Erik B. Simonsen

A mechanism for increased contractile strength of human pennate muscle in response to strength training: changes in muscle architecture

1 In human pennate muscle, changes in anatomical cross‐sectional area (CSA) or volume caused by training or inactivity may not necessarily reflect the change in physiological CSA, and thereby in maximal contractile force, since a simultaneous change in muscle fibre pennation angle could also occur. 2 Eleven male subjects undertook 14 weeks of heavy‐resistance strength training of the lower limb muscles. Before and after training anatomical CSA and volume of the human quadriceps femoris muscle were assessed by use of magnetic resonance imaging (MRI), muscle fibre pennation angle (θp) was measured in the vastus lateralis (VL) by use of ultrasonography, and muscle fibre CSA (CSAfibre) was obtained by needle biopsy sampling in VL. 3 Anatomical muscle CSA and volume increased with training from 77.5 ± 3.0 to 85.0 ± 2.7 cm2 and 1676 ± 63 to 1841 ± 57 cm3, respectively (±s.e.m.). Furthermore, VL pennation angle increased from 8.0 ± 0.4 to 10.7 ± 0.6 deg and CSAfibre increased from 3754 ± 271 to 4238 ± 202 μm2. Isometric quadriceps strength increased from 282.6 ± 11.7 to 327.0 ± 12.4 N m. 4 A positive relationship was observed between θp and quadriceps volume prior to training (r = 0.622). Multifactor regression analysis revealed a stronger relationship when θp and CSAfibre were combined (R= 0.728). Post‐training increases in CSAfibre were related to the increase in quadriceps volume (r = 0.749). 5 Myosin heavy chain (MHC) isoform distribution (type I and II) remained unaltered with training. 6 VL muscle fibre pennation angle was observed to increase in response to resistance training. This allowed single muscle fibre CSA and maximal contractile strength to increase more (+16 %) than anatomical muscle CSA and volume (+10 %). 7 Collectively, the present data suggest that the morphology, architecture and contractile capacity of human pennate muscle are interrelated, in vivo. This interaction seems to include the specific adaptation responses evoked by intensive resistance training.

A New Concept For Isokinetic Hamstring: Quadriceps Muscle Strength Ratio

Conventionally, the hamstring:quadriceps strength ratio is calculated by dividing the maximal knee flexor (hamstring) moment by the maximal knee extensor (quadriceps) moment measured at identical angular velocity and contraction mode. The agonist-antagonist strength relationship for knee extension and flexion may, however, be better described by the more functional ratios of eccentric hamstring to concentric quadriceps moments (extension), and concentric hamstring to eccentric quadriceps moments (flexion). We compared functional and conventional isokinetic hamstring: quadriceps strength ratios and examined their relation to knee joint angle and joint angular velocity. Peak and angle-specific (50°, 40°, and 30° of knee flexion) moments were determined during maximal concentric and eccentric muscle contractions (10° to 90° of motion; 30 and 240 deg/sec). Across movement speeds and contraction modes the functional ratios for different moments varied between 0.3 and 1.0 (peak and 50°), 0.4 and 1.1 (40°), and 0.4 and 1.4 (30°). In contrast, conventional hamstring:quadriceps ratios were 0.5 to 0.6 based on peak and 50° moments, 0.6 to 0.7 based on 40° moment, and 0.6 to 0.8 based on 30° moment. The functional hamstring:quadriceps ratio for fast knee extension yielded a 1:1 relationship, which increased with extended knee joint position, indicating a significant capacity of the hamstring muscles to provide dynamic knee joint stability in these conditions. The evaluation of knee joint function by use of isokinetic dynamometry should comprise data on functional and conventional hamstring:quadriceps ratios as well as data on absolute muscle strength.

A mechanism for altered flexibility in human skeletal muscle.

