Landing from different heights: Biomechanical and neuromuscular strategies in trained gymnasts and untrained prepubescent girls

The purpose of this study was to examine the biomechanics of the lower limb, during landing in female prepubertal gymnasts and prepubertal untrained girls, aged 9–12 years. Ten healthy participants were included in each group and performed five landings from 20, 40, and 60 cm. Kinematics, ground reaction forces (GRF) and electromyogram (EMG) from the lateral gastrocnemius, tibialis anterior, and vastus lateralis are presented. Gymnasts had higher vertical GRF and shorter braking phase during landing. Compared to untrained girls, gymnasts exhibited for all examined drop heights more knee flexion before and at ground contact, but less knee flexion at maximum knee flexion position. Especially when increasing drop heights the gymnasts activated their examined muscles earlier, and generally they had higher pre- and post landing EMG amplitudes normalized to the peak EMG at 60 cm drop height. Furthermore, gymnasts had lower antagonist EMG for the tibialis anterior compared to untrained girls, especially when landing from higher heights. It is concluded that the landing strategy preferred by gymnasts is influenced by long-term and specialized training and induces a stiffer landing pattern. This could have implications in injury prevention, which requires further investigation.     

Modification of landing conditions at contact via flight

Weight-bearing tasks performed by humans consist of a series of phases with multiple objectives. Analysis of the relationship between control and dynamics during successive phases of the tasks is essential for improving performance without sustaining injury. Experimental evidence regarding foot landings suggests that the distribution of momentum among segments at contact influences stability during interaction with the landing surface. In this study, we hypothesized that modification of control in one subsystem, in our case shoulder torque, during the flight phase of an aerial task would enable the performer to maintain behavior of other subsystems (e.g.lower extremity kinematics) and initiate contact with momentum conditions consistent with successful task performance. To test this hypothesis, an experimentally validated multilink dynamic model that incorporated modifications in shoulder torque was used to simulate the flight phase dynamics of overrotated landings. The simulation results indicate that modification in shoulder torque during the flight phase enables gymnasts to maintain lower extremity kinematics and initiate contact with trunk angular velocities consistent with those observed during successful landings. These results suggest that modifications in the control logic of one subsystem may be sufficient for achieving both global and local task objectives of landing.     

JUMP LANDING CHARACTERISTICS IN ELITE SOCCER PLAYERS WITH CEREBRAL PALSY

The aim of the present study was to analyse the parameters that characterize the vertical ground reaction force during the landing phase of a jump, and to determine the relationship among these parameters in elite soccer players with cerebral palsy (CP). Thirteen male members of the Spanish national soccer team for people with CP (mean age: 27.1 ± 4.7 years) volunteered for the study. Each participant performed three counter movement jumps. The characteristics of the first peak of the vertical ground reaction force during the landing phase of a jump, which corresponds to the forefoot contact with the ground, were similar to the results obtained in previous studies. However, a higher magnitude of rearfoot contact with the ground (F2) was observed in participants with CP than in participants without CP. Furthermore, a significant correlation between F2 magnitude and the elapsed time until its production (T2) was not observed (r = -0.474 for p = 0.102). This result implies that a landing technique based on a delay in the production of F2 might not be effective to reduce its magnitude, contrary to what has been observed in participants without CP. The absence of a significant correlation between these two parameters in the present study, and the high magnitude of F2, suggest that elite soccer players with CP should use footwear with proper cushioning characteristics.     

A six degree of freedom head acceleration measurement device for use in football

The high incidence rate of concussions in football provides a unique opportunity to collect biomechanical data to characterize mild traumatic brain injury. The goal of this study was to validate a six degree of freedom (6DOF) measurement device with 12 single-axis accelerometers that uses a novel algorithm to compute linear and angular head accelerations for each axis of the head. The 6DOF device can be integrated into existing football helmets and is capable of wireless data transmission. A football helmet equipped with the 6DOF device was fitted to a Hybrid III head instrumented with a 9 accelerometer array. The helmet was impacted using a pneumatic linear impactor. Hybrid III head accelerations were compared with that of the 6DOF device. For all impacts, peak Hybrid III head accelerations ranged from 24 g to 176 g and 1,506 rad/s(2) to 14,431 rad/s(2). Average errors for peak linear and angular head acceleration were 1% ± 18% and 3% ± 24%, respectively. The average RMS error of the temporal response for each impact was 12.5 g and 907 rad/s(2).     

Effects of added trunk load and corresponding trunk position adaptations on lower extremity biomechanics during drop-landings

Although both trunk mass and trunk position have the potential to affect lower extremity biomechanics during landing, these effects are not well understood. Our overall hypothesis stated that both trunk mass and trunk position affect lower extremity biomechanics in landing. Thus, our purpose was to determine the effects of an added trunk load and kinematic trunk adaptation groups on lower extremity joint kinematics, kinetics, and energetics during drop-landings. Twenty-one recreationally active subjects were instrumented for biomechanical analysis. Subjects performed two sets of eight double-limb landings with and without 10% body weight added to the trunk. On lower extremity dependent variables, 2(condition: no load, trunk load)x2(group: trunk extensors vs. trunk flexors) ANOVAs were performed. Condition by group interactions at the hip showed differing responses to the added trunk load between groups where the trunk extensor group decreased hip extensor efforts ( downward decrease 11-18%) while the trunk flexor group increased hip extensor efforts ( upward increase 14-19%). The trunk load increased biomechanical demands at the knee and ankle regardless of trunk adaptation group. However, the percent increases in angular impulses and energy absorption in the trunk extensor group were 14-28% while increases in the trunk flexor group were 4-9%. Given the 10% body weight added to the trunk, the 14-28% increases at the knee and ankle in the trