load-deformation curves.21 Tendon forceextension data were collected in conjunction with electromyography (EMG) and tendon perturbations as outlined below. Measurement of Tendon Force Tendon force was calculated from torque around the ankle, using inverse dynamics. The 3D ground reaction force (GrF) was captured using a 600 × 400 mm2 force plate at 1000 Hz. An active infrared Figure 1 — Schematic highlighting the range of motion adopted during eccentric (EL) and concentric (CL), from full plantar flexion to full dorsiflexion. Each exercise was performed on a wooden box, placed on a force plate embedded in the ground. Electromyography (EMG) was recorded by electrodes placed on the calf muscle (denoted with x) and joint motion was tracked by placing CODA markers on the leg joints (denoted by •). An ultrasound probe tracked the muscle-tendon junction; markers were placed on the probe to monitor its movement. JAB Vol. 31, No. 2, 2015 Eccentric and Concentric Exercise of Triceps Surae 71 motion analysis system (CODA, CX1, Charnwood Dynamics, Rothley, UK) with four cameras was used to determine the moments around the ankle and knee joint by placing markers on the lateral and medial malleoli, metatarsal 1 and 5, calcaneus, tendon (midpoint), medial and lateral tibial condyle, tibial tuberosity, femoral epicondyle, and hip joint (Figure 1). The motion analysis global coordinate system was aligned to the force plate. The ankle joint rotation center was estimated as a virtual point half the distance between the lateral and medial malleoli.22 The perpendicular distance to the ankle joint center from the line joining the calcaneus marker and the Achilles tendon marker was taken as the moment arm after correction for skin thickness measured by ultrasound (US). Tendon force was calculated by dividing the externally applied ankle joint moment by the moment arm and normalized across subjects by body weight. Torque due to inertia around the foot was ignored, as movements were slow. However, a complete calculation of angular torque around the ankle, including forces exerted on the foot by gravity and acceleration, was carried out on three subjects to validate this assumption. Measurement of Tendon Elongation Advances in ultrasonography and motion analysis have enabled simultaneous use to determine in vivo human muscle-tendon complex dynamics.23 In the current study, a technique to measure continual tendon elongation was developed to measure tendon mechanical behavior throughout exercise. Tendon length was defined as the distance between the Achilles tendon insertion and the distal MTJ of the medial gastrocnemius (Figure 2). The point of Achilles tendon insertion was tracked using a single marker placed on the calcaneus. The MTJ was tracked using combined CODA and an ultrasound system (Voluson e, GE Healthcare, UK; wide band, multifrequency linear transducer, frequency = 3.7–11.3 MHz, FOV = 37.4 mm, FR = 17Hz). Two markers were placed along the ultrasound probe axis while a third marker was placed approximately 60 mm from the midpoint of the line intersecting the initial markers (Figure 2). This enabled the probe position to be tracked and spatially synchronized with the motion data. A synchronizing system was used to trigger motion capture to ensure accurate ultrasound frame attribution. This consisted of a TTL switch coupled to the digital input of the CODA active hub, providing a triggering pulse to start data acquisition at the commencement of the US scan with a maximum temporal lag of 55 ms (manufacturer values). The CODA active hub was electronically integrated with the force plates and EMG system, enabling synchronization of all four devices. Tracking of the Muscle-Tendon Joint Tracking the MTJ throughout the US video required identifying the precise location of the intersection of the superficial and deep connective tissue in every frame. To reduce human error, improve accuracy, and increase speed, an algorithm was developed and implemented in MatLab (Version 7.9.0.529 [R2009b], 32-bit, MathWorks, Natick, MA). To automate tracking, the projection of the two brightest lines (Figure 3A) was established along the radial line, and their relative orientation angle determined using the Radon transform. Briefly, the Radon transform provides an intensity map of an image, from which the 2D location and angular orientation of the most intense image points can be determined. Extracting data from the most intense points enabled the MTJ to be located from the intersection of the eight brightest lines each side of the junction (Figure 3B). From the 8 × 8 matrix of brightest lines, the 64 intersection Figure 2 — Muscle-tendon junction (MTJ) tracking to establish Achilles tendon length. C and C’ denote the calcaneus position at full plantar flexion and dorsiflexion, respectively. The central images indicate the movement of the MTJ during exercise, from MTJ to MTJ’ at the two extreme positions. The distance between C and MTJ at any point in time is defined as the Achilles tendon length. A force record is also shown demonstrating the change in force with the movement. US = ultrasound. JAB Vol. 31, No. 2, 2015 72 Chaudhry et al. Figure 3 — (A) Part of a single ultrasound video frame (480 × 190 pixels) showing the muscle-tendon junction (MTJ). (B) The same image with superimposed lines. The lower lines are the eight brightest lines selected from the 231 Radon transform. These lie within the angle range 93º–103º with intercepts within the range of 54–74 pixels. The upper lines are a similar set for the angle range 79º–89º and intercept range of 107–127. The mean angle and mean intercepts of these lines is used to set the target range for analyzing the next frame of the video. (Note. Angles in the Radon transform are measured with respect to the vertical in these figures, and the intercepts are with a line perpendicular to each Radon line passing through the center of the part of the frame analyzed (white + in B). (C) The same image marked in white with the 64 points at which each pair of lines (ie, one from the top set and one from the bottom set) approach to within 10 pixels. The black cross shows the median coordinates of these 64 points, and is the value used for the position of the MTJ within this video frame. points were located, and the junction taken as the median of these (Figure 3C). Reliability of this algorithm was assessed by comparing results from three separate attempts to track each video. In addition, it was compared with the manual method of MTJ tracking using Image J software (version 1.44o, NIH, Bethesda, MD). Assessment of Accuracy in Ultrasound Measurements The accuracy of the US setup for tracking movement was further assessed using a phantom. A metallic wand of known dimensions was immersed in a container of ultrasound gel, such that the container of gel represented the leg and the wand represented the tendon. Along the length of the wand, a metallic coil was placed to represent the MTJ. Moving the wand within the container, the coil was tracked throughout the movement using the ultrasound system. To assess US tracking accuracy, wand and coil displacement were also measured directly using CODA markers, and the two data sets compared. Measurement of Muscle Activation EMG recordings were made using Ag/AgCl dual snap electrodes with a 20 mm interelectrode distance. After careful skin preparation, electrodes were placed on the belly of the soleus, lateral gastrocnemius, medial gastrocnemius, and tibialis anterior muscles following the SENIAM guidelines24 and a single, self-adhesive Ag/ AgCl snap electrode placed on the anterior tibia as a reference. EMG signals were preamplified and band-pass filtered between 10 and 1000 Hz before sampling at 1500 Hz using a wireless EMG system (Telemyo 2400T G2, Noraxon, Scottsdale, AZ; input impedance > 100 MΩ, common mode rejection ratio > 100 dB, base gain 500) with 16-bit analog-to-digital resolution. Data Analysis Data analysis was carried out in MatLab. For each exercise task, only cycles within 10% of the prescribed speed were retained for analysis. JAB Vol. 31, No. 2, 2015 Eccentric and Concentric Exercise of Triceps Surae 73 To ensure only the loading phase of each exercise was assessed, points of minimum and maximum heel height were identified and the portion of the record between them resampled by interpolation to 111 uniformly spaced points. Times of single-leg support were confirmed by absence of a force reading from the plate under the contralateral foot. For descriptive purposes, ankle angles were extracted during each test. After registration, GrF and EMG data were obtained for each subject. To establish muscle activation, the EMG signal was full-wave rectified, then smoothed using a moving median filter with a window size of 121. To compare posterior and anterior compartment activation, the contribution of the gastrocnemius and soleus were summed for posterior activation. Anterior data were taken directly from the tibialis anterior. Having established appropriate subject repeatability over repeat exercises, mean force-length curves for each subject were produced over the three repeats of eccentric and concentric. Apparent tendon stiffness was measured from the slope of the mean force-length curve for each individual, placing a linear regression line through the data. After registration, the vertical component of the GrF vector was further analyzed to investigate for tremor (defined throughout the text as perturbations) in the GrF, and subsequently in the Achilles. First, the mean value was subtracted to make GrF independent of the body mass. To obtain the magnitude and frequency of the perturbations, power spectrum densities were calculated using a fast Fourier transformation (FFT) method, after elimination of the dc component. The power was subsequently summed within 1 Hz windows across the frequency range 0–16 Hz. Finally, any correlation between tendon stiffness and the extent of perturbation in the GrF and Achilles was investigated across subjects. Comparing eccentric and concentric across the subject population showed no significant differences in force-displacement behavior during the two exercises for most subjects. Range of motion was also similar; 47.94 ± 1.83° during eccentric and 47.05 ± 1.93° during concentric, with a shortening velocity of 8.4 ± 1.5 mm/s during eccentric and 7.9 ± 1.1 mm/s during concentric. Unsurprisingly, this resulted in no significant