1. We investigated the effect of a long‐term stretching regimen on the tissue properties and stretch tolerance of human skeletal muscle. 2. Resistance to stretch was measured as torque (in N m) offered by the hamstring muscle group during passive knee extension while electromyographic (EMG) activity, knee joint angle and velocity were continuously monitored during a standardized stretch manoeuvre. Seven healthy subjects were tested before and after a 3 week training period using two separate protocols. Protocol 1 consisted of a slow stretch at 0.087 rad s‐1 to a predetermined angle followed by a 90 s holding phase. Subjects were brought to the same angle before and after the training period. Protocol 2 was a similar stretch, but continued to the point of pain. 3. During protocol 1 the torque rose during the stretch and then declined during the holding phase. EMG activity was small and did not change significantly during the protocol. No significant differences in stiffness, energy and peak torque about the knee joint were seen as a result of the training. During protocol 2 the angle to which the knee could be extended was significantly increased as a result of the training. This was accompanied by a comparable increase in peak torque and energy. EMG activity was small and not affected by training. 4. It is concluded that reflex EMG activity does not limit the range of movement during slow stretches and that the increased range of motion achieved from training is a consequence of increased stretch tolerance on the part of the subject rather than a change in the mechanical or viscoelastic properties of the muscle.

Dynamic control of muscle stiffness and H reflex modulation during hopping and jumping in man.

1. The objective of the study was to evaluate the functional effects of reflexes on muscle mechanics during natural voluntary movements. The excitability of the H (Hoffmann) reflex was used as a measure of the excitability of the central component of the stretch reflex. 2. We recorded EMG, ground reaction forces and the H reflex in the soleus muscle in humans while landing from a downward jump, during drop jumping and during hopping. The movements were also recorded by high‐speed cinematography. 3. The EMG pattern was adapted to the motor task. When landing the EMG in the soleus muscle and in the anterior tibial muscle showed preinnervation and alternating activity after touch down. When hopping there was little preinnervation in the soleus muscle, and the activity was initiated about 45 ms after touch down by a peak and continued unbroken until lift off. In the drop jumps the EMG pattern depended on the jumping style used by the subject. 4. The H reflex in the soleus muscle was strongly modulated in a manner appropriate to the requirements of the motor task. During landing from a downward jump the H reflex was low at touch down whereas while hopping it was high at touch down. During drop jumping it was variable and influenced by the jumping technique. 5. Muscle stiffness in the ankle joint was negative after touch down when landing, but always positive when hopping. 6. It is suggested that during landing the alternating EMG pattern after touch down was programmed and little influenced by reflexes. During hopping reflexes could contribute to the initial peak and the EMG during lift off. 7. The programmed EMG activity and the suppression of the H reflex while landing probably contribute to the development of the negative stiffness and change the muscles from a spring to a damping unit.

Antagonist muscle coactivation during isokinetic knee extension

The aim of the present study was to quantify the amount of antagonist coactivation and the resultant moment of force generated by the hamstring muscles during maximal quadriceps contraction in slow isokinetic knee extension. The net joint moment at the knee joint and electromyographic (EMG) signals of the vastus medialis, vastus lateralis, rectus femoris muscles (quadriceps) and the biceps femoris caput longum and semitendinosus muscles (hamstrings) were obtained in 16 male subjects during maximal isokinetic knee joint extension (KinCom, ROM 90–10°, 30°u2003·u2003s−1). Two types of extension were performed: [1] maximal concentric quadriceps contractions and [2] maximal eccentric hamstring contractions. Hamstring antagonist EMG in [1] were converted into antagonist moment based on the EMG‐moment relationships determined in [2] and vice versa. Since antagonist muscle coactivation was present in both [1] and [2] a set of related equations was constructed to yield the moment/EMG relationships for the hamstring and quadriceps muscles, respectively. The equations were solved separately for every 0.05° knee joint angle in the 90–10° range of excursion (0°=full extension) ensuring that the specificity of muscle length and internal muscle lever arms were incorporated into the moment/EMG relationships established. Substantial hamstring coactivation was observed during quadriceps agonist contraction. This resulted in a constant level of antagonist hamstring moment of about 30 Nm throughout the range of motion. In the range of 30–10° from full knee extension this antagonist hamstring moment corresponded to 30–75% of the measured knee extensor moment. The level of antagonist coactivation was 3‐fold higher for the lateral (Bfcl) compared to medial (ST) hamstring muscles. The amount of EMG crosstalk between agonist–antagonist muscle pairs was negligible (RXY2<0.02–0.06). The present data show that substantial antagonist coactivation of the hamstring muscles may be present during slow isokinetic knee extension. In consequence substantial antagonist flexor moments are generated. The antagonist hamstring moments potentially counteract the anterior tibial shear and excessive internal tibial rotation induced by the contractile forces of the quadriceps near full knee extension. In doing so the hamstring coactivation is suggested to assist the mechanical and neurosensory functions of the anterior cruciate ligament (ACL).

