Active responses decrease impact forces at the hip and shoulder in falls to the side.

Active responses, such as using the arm to break the fall, may be an effective means of decreasing likelihood of injury in a fall and may help explain why only a small percentage of falls result in a fracture. We quantified the impact force at the hip and shoulder in falls to the side from a kneeling position under three conditions: (1) attempting to break the fall by using an arm; (2) falling with the body relaxed; and (3) falling with the body tensed. Subjects fell from a kneeling position onto a force platform array covered with foam padding and impact force data were recorded. The ground reaction force-time curve was generally bimodal due to sequential impacts of the hip and shoulder. Impact forces at the hip and shoulder were 12 and 16% less for the slap condition (p < 0.05) than for the tensed condition. The impact forces for the relaxed and tensed conditions were not significantly different, although impact forces tended to be less in the relaxed condition. We concluded that active responses reduce the impact forces experienced at the hip and shoulder in falls to the side. Decreased effectiveness of protective responses, due to increases in reaction time and decreases in strength with age, may help explain why so many hip fractures occur in the elderly but so few occur in younger people.     

The axial injury tolerance of the human foot/ankle complex and the effect of Achilles tension

Axial loading of the foot/ankle complex is an important injury mechanism in vehicular trauma that is responsible for severe injuries such as calcaneal and tibial pilon fractures. Axial loading may be applied to the leg externally, by the toepan and/or pedals, as well as internally, by active muscle tension applied through the Achilles tendon during pre-impact bracing. The objectives of this study were to investigate the effect of Achilles tension on fracture mode and to empirically model the axial loading tolerance of the foot/ankle complex. Blunt axial impact tests were performed on forty-three (43) isolated lower extremities with and without experimentally simulated Achilles tension. The primary fracture mode was calcaneal fracture in both groups. However, fracture initiated at the distal tibia more frequently with the addition of Achilles tension (p < 0.05). Acoustic sensors mounted to the bone demonstrated that fracture initiated at the time of peak local axial force. A survival analysis was performed on the injury data set using a Weibull regression model with specimen age, gender, body mass, and peak Achilles tension as predictor variables (R2 = 0.90). A closed-form survivor function was developed to predict the risk of fracture to the foot/ankle complex in terms of axial tibial force. The axial tibial force associated with a 50% risk of injury ranged from 3.7 kN for a 65 year-old 5th percentile female to 8.3 kN for a 45 year-old 50th percentile male, assuming no Achilles tension. The survivor function presented here may be used to estimate the risk of foot/ankle fracture that a blunt axial impact would pose to a human based on the peak tibial axial force measured by an anthropomorphic test device.     

Tonic finite element model of the lower limb

It is widely admitted that muscle bracing influences the result of an impact, facilitating fractures by enhancing load transmission and reducing energy dissipation. However, human numerical models used to identify injury mechanisms involved in car crashes hardly take into account this particular mechanical behavior of muscles. In this context, in this work we aim to develop a numerical model, including muscle architecture and bracing capability, focusing on lower limbs. The three-dimensional (3-D) geometry of the musculoskeletal system was extracted from MRI images, where muscular heads were separated into individual entities. Muscle mechanical behavior is based on a phenomenological approach, and depends on a reduced number of input parameters, i.e., the muscle optimal length and its corresponding maximal force. In terms of geometry, muscles are modeled with 3-D viscoelastic solids, guided in the direction of fibers with a set of contractile springs. Validation was first achieved on an isolated bundle and then by comparing emergency braking forces resulting from both numerical simulations and experimental tests on volunteers. Frontal impact simulation showed that the inclusion of muscle bracing in modeling dynamic impact situations can alter bone stresses to potentially injury-inducing levels.     

Influence of tension time on muscle fiber sarcolemmal injury in rat diaphragm.

We hypothesized that the amount of sarcolemmal injury is directly related to the total tension time (TT(tot)), calculated as mean tension x total stimulation time. Diaphragm strips from Sprague-Dawley rats were superfused at optimal muscle length with Krebs containing procion orange to identify sarcolemmal injury. TT(tot) was induced by stimulation with 100 Hz for 3 min at duty cycles of 0.02, 0.15, 0.3, and 0.6, or with continuous contractions at 0.2, 0.4, 0.6, and 1.0 of maximal tension. A significant positive correlation between TT(tot) and the percentage of fibers with injured sarcolemma (r(2) = 0.63, P < 0.05) is seen. Stimulation (at 100 Hz, duty cycle = 1) resulted in fast fatigue with low injury, likely caused by altered membrane conductivity. Stimulations inducing the largest injury are those showing progressive force loss and high TT(tot), where injury may be due to activation of membrane degradative enzymes. The maximal tension measured at 20 min poststimulation was inversely related to the number of fibers injured, suggesting loss of force is caused by cellular injury.     