variation in tendon stiffness between eccentric and concentric. Intrasubject variability was far greater, and concentric and eccentric resulted in distinctly different temporal loading patterns across subjects. Eccentric and concentric mean force-length curves were produced for each individual, with example data for six individuals shown in Figure 4. It is interesting to note that for some individuals there is a clear difference in tendon behavior between eccentric and concentric. There were also notable variations in tendon stiffness between subjects. To enable a direct comparison of tendon properties during eccentric and concentric across all subjects, normalized force and displacement were derived. Figure 5 shows mean forcelength behavior across all subjects during eccentric and concentric, with the time axis reversed for eccentric, for ease of comparison. Interestingly, clear temporal differences between eccentric and concentric were apparent for both tendon force and length (Figure 5A, B). However, these differences were lost in mean force-length data (Figure 5C). After carrying out paired comparisons of eccentric and concentric across all subjects, no significant differences were observed in maximum extension (P = .07) or stiffness (P = .06). Statistical Analysis Normality tests were performed on the data using the Shapiro-Wilk method (OriginPro version 8, OriginLab, Northampton, MA). Once the data were found to be normally distributed, paired t tests were used to examine differences in tendon force, perturbations, muscle activation, and tendon extension for each exercise. For all statistical tests, significance was established at P ≤ .05. Data are presented as mean (SD). To assess accuracy of US tracking, the correlations between tracking distances measured by ultrasound and motion markers were determined, and the intraclass correlation was calculated. Systematic bias between the two measures was also investigated using a paired t test. Results Using the combined motion analysis and US capture techniques to scan movement of the phantom revealed that US could determine position to an accuracy of ±1.65 mm. The repeatability of the MTJ tracking algorithm was found to be ±0.59 × 0.36 mm for tracking a video multiple times. Good correlation was found between automatic and standard tracking methods with an intraclass correlation coefficient of 0.98 for the x axis (95% confidence interval 0.94–0.99) and 0.62 (95% confidence interval 0.26–0.88) for the y axis. No systematic difference between data sets was observed (P = .12 in x direction and P = .09 in y direction). There was less than 1% difference in the tendon forces calculated from torque around the ankle with and without the inclusion of inertial effects, hence inertia around the foot was ignored. Figure 4 — Graphs of the mean Achilles tendon force-length (ATF-ATL) relationship for six subjects, shown in separate panels. White squares report eccentric data and black circles report concentric data. Some subjects show different loading behavior between eccentric and concentric. JAB Vol. 31, No. 2, 2015 74 Chaudhry et al. Figure 5 — A comparison of (A) normalized mean Achilles tendon force (± SE) and (B) normalized mean Achilles tendon length (± SE) against time during the loading cycle. (C) Achilles tendon force versus Achilles tendon length and (D) electromyography (EMG) data comparing activation in the posterior muscles (circles) and anterior muscles (squares) during eccentric and concentric across all participants. Eccentric data are shown as open circles/squares whereas concentric data are filled circles/squares. Tendon apparent stiffness varied across the subject population, but not between exercise types (t = .82, P = .43), with mean (± SE) values of 62.79 N∙mm–1 (9.37) during concentric and 58.89 N∙mm–1 (9.21) during eccentric. Considering the EMG data (Figure 6D), muscle activation during the cycle and maximal muscle activation values were significantly higher in concentric than eccentric (P < .01) for both the anterior compartment (concentric = 0.030 mV [SE 0.004], eccentric = 0.025 mV [SE 0.003]) and the posterior compartment (concentric = 0.140 mV [SE 0.014], eccentric = 0.118mV [SE 0.011]) of the triceps surae muscle group. A greater amplitude of higher frequency force perturbations were present during eccentric (P = .02) at 10 Hz and showed negative correlation with tendon stiffness at 10 Hz. Summary data showing the FFT of tendon force (Figure 6C) demonstrated that the main amplitude of the power component was typically seen at frequencies below 5 Hz, with a peak around 2–3 Hz, representative of low frequency movements. However, a range of vibration frequencies of JAB Vol. 31, No. 2, 2015 Eccentric and Concentric Exercise of Triceps Surae 75 A C B D Figure 6 — (A) Example data set showing normalized Achilles tendon force (ATF) and normalized ground reaction force (GrF) for one subject during eccentric. The data are detrended, removing low frequency force change corresponding with movement, to focus on perturbations. (B) The corresponding fast Fourier transformation (FFT) of the GrF data. (C) Mean FFT of Achilles tendon force across all subjects for eccentric (empty circles) and concentric (filled squares) (± SE). (D) The correlation between magnitude of perturbations at 10 Hz and tendon stiffness during eccentric across the subject population. A negative correlation was observed (R2 = .59). up to 15 Hz were present in all subjects. Statistically significant differences between eccentric and concentric were observed at around 10 Hz, where tremor during eccentric was greater. Further, the magnitude of 10 Hz perturbations was negatively correlated with tendon stiffness (Figure 6D). However, not every subject showed a higher power of perturbations at around 10 Hz during eccentric (7 out of 11). Discussion The current study shows that eccentric and concentric do not differ in terms of tendon force and stiffness behavior during exercise. However, there is a clear difference in the magnitude of perturbations generated by these exercises, with perturbations at around 10 JAB Vol. 31, No. 2, 2015 76 Chaudhry et al. Hz generated specifically during eccentric exercise. Furthermore, data highlights that the magnitude of tendon perturbations is inversely correlated to tendon stiffness. To collate these findings, this paper presents a novel semiautomated method of measuring of tendon extension during high-strain movement, which increased the efficiency of analyzing one video of 200 frames 10 times as compared with manual tracking. These findings also provide a solution, independent of the human eye, able to track large dynamic movements when the region of interest moves out of the field of view. Such an algorithm can be very useful in different biomechanical studies based around MTJ tracking. Systematic error is always to be considered with any such measurement technique. However, by placing markers on the US probe to account for its movement, the use of US to measure tendon extension has previously been validated and considered reliable.23 However, the US image tracking technique developed and tested during this study can still be sensitive to temporal lag and image quality, and skill and practice are required to track the MTJ accurately. Temporal lag was less than 2% of the movement cycle and unlikely to significantly affect data. Blurred or low intensity images prevented accurate tracking, thus videos that did not meet the quality standards were not included in the analysis. However, of 88 videos collected, only 8 were below the required standard. In addition, while the technique was shown to be reproducible for one participant, individual to individual variability may occur, owing to factors such as differences in individual muscle architecture and changes in tissue shape during movement. These may alter the apparent location of the MTJ which would in turn influence strain and stiffness measures. Furthermore, the use of a constant value for tendon thickness may increase interindividual variability. However, using identical test procedures and a single tester in the current study minimized error and enabled eccentric and concentric to be successfully compared. Indeed, no significant differences in tendon mechanical properties or muscle activity data were observed, even when the same subject was tested on two consecutive days, as long as the ROM, subject foot placement, and US probe position were consistent between the measures. This was ensured by carefully marking the positions for the second day of measures. At the start of a concentric movement from full dorsiflexion, the calf muscles are activated, increasing in activation as they accelerate the subject upwards. Tendon force is at its peak at the start of the concentric movement, possibly as the triceps surae moment arm is furthest from the ankle joint axis for sagittal plane movement at this point. As the subject rises to plantar flexion, force reduces. In contrast, subjects start an eccentric movement with the heel at its highest point, and lower ‘under control’. The movement is controlled (resisted) by lengthening of the activated calf muscle and stretch of the Achilles tendon. Maximal force occurs at the end of the movement, again at full dorsiflexion, when maximum force is required to decelerate the subject against gravity. It should be noted that the tendon was always loaded during the movement and therefore the force-extension behavior seen is similar to that evident in the linear region of an in vitro test curve, with muscle contraction likely acting to strain the tendon beyond the low stiffness toe region seen when testing tendon in vitro. Normalizing force curves to heel height enables the change in force during each movement to be readily compared. From these data, significantly higher forces in the tendon were observed during concentric for a small range of ankle angles when the foot was fully dorsiflexed, and the forces passing through the tendon are maximum. However, no overall significant differences in force during the loading cycle were observed between eccentric and concentric. Such a finding is consistent with a previous study,10 comparing similar eccentric and concentric exercises. However, Rees et al10 only reported the maximum forces during eccentric and concentric. This study has additionally reported tendon stiffness and its relationship with 10 Hz perturbations. Also in