Jussi Peltonen

Age-related differences in Achilles tendon properties and triceps surae muscle architecture in vivo

This study examined the concurrent age-related differences in muscle and tendon structure and properties. Achilles tendon morphology and mechanical properties and triceps surae muscle architecture were measured from 100 subjects [33 young (24 ± 2 yr) and 67 old (75 ± 3 yr)]. Motion analysis-assisted ultrasonography was used to determine tendon stiffness, Youngs modulus, and hysteresis during isometric ramp contractions. Ultrasonography was used to measure muscle architectural features and size and tendon cross-sectional area. Older participants had 17% lower (P < 0.01) Achilles tendon stiffness and 32% lower (P < 0.001) Youngs modulus than young participants. Tendon cross-sectional area was also 16% larger (P < 0.001) in older participants. Triceps surae muscle size was smaller (P < 0.05) and gastrocnemius medialis muscle fascicle length shorter (P < 0.05) in old compared with young. Maximal plantarflexion force was associated with tendon stiffness and Youngs modulus (r = 0.580, P < 0.001 and r = 0.561, P < 0.001, respectively). Comparison between old and young subjects with similar strengths did not reveal a difference in tendon stiffness. The results suggest that regardless of age, Achilles tendon mechanical properties adapt to match the level of muscle performance. Old people may compensate for lower tendon material properties by increasing tendon cross-sectional area. Lower tendon stiffness in older subjects might be beneficial for movement economy in low-intensity locomotion and thus optimized for their daily activities.

In vivo mechanical response of human Achilles tendon to a single bout of hopping exercise

SUMMARY Stiffness of the human Achilles tendon (AT) was determined in vivo before and after a single bout of hopping exercise. It was hypothesized, based on published data using in vitro specimens, that a reduction in AT stiffness may occur after just 1000 loading cycles at physiological stress levels. Ten healthy subjects performed two-legged hopping exercise consisting of 1150–2600 high impacts. Tendon stiffness was determined in several isometric ramp contractions [20%, 40%, 60%, 80% and 100% maximum voluntary contraction (MVC)] during which tendon elongation was measured using ultrasonography and two cameras. Tendon force was calculated by dividing measured ankle torque by magnetic resonance imaging-derived AT lever arm length. Tendon stiffness remained unchanged, being 430±200 N mm−1 before and 390±190 N mm−1 after the exercise [not significant (n.s.)]. Despite the lack of changes in stiffness, maximum tendon force during MVC was reduced from 3.5±0.6 kN to 2.8±0.7 kN (P<0.01). As the proposed decline in stiffness was not observed, it is concluded that mechanical fatigue did not take place in the AT of healthy individuals after a single bout of high-impact exercise performed until exhaustion.

Effect of time-of-day-specific strength training on maximum strength and EMG activity of the leg extensors in men

Abstract In this study, we examined the effects of time-of-day-specific strength training on maximum strength and electromyography (EMG) of the knee extensors in men. After a 10-week preparatory training period (training times 17:00–19:00 h), 27 participants were randomized into a morning (07:00–09:00 h, n = 14) and an evening group (17:00–19.00 h, n = 13). Both groups then underwent 10 weeks of time-of-day-specific training. A matched control group (n = 7) completed all testing but did not train. Unilateral isometric knee extension peak torque (MVC) and one-repetition maximum half-squat were assessed before and after the preparatory training and after the time-of-day-specific training at times that were not training-specific (between 09:00 and 16:00 h). During training-specific hours, peak torque and EMG during MVC and submaximum isometric contraction at 40% MVC were assessed before and after the time-of-day-specific training. The main finding was that a significant diurnal difference (P < 0.01) in peak torque between the 07:00 and 17:00 h tests decreased after time-of-day-specific training in the morning group but not in the evening or control groups. However, the extent of this time-of-day-specific adaptation varied between individuals. Electromyography during MVC did not show any time-of-day-specific adaptation, suggesting that peripheral rather than neural adaptations are the main source of temporal specificity in strength training.

Effects of contraction intensity on muscle fascicle and stretch reflex behavior in the human triceps surae

The aims of this study were to examine changes in the distribution of a stretch to the muscle fascicles with changes in contraction intensity in the human triceps surae and to relate fascicle stretch responses to short-latency stretch reflex behavior. Thirteen healthy subjects were seated in an ankle ergometer, and dorsiflexion stretches (8 degrees ; 250 degrees /s) were applied to the triceps surae at different moment levels (0-100% of maximal voluntary contraction). Surface EMG was recorded in the medial gastrocnemius, soleus, and tibialis anterior muscles, and ultrasound was used to measure medial gastrocnemius and soleus fascicle lengths. At low forces, reflex amplitudes increased despite a lack of change or even a decrease in fascicle stretch velocities. At high forces, lower fascicle stretch velocities coincided with smaller stretch reflexes. The results revealed a decline in fascicle stretch velocity of over 50% between passive conditions and maximal force levels in the major muscles of the triceps surae. This is likely to be an important factor related to the decline in stretch reflex amplitudes at high forces. Because short-latency stretch reflexes contribute to force production and stiffness regulation of human muscle fibers, a reduction in afferent feedback from muscle spindles could decrease the efficacy of human movements involving the triceps surae, particularly where high force production is required.

Differences in contractile behaviour between the soleus and medial gastrocnemius muscles during human walking

SUMMARY The functional roles of individual lower limb muscles during human walking may differ depending on walking speed or duration. In this study, 11 volunteers walked on a treadmill for 60 min at speeds corresponding to both optimal and 20% above optimal energetic cost of transport whilst oxygen consumption and medial gastrocnemius (MG) and soleus fascicle lengths were measured. Although energetic cost of transport was ∼12% higher at the faster speed, it remained constant over 60 min at both speeds, suggesting that humans can walk for prolonged periods at a range of speeds without compromising energetic efficiency. The fascicles of both muscles exhibited rather ‘isometric’ behaviour during the early to mid stance phase of walking, which appears to be independent of walking speed or movement efficiency. However, several functional differences were observed between muscles. MG exhibited time- and speed-dependent decreases in operating length, and shortened faster during the pushoff phase at the faster walking speed. Conversely, soleus exhibited consistent contractile behaviour regardless of walking speed or duration, and always shortened slower than MG during pushoff. Soleus appears to play a more important functional role than MG during walking. This may be especially true when walking for prolonged periods or at speeds above the most energetically efficient, where the force potential and thus the functional importance of MG appears to decline.

Viewpoint: On the hysteresis in the human Achilles tendon

This viewpoint was stimulated by two observations: 1) the statistical skewness whereby numerous articles have reported tendon stiffness and Young9s modulus, but far fewer have reported tendon hysteresis; 2) in vivo human studies seem very often to report hysteresis values greater than 10%, suggesting either that there are methodological differences between human and animal studies, or that human tendons have a much poorer capacity to store and reutilize elastic energy. In this article we focus on the healthy human Achilles/gastrocnemius tendon (AT) since it has an important locomotor function and clearly a low AT hysteresis would allow elastic recoil for efficient locomotion. We discuss that both the measurement of tendon length and force and their correct synchronization can contribute to the variability of measures of tendon properties. Within the large variability the lower values are likely to be more valid, being consistent with animal studies.

Achilles tendon length changes during walking in long-term diabetes patients

BACKGROUND Diabetes leads to numerous side effects, including an increased density of collagen fibrils and thickening of the Achilles tendon. This may increase tissue stiffness and could affect stretch distribution between muscle and tendinous tissues during walking. The primary aim of this study was to examine stretch distribution between muscle and tendinous tissues in the medial gastrocnemius muscle-tendon unit in long-term diabetes patients and control subjects during walking. METHODS Achilles tendon length changes were investigated in 13 non-neuropathic diabetes patients and 12 controls, whilst walking at a self selected speed across a 10 m force platform. Electromyographic activity was recorded in the medial gastrocnemius, soleus and tibialis anterior muscles, goniometers were used to detect joint angle changes, and ultrasound was used to estimate tendon length changes. FINDINGS Achilles tendon length changes were attenuated in diabetes patients compared to controls, and were inversely correlated with diabetes duration (r=-0.628; P<0.05), as was ankle range of motion (r=-0.693; P<0.01). Tendon length changes were also independent of walking speed (r=-0.299; P=0.224) and age (r=0.115; P=0.721) in the diabetic group. INTERPRETATION Stretch distribution between muscle and tendon during walking is altered in diabetic patients, which could decrease walking efficiency, a factor that may be exacerbated with increasing diabetes duration. Diabetes-induced changes in mechanical tendon properties may be at least partly responsible for attenuated tendon length changes during walking in this patient group.

Achilles tendon stiffness is unchanged one hour after a marathon

SUMMARY Overuse-induced injuries have been proposed as a predisposing factor for Achilles tendon (AT) ruptures. If tendons can be overloaded, their mechanical properties should change during exercise. Because there data are lacking on the effects of a single bout of long-lasting exercise on AT mechanical properties, the present study measured AT stiffness before and after a marathon. AT stiffness was determined as the slope of the force–elongation curve between 10 and 80% of maximum voluntary force. AT force–elongation characteristics were measured in an ankle dynamometer using simultaneous motion-capture-assisted ultrasonography. Oxygen consumption and ankle kinematics were also measured on a treadmill at the marathon pace. All measurements were performed before and after the marathon. AT stiffness did not change significantly from the pre-race value of 197±62 N mm−1 (mean ± s.d.) to the post-race value of 206±59 N mm−1 (N=12, P=0.312). Oxygen consumption increased after the race by 7±10% (P<0.05) and ankle kinematic data revealed that in nine out of 12 subjects, the marathon induced a change in their foot strike technique. The AT of the physically active individuals seems to be able to resist mechanical changes under physiological stress. We therefore suggest that natural loading, like in running, may not overstress the AT or predispose it to injury. In addition, decreased running economy, as well as altered foot strike technique, was probably attributable to muscle fatigue.

Neuromuscular fatigue induced by an isotonic heavy-resistance loading protocol in knee extensors.

Abstract The main aim of this study was to assess neuromuscular fatigue during a typical high-load, low-repetition loading protocol. Muscle stimulations were used to assess maximum voluntary contraction, resting single- and double-pulse twitch characteristics, and superimposed double-pulse twitch force (used to calculate voluntary activation) before and after an acute knee extension loading protocol. In our participants, who had previous resistance training experience, the mean voluntary activation level was 96.2% in an unfatigued state. Maximum voluntary contraction (−11.8%), resting double-pulse twitch force (−10.6%), and voluntary activation (−2.1%) were markedly decreased as a consequence of loading (P < 0.05). In addition, although potentiated twitch characteristics were observed during the loading protocol, this was short-lived, as fatigue surpassed the potentiation mechanisms. Our results show that both central and peripheral mechanisms contributed to neuromuscular fatigue during the present loading protocol.

Triceps surae muscle-tendon properties in older endurance- and sprint-trained athletes

Previous studies have shown that aging is associated with alterations in muscle architecture and tendon properties (Morse CI, Thom JM, Birch KM, Narici MV. Acta Physiol Scand 183: 291-298, 2005; Narici MV, Maganaris CN, Reeves ND, Capodaglio P. J Appl Physiol 95: 2229-2234, 2003; Stenroth L, Peltonen J, Cronin NJ, Sipila S, Finni T. J Appl Physiol 113: 1537-1544, 2012). However, the possible influence of different types of regular exercise loading on muscle architecture and tendon properties in older adults is poorly understood. To address this, triceps surae muscle-tendon properties were examined in older male endurance (OE, n = 10, age = 74.0 ± 2.8 yr) and sprint runners (OS, n = 10, age = 74.4 ± 2.8 yr), with an average of 42 yr of regular training experience, and compared with age-matched [older control (OC), n = 33, age = 74.8 ± 3.6 yr] and young untrained controls (YC, n = 18, age = 23.7 ± 2.0 yr). Compared with YC, Achilles tendon cross-sectional area (CSA) was 22% (P = 0.022), 45% (P = 0.001), and 71% (P < 0.001) larger in OC, OE, and OS, respectively. Among older groups, OS had significantly larger tendon CSA compared with OC (P = 0.033). No significant between-group differences were observed in Achilles tendon stiffness. In older groups, Youngs modulus was 31-44%, and maximal tendon stress 44-55% lower, than in YC (P ≤ 0.001). OE showed shorter soleus fascicle length than both OC (P < 0.05) and YC (P < 0.05). These data suggest that long-term running does not counteract the previously reported age-related increase in tendon CSA, but, instead, may have an additive effect. The greatest Achilles tendon CSA was observed in OS followed by OE and OC, suggesting that adaptation to running exercise is loading intensity dependent. Achilles tendon stiffness was maintained in older groups, even though all older groups displayed larger tendon CSA and lower tendon Youngs modulus. Shorter soleus muscle fascicles in OE runners may be an adaptation to life-long endurance running.