Metabolic underpinnings of the paradoxical net phosphocreatine resynthesis in contracting rat gastrocnemius muscle

Net phosphocreatine (PCr) resynthesis during muscle contraction is a paradoxical phenomenon because it occurs under conditions of high energy demand. The metabolic underpinnings of this phenomenon were analyzed non-invasively using 31P-magnetic resonance spectroscopy in rat gastrocnemius muscle (n=11) electrically stimulated (7.6 Hz, 6 min duration) in situ under ischemic and normoxic conditions. During ischemic stimulation, [PCr] initially fell to a steady state (9+/-5% of resting concentration) which was maintained for the last 5 min of stimulation, whereas isometric force production decreased to a non-measurable level beyond 3 min. Throughout normoxic stimulation, [PCr] and force production declined to a steady state after respectively 1 min (5+/-3% of resting concentration) and 3.25 min (21+/-8% of initial value) of stimulation. Contrary to the observations under ischemia, a paradoxical net PCr resynthesis was recorded during the last 2 min of normoxic stimulation and was not accompanied by any improvement in force production. These results demonstrate that the paradoxical net PCr resynthesis recorded in contracting muscle relies exclusively on oxidative energy production and could occur in inactivated fibers, similarly to PCr resynthesis during post-exercise recovery.     

Metabolic underpinnings of the paradoxical net phosphocreatine resynthesis in contracting rat gastrocnemius muscle

Net phosphocreatine (PCr) resynthesis during muscle contraction is a paradoxical phenomenon because it occurs under conditions of high energy demand. The metabolic underpinnings of this phenomenon were analyzed non-invasively using ³¹P-magnetic resonance spectroscopy in rat gastrocnemius muscle (n=11) electrically stimulated (7.6 Hz, 6 min duration) in situ under ischemic and normoxic conditions. During ischemic stimulation, [PCr] initially fell to a steady state (9±5% of resting concentration) which was maintained for the last 5 min of stimulation, whereas isometric force production decreased to a non-measurable level beyond 3 min. Throughout normoxic stimulation, [PCr] and force production declined to a steady state after respectively 1 min (5±3% of resting concentration) and 3.25 min (21±8% of initial value) of stimulation. Contrary to the observations under ischemia, a paradoxical net PCr resynthesis was recorded during the last 2 min of normoxic stimulation and was not accompanied by any improvement in force production. These results demonstrate that the paradoxical net PCr resynthesis recorded in contracting muscle relies exclusively on oxidative energy production and could occur in inactivated fibers, similarly to PCr resynthesis during post-exercise recovery.     

Neuromuscular fatigue during maximal concurrent hand grip and elbow flexion or extension.

The objective was to investigate muscle fatigue measuring changes in force output and force tremor and electromyographic activity (EMG) during two sustained maximal isometric contractions for 60s: (1) concurrent hand grip and elbow flexion (HG and EF); or (2) hand grip and elbow extension (HG and EE). Each force tremor amplitude was decomposed into four frequency bands (1-3, 4-10, 11-20, and 21-50Hz). Surface EMGs were recorded from the flexor digitorum superficialis (FDS), extensor digitorum (ED), biceps brachii (BB) and lateral head of triceps brachii (TB). The HG and EF forces for the HG and EF and the HG force for the HG and EE declined rapidly, whereas the EE force remained almost constant near to the initial value for the first 40s and then declined. The decrease in EMG amplitude was observed not for the FDS muscle but for the ED muscle. The HG tremor amplitude for each frequency band showed similar decreasing rate, whereas the decreases in EF and EE tremor amplitudes for the lower band (below 10Hz) were slower than those for the higher band (above 11Hz). The neuromuscular mechanisms underlying muscle fatigue during sustained maximal concurrent contractions of hand grip and elbow flexion or extension are discussed.     