Motor unit recruitment during prolonged isometric contractions

SummaryMotor unit recruitment patterns were studied during prolonged isometric contraction using fine wire electrodes. Single motor unit potentials were recorded from the brachial biceps muscle of eight male subjects, during isometric endurance experiments conducted at relative workloads corresponding to 10% and 40% of maximal voluntary contraction (MVC), respectively. The recordings from the 10% MVC experiment demonstrated a characteristic time-dependent recruitment. As the contraction progressed both the mean number of motor unit spikes counted and the mean amplitude of the spikes increased significantly (P<0.01). This progressive increase in spike activity was the result of a discontinuous process with periods of increasing and decreasing activity. The phenomenon in which newly recruited motor units replace previously active units is termed “motor unit rotation” and appeared to be an important characteristic of motor control during a prolonged low level contraction. In contrast to the 10% MVC experiment, there was no indication of de novo recruitment in the 40% MVC experiment. Near the point of exhaustion a marked change in action potential shape and duration dominated the recordings. These findings demonstrate a conspicuous difference in the patterns of motor unit recruitment during a 10% and a 40% MVC sustained contraction. It is suggested that there is a close relationship between intrinsic muscle properties and central nervous system recruitment strategies which is entirely different in fatiguing high and low level isometric contractions.

Determinants of musculoskeletal flexibility: viscoelastic properties, cross-sectional area, EMG and stretch tolerance.

Cross‐sectional area, stiffness, viscoelastic stress relaxation, stretch tolerance and EMG activity of the human hamstring muscle group were examined in endurance‐trained athletes with varying flexibility. Subjects were defined as tight (n=10) or normal (n=8) based on a clinical toe‐touch test. Cross‐sectional area was computed from magnetic resonance imagining (MRI) images. Torque (Nm) offered by the hamstring muscle group, electromyographic (EMG) activity, knee joint angle and velocity were continuously monitored during two standardized stretch protocols. Protocol 1 consisted of a slow stretch at 0.087 rad/s (dynamic phase) to a pre‐determined final angle followed by a 90‐s static phase. In the dynamic phase final angle and stiffness was lower in tight (28.0±2.9 Nm/rad) than normal subjects (54.9±6.5 Nm/rad), P<0.01. In the static phase tight subjects had lower peak (15.4±1.8 Nm) and final torque (10.8±1.6 Nm) than normal subjects (31.6±4.1 Nm, 24.1±3.7 Nm, respectively)(P<0.01), but torque decline was similar. Protocol 2 consisted of a slow stretch to the point of pain and here tight subjects reached a lower maximal angle, torque, stiffness and energy than normal subjects (P<0.01). On the other hand, stiffness was greater in tight subjects in the common range (P<0.01). Cross‐sectional area of the hamstring muscles and EMG activity during the stretch did not differ between the groups. However, lateral hamstring cross‐sectional area was positively related to mid‐range stiffness (P<0.05), but inversely related to final stiffness, peak torque and the toe‐touch test (P<0.01). Final angle and peak torque in protocol 1 combined to improve the predictability of the toe‐touch test (R2=0.77, P<0.001). These data show that the toe‐touch test (R2=0.77, P<0.001). These data show that the toe‐touch test is largely a measure of hamstring flexibility. Further, subjects with a restricted joint range of movement on a clinical toe‐touch test have stiffer hamstring muscles and a lower stretch tolerance.

Viscoelastic stress relaxation during static stretch in human skeletal muscle in the absence of EMG activity

The present study sought to investigate the role of EMG activity during passive static stretch. EMG and passive resistance were measured during static stretching of human skeletal muscle in eight neurologically intact control subjects and six spinal cord‐injured (SCI) subjects with complete motor loss. Resistance to stretch offered by the hamstring muscles during passive knee extension was defined as passive torque (Nm). The knee was passively extended at 5o/s to a predetermined final position, where it remained stationary for 90 s (static phase) while force and integrated EMG of the hamstring muscle were recorded. EMG was sampled for frequency domain analysis in a second stretch maneuver in five control and three SCI subjects. There was a decline in passive torque in the 90‐s static phase for both control and SCI subjects, P<0.05. Although peak passive torque was greater in control subjects, P<0.05, there was no difference in time‐dependent passive torque response between control (33%) and SCI (38%) subjects. Initial and final 5‐s IEMG ranged from 1.8 to 3.4 μ V.s and did not change during a stretch or differ between control and SCI subjects. Frequency domain analysis yielded similar results in both groups, with an equal energy distribution in all harmonics, indicative of ‘white noise’. The present data demonstrate that no measurable EMG activity was detected in either group during the static stretch maneuver. Therefore, the decline in resistance to static stretch was a viscoelastic stress relaxation response.

Activity of mono- and biarticular leg muscles during sprint running

SummaryA cinematographic recording of the movements of the lower limbs together with simultaneous emg tracings from nine lower limb muscles were obtained from two male track sprinters during three phases of a 100 m sprint run. The extensor muscles of the hip joint were found to be the primary movers by acceleration of the bodys center of gravity (C.G.) during the ground phase of the running cycle. The extensors of the knee joint were also important in this, but to a minor extent, while the plantar flexors of the ankle joint showed the least contribution. The biarticular muscles functioned in a way different from the monoarticular muscles in the sense that they perform eccentric work during the flight and recovery phases and concentric work during the whole ground phase (support), whereas the monoarticular muscles are restricted first to eccentric work and then to concentric work during the ground phase. Furthermore, the biarticular muscles show variation (and rate of variation) in muscle length to a larger extent than the monoarticular muscles. Paradoxical muscle actions appear to take place around the knee joint, where the hamstring muscles, m. gastrocnemius, m. vastus laterialis and m. vastus medialis act as synergists by extending the knee joint during the last part of the ground phase.

Anatomy and biomechanics of the back muscles in the lumbar spine with reference to biomechanical modeling.

Study Design. This article describes the development of a musculoskeletal model of the human lumbar spine with focus on back muscles. It includes data from literature in a structured form. Objective. To review the anatomy and biomechanics of the back muscles related to the lumbar spine with relevance for biomechanical modeling. Summary of Background Data. To reduce complexity, muscle units have been incorporated in an abridged manner, reducing their actions more or less to a single force equivalent. In early models of the lumbar spine, this may have been a necessary step to reduce complexity and, thereby, calculation time. The muscles of the spine are well described in the literature, but mainly qualitatively. Most of the literature provides a description of the structures without precise data of fiber length, muscle length, cross-sectional areas, moment arms, forces, etc. The predicted output of musculoskeletal models is very much dependent on the input parameters. The information needed to improve models consists of better approximations of the attachments to the vertebrae, and more precise data. Method. Review of literature. Results. The predicted output of musculoskeletal models is very much dependent on the input parameters. Moderate changes in the assumed muscle line-of-action (i.e., moment arm) could substantially alter the magnitudes of predicted muscle and spinal forces, while the choice of optimization formulation is less sensitive. Conclusions. Input parameters, moment arms, as well as physiologic cross-sectional areas have a profound effect on the predicted muscle forces. Therefore, it is important to choose the values for moment arm and physiologic cross-sectional area carefully because they are essential input parameters to biomechanical models.