agreement with previous studies,25,26 our data highlighted comparatively lower EMG activity during eccentric than concentric. Muscle activation within the medial gastrocnemius, lateral gastrocnemius, and soleus were all significantly lower during eccentric (P < .01). By contrast, there was no significant difference in the antagonistic tibialis anterior muscle activation between eccentric and concentric. Such a response from the tibialis anterior also follows previously reported findings.27 However, there is variability considering data from a range of different studies, as a result of different test methodologies.28,29 Individual variations in tendon properties, particularly stiffness, were evident across the group of participants. However, no significant differences in tendon stiffness between eccentric and concentric were reported. Repeats by an individual were similar, and reported tendon properties were within the range of previously published data.15,30,31 The variation in Achilles tendon stiffness between individuals could be partly dependent on previous training;27 although all subjects in the study performed only moderate activity, the effect of specific training cannot be ruled out. In addition, measurements were undertaken following a clinicallyrelevant protocol, where no control for the muscle activation was considered, a factor previously shown to affect stiffness measurements.32,33 However, to get a true understanding of the differences between eccentric and concentric, it was necessary to carry out measurements during the typical protocol and not under isometric conditions. By reporting whole curves, we have shown for the first time the extent of variability in tendon loading curves between individuals, which could be relevant to understanding the variable patient responses to therapy. We have also shown temporal differences between eccentric and concentric for the first time, which could be of further interest. Strain is an important stimulus for tendon repair. A number of studies have indicated that mechanical loading is an essential stimulus for tendon repair,34,35 initiating tenocyte signaling mechanisms in a strain dependent manner.36 Results from the current study indicate that eccentric and concentric, performed under equal load and at the same speed, result in different temporal strains within the tendon (Figure 5A, B). However, if the data are plotted for normalized position (Figure 5C), this difference disappears. Consequently, differences in strain alone are unlikely to be the differentiating trigger for repair during eccentric exercise, although the situation may differ in patients with tendinopathy. By contrast, tendon perturbations have been shown to differ between eccentric and concentric, and were higher during eccentric, particularly around 10 Hz (P < .05). Significantly higher perturbations during eccentric, in the 8–12 Hz range, have been reported previously.11 However, the origin of these perturbations has not previously been considered. The current study assessed the correlation between perturbations and tendon stiffness and found that while perturbation frequency does not change with tendon stiffness, the magnitude of tendon perturbations at 10 Hz is inversely correlated to tendon stiffness. This suggests the perturbations are not a result of mechanical resonance, as previous studies suggest a correlation between perturbation frequency and tendon stiffness should be present in such a situation.12,13,37,38 However, the negative correla- JAB Vol. 31, No. 2, 2015 Eccentric and Concentric Exercise of Triceps Surae 77 tion between tendon stiffness and perturbation magnitude at 10 Hz during eccentric does show some influence of tendon mechanics on resulting perturbations. This relationship is interesting, as tendon stiffness has previously been reported to decrease in tendinopathic tissue,39 which would imply that 10 Hz perturbations would be greater in injured tendon.40 Indeed, an increased magnitude perturbation has recently been reported.41 It is important to note that not every subject showed 10 Hz perturbations, implying that not every patient performing eccentric would receive this stimulus. If this stimulus has some relationship with tendinopathy, this may partly explain why not all subjects respond to eccentric. However, further analysis of perturbations is needed to draw firm conclusions. It is still unclear whether perturbations are a beneficial stimulus for tendon repair or, in fact, indicate a muscle weakness or poorly synchronized muscle activity in the triceps surae, both of which may be risk factors for tendinopathy. It may be that eccentric treats tendinopathy by training the triceps surae to function more effectively and minimize perturbation, with a progressive load and speed challenge incorporated as the triceps surae adapts to the training stimuli. There is a lack of consensus concerning the management of tendinopathy, which has limited options for developing new and more effective treatments. This study has given further insight into the biomechanical behavior of the triceps surae during physiologically representative eccentric and concentric exercises. 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