Anaerobic energy provision in aged skeletal muscle during tetanic stimulation

The effect of age on skeletal muscle anaerobic energy metabolism was investigated in adult (11 mo) and aged (25 mo) Fischer 344 rats. Hindlimb skeletal muscles innervated by the sciatic nerve were stimulated to contract with trains of supramaximal impulses (100 ms, 80 Hz) at a train rate of 1 Hz for 60 s, with an occluded circulation. Soleus, plantaris, and red and white gastrocnemius (WG) were sampled from control and stimulated limbs. All muscle masses were reduced with age (9-13%). Peak isometric tensions, normalized per gram of wet muscle, were lower throughout the stimulation in the aged animals (28%). The potential for anaerobic ATP provision was unaltered with age in all muscles, because resting high-energy phosphates and glycogen contents were similar to adult values. Anaerobic ATP provision during stimulation was unaltered by aging in soleus, plantaris, and red gastrocnemius muscles. In the WG, containing mainly fast glycolytic (FG) fibers, ATP and phosphocreatine contents were depleted less in aged muscle. In situ glycogenolysis and glycolysis were 90.0 +/- 4.8 and 69.3 +/- 2.6 mumol/g dry muscle (dm) in adult WG and reduced to 62.3 +/- 6.9 and 51.5 +/- 5.5 mumol/g dm, respectively, in aged WG. Consequently, total anaerobic ATP provision was lower in aged WG (224.5 +/- 20.9 mumol/g dm) vs. adult (292.6 +/- 7.6 mumol/g dm) WG muscle. In summary, the decreased tetanic tension production in aged animals was associated with a decreased anaerobic energy production in FG fibers. Reduced high-energy phosphate use and a greater energy charge potential after stimulation suggested that the energy demand was reduced in aged FG fibers.(ABSTRACT TRUNCATED AT 250 WORDS)     

Development of biomechanical response corridors of the thorax to blunt ballistic impacts

Human responses are critical to understanding injury biomechanics in blunt ballistic impacts, which are defined as 20-200 g projectiles impacting at 20-250 m/s. 13 human cadavers were exposed to three distinct ballistic impacts of the chest to determine force-time, deflection-time and force-deflection responses. Comparisons were made between biomechanical responses for ballistic impacts and those previously reported for lower speed, higher mass impacts. Impact condition B (140 g at 40 m/s) gave the largest peak force 10,602+/-2226 N and deflection 54.7+/-14.6 mm. Impact condition A (140 g at 20 m/s) involved lower impact energy and produced lower peak force 3383+/-761 N and deflection 25.9+/-3.1 mm, as did impact condition C (40 g at 60 m/s), which gave 3158+/-309 N and 20.1+/-7.8 mm. The results indicate each impact condition gives distinctive responses, which differ from those previously reported in the automotive literature for lower speed impacts. This information provides the foundation for future biomechanical research in the area of blunt ballistic impacts, specifically the development of test surrogates and evaluation of protective equipment.     

The influence of muscle activation on impact dynamics during lateral falls on the hip

Muscle activation has been demonstrated to influence impact dynamics during scenarios including running, automotive impacts, and head impacts. This study investigated the effects of targeted muscle activation magnitude on impact dynamics during low energy falls on the hip with human volunteers. Fifteen university-aged participants (eight females, seven males) underwent 12 lateral pelvis release trials. Half of the trials were muscle-‘relaxed’; in the remaining ‘contracted’ trials participants isometrically contracted their gluteus medius to 20–30% of maximal voluntary contraction before the drop was initiated onto a force plate. Peak force applied to the femur-pelvis complex averaged 9.3% higher in contracted compared to relaxed trials (F = 6.798, p = .022). Muscle activation effects were greater for females, resulting in (on average) an 18.5% increase in effective pelvic stiffness (F = 5.838, p = .046) and a 23.4% decrease in time-to-peak-force (F = 5.109, p = .042). In the relaxed trials, muscle activation naturally increased during the impact event, reaching levels of 12.8, 7.5, 11.1, and 19.1% MVC at the time of peak force for the gluteus medias, vastus lateralis, erector spinae, and external oblique, respectively. These findings demonstrated that contraction of trunk and hip musculature increased peak impact force across sexes. In females, increases in the magnitude and rate of loading were accompanied (and likely driven) by increases in system stiffness. Accordingly, incorporating muscle activation contributions into biomechanical models that investigate loading dynamics in the femur and/or pelvis during lateral impacts may improve estimate accuracy.     

Electromechanical response times and muscle elasticity in men and women

The purpose of this study was to compare the delay in performance attributable to muscle elasticity in men and women. A group of 11 active young men age (mean, SE) 21.9, 0.7 years, stature 1.780, 0.020 m, body mass 76.4, 3.2 kg and 11 women age 20.9, 0.4 years, stature 1.670, 0.020 m and body mass 61.9, 2.6 kg provided written informed consent and were recruited to the study. In response to an acoustic signal delivered via headphones, the subjects performed a plantar flexion movement of the preferred leg as quickly as possible. A seated position ensured that the knee of the subject was flexed at a right angle and that the shank was vertical. The ball of the foot was on a force platform which was used detect the onset of muscle tension and the heel rested on a pressure pad which was used to identify movement. Surface electrodes sensed electromyographic activity (EMG) in the soleus muscle. Force platform output was captured by a digital storage oscilloscope and recorded via a y-t pen recorder or subsequent analysis. A separate timer was used to determine total reaction time (TRT). Premotor time (EMGT) was taken to be the time interval from the delivery of the signal to change in EMG. Electromechanical delay (EMD) was the time interval between the change in EMG and movement and was subdivided into force time (FT) i.e. the time interval between EMG and the onset of muscle tension and elastic charge time (CT) i.e. the time interval between the onset of muscle tension and movement. The subjects performed ten trials and in most cases the mean of ten readings was used to determine TRT, EMGT, EMD, FT and CT.(ABSTRACT TRUNCATED AT 250 WORDS)     

Strain in the braincase and its sutures during function

The skull is distinguished from other parts of the skeleton by its composite construction. The sutures between bony elements provide for interstitial growth of the cranium, but at the same time they alter the transmission of stress and strain through the skull. Strain gages were bonded to the frontal and parietal bones of miniature pigs and across the interfrontal, interparietal and coronal sutures. Strains were recorded 1) during natural mastication in conjunction with electromyographic activity from the jaw muscles and 2) during stimulation of various cranial muscles in anesthetized animals. Vault sutures exhibited vastly higher strains than did the adjoining bones. Further, bone strain primarily reflected torsion of the braincase set up by asymmetrical muscle contraction; the tensile axis alternated between +45 degrees and -45 degrees depending on which diagonal masseter/temporalis pair was most active. However, suture strains were not related to overall torsion but instead were responses to local muscle actions. Only the coronal suture showed significant strain (tension) during jaw opening; this was caused by the contraction of neck muscles. All sutures showed strain during jaw closing, but polarity depended on the pattern of muscle usage. For example, masseter contraction tensed the coronal suture and the anterior part of the interfrontal suture, whereas the temporalis caused compression in these locations. Peak tensile strains were larger than peak compressive strains. Histology suggested that the skull is bent at the sutures, with the ectocranial surface tensed and the endocranial surface predominantly compressed. Collectively, these results indicate that skulls with patent sutures should be analyzed as complexes of independent parts rather than solid structures.     

Fatigue and recovery contractile properties of young and elderly men

The 24 h recovery pattern of contractile properties of the triceps surae muscle, following a period of muscle fatigue, was compared in physically active young (25 years, n = 10) and elderly (66 years, n = 7) men. The fatigue test protocol consisted of 10 min of intermittent submaximal 20 Hz tetani. The maximal twitch (Pt) and tetanic force at 3 frequencies (10, 20 and 50 Hz) were determined at baseline and at 15 min, 1, 4 and 24 h after fatiguing the muscle. Maximal voluntary contraction (MVC) and vertical jump (MVJ) were also assessed. The loss of force during the fatigue test was not significantly different between the young (18 +/- 13%) and elderly (22 +/- 15%). Both groups showed similar and significant reductions of Pt (15%), tetanic force (10 to 35%) and rate of force development (dp/dt) (20%) 15 min and 1 h into recovery. The loss of force was greater at the lower stimulation frequencies of 10 and 20 Hz. Time-to-peak tension was unchanged from baseline during recovery in either group. The average rate of relaxation of twitch force (-dPt/dt) was decreased (p less than 0.05) and half-relaxation time significantly increased at 15 min and 1 h in the elderly but not the young. The findings indicate that after fatiguing contractions, elderly muscle demonstrates a slower return to resting levels of the rate and time course of twitch relaxation compared to the young.     

Theophylline minimally alters contractile properties of canine diaphragm in vitro

We examined the effects of theophylline on contractile properties and high-frequency fatigue of canine diaphragm in vitro. Eighteen diaphragm muscle bundles were obtained from 10 anesthetized dogs and equilibrated in oxygenated Krebs solution to 100, 200, or 300 mg/l theophylline. These bundles were compared with 18 matched control bundles from the contralateral hemidiaphragm. No statistically significant differences in twitch tension, tetanic tension, twitch-to-tetanus ratio, time to peak tension, or half-relaxation time were observed. Concentrations of 300 mg/l theophylline, however, significantly (P less than 0.05) increased force production at 10 Hz by 32%. A similar tendency was present at lower concentrations and exhibited a clear dose-response behavior. High-frequency fatigue was similar in control and theophylline-treated bundles. We conclude that supratherapeutic in vitro concentrations of theophylline do not increase maximal tetanic tension and do not protect against muscle fatigue but potentiate relative force production at low stimulation frequencies. This relatively small effect cannot be explained by poor diffusion of the drug in the muscle bundle, because theophylline concentrations in the muscle bath and in the muscle bundle were virtually identical. Moreover, it remains unclear whether this potentially beneficial effect can be achieved at in vivo attainable serum concentrations.     

Contribution of muscle series elasticity to maximum performance in drop jumping

The contribution of muscle in-series compliance on maximum performance of the muscle tendon complex was investigated using a forward dynamic computer simulation. The model of the human body contains 8 Hill-type muscles of the lower extremities. Muscle activation is optimized as a function of time, so that maximum drop jump height is achieved by the model. It is shown that the muscle series elastic energy stored in the downward phase provides a considerable contribution (32%) to the total muscle energy in the push-off phase. Furthermore, by the return of stored elastic energy all muscle contractile elements can reduce their shortening velocity up to 63% during push-off to develop a higher force due to their force velocity properties. The additional stretch taken up by the muscle series elastic element allows only m. rectus femoris to work closer to its optimal length, due to its force length properties. Therefore the contribution of the series elastic element to muscle performance in maximum height drop jumping is to store and return energy, and at the same time to increase the force producing ability of the contractile elements during push-off.     

Adaptations in human neuromuscular function following prolonged unweighting: I. Skeletal muscle contractile properties and applied ischemia efficacy.

Strength loss following disuse may result from alterations in muscle and/or neurological properties. In this paper, we report our findings on human plantar flexor muscle properties following 4 wk of limb suspension (unilateral lower limb suspension), along with the effect of applied ischemia (Isc) on these properties. In the companion paper (Part II), we report our findings on the changes in neurological properties. Measurements of voluntary and evoked forces, the compound muscle fiber action potential (CMAP), and muscle cross-sectional area (CSA) were collected before and after 4 wk of unilateral lower limb suspension in adults (n = 18; 19-28 yr). A subset of subjects (n = 6) received applications of Isc 3 days/wk (3 sets; 5-min duration). In the subjects not receiving Isc, the loss in CSA and strength was as expected ( approximately 9 and 14%). We observed a 30% slowing in the duration of the CMAP, a 10% decrease in evoked doublet force, a 12% increase in the twitch-to-doublet force ratio, and an altered postactivation potentiation response (11% increase in the postactivation potentiation-to-doublet ratio). We also detected a 10% slowing in the ability of the plantar flexor to develop force during the initial phase of an evoked contraction, along with a 6% reduction in in vivo specific doublet force. In the Isc subjects, no preservation was observed in strength or the evoked muscle properties. However, the Isc group did maintain CSA of the lateral gastrocnemius, as the control subjects' lateral gastrocnemius atrophied 10.2%, whereas the subjects receiving Isc atrophied 4.7%. Additionally, Isc abolished the unweighting-induced slowing in the CMAP. These findings suggest that unweighting alters the contractile properties involved in the excitation-contraction coupling processes and that Isc impacts the sarcolemma.     

Force fluctuations are modulated by alternate muscle activity of knee extensor synergists during low-level sustained contraction.

The study examined the hypothesis that altered synergistic activation of the knee extensors leads to cyclic modulation of the force fluctuations. To test this hypothesis, the force fluctuations were investigated during sustained knee extension at 2.5% of maximal voluntary contraction force for 60 min in 11 men. Surface electromyograms (EMG) were recorded from the rectus femoris (RF), vastus lateralis (VL), and vastus medialis (VM) muscles. The SD of force and average EMG (AEMG) of each muscle were calculated for 30-s periods during alternate muscle activity. Power spectrum of force was calculated for the low- (< or =3 Hz), middle- (4-6 Hz), and high-frequency (8-12 Hz) components. Alternate muscle activity was observed between RF and the set of VL and VM muscles. The SD of force was not constant but variable due to the alternate muscle activity. The SD was significantly greater during high RF activity compared with high VL and VM activity (P < 0.05), and the correlation coefficient between the SD and AEMG was significantly greater in RF [0.736 (SD 0.095), P < 0.05] compared with VL and VM. Large changes were found in the high-frequency component. During high RF activity, the correlation coefficient between the SD and high-frequency component [0.832 (SD 0.087)] was significantly (P < 0.05) greater compared with other frequency components. It is suggested that modulations in knee extension force fluctuations are caused by the unique muscle activity in RF during the alternate muscle activity, which augments the high-frequency component of the fluctuations.     

A joint kinetic analysis of rugby place kicking technique to understand why kickers achieve different performance outcomes

We aimed to identify differences in kicking leg and torso mechanics between groups of rugby place kickers who achieve different performance outcomes, and to understand why these features are associated with varying levels of success. Thirty-three experienced place kickers performed maximum effort place kicks, whilst three-dimensional kinematic (240 Hz) and ground reaction force (960 Hz) data were recorded. Kicking leg and torso mechanics were compared between the more successful (‘long’) kickers and two sub groups of less successful kickers (’short’ and ‘wide-left’) using magnitude-based inferences and statistical parametric mapping. Short kickers achieved substantially slower ball velocities compared with the long kickers (20.8 ± 2.2 m/s vs. 27.6 ± 1.7 m/s, respectively) due to performing substantially less positive hip flexor (normalised mean values = 0.071 vs. 0.092) and knee extensor (0.004 vs. 0.009) joint work throughout the downswing, which may be associated with their more front-on body orientation, and potentially a lack of strength or intent. Wide-left kickers achieved comparable ball velocities (26.9 ± 1.6 m/s) to the long kickers, but they were less accurate due to substantially more longitudinal ball spin and a misdirected linear ball velocity. Wide-left kickers created a tension arc across the torso and therefore greater positive hip flexor joint work (normalised mean = 0.112) throughout the downswing than the long kickers. Whilst this may have assisted kicking foot velocity, it also induced greater longitudinal torso rotation during the downswing, and may have affected the ability of the hip to control the direction of the foot trajectory.     

Characterization of cross-bridge elasticity and kinetics of cross-bridge cycling during force development in single smooth muscle cells

Force development in smooth muscle, as in skeletal muscle, is believed to reflect recruitment of force-generating myosin cross-bridges. However, little is known about the events underlying cross-bridge recruitment as the muscle cell approaches peak isometric force and then enters a period of tension maintenance. In the present studies on single smooth muscle cells isolated from the toad (Bufo marinus) stomach muscularis, active muscle stiffness, calculated from the force response to small sinusoidal length changes (0.5% cell length, 250 Hz), was utilized to estimate the relative number of attached cross-bridges. By comparing stiffness during initial force development to stiffness during force redevelopment immediately after a quick release imposed at peak force, we propose that the instantaneous active stiffness of the cell reflects both a linearly elastic cross-bridge element having 1.5 times the compliance of the cross-bridge in frog skeletal muscle and a series elastic component having an exponential length-force relationship. At the onset of force development, the ratio of stiffness to force was 2.5 times greater than at peak isometric force. These data suggest that, upon activation, cross-bridges attach in at least two states (i.e., low-force-producing and high-force-producing) and redistribute to a steady state distribution at peak isometric force. The possibility that the cross-bridge cycling rate was modulated with time was also investigated by analyzing the time course of tension recovery to small, rapid step length changes (0.5% cell length in 2.5 ms) imposed during initial force development, at peak force, and after 15 s of tension maintenance. The rate of tension recovery slowed continuously throughout force development following activation and slowed further as force was maintained. Our results suggest that the kinetics of force production in smooth muscle may involve a redistribution of cross-bridge populations between two attached states and that the average cycling rate of these cross-bridges becomes slower with time during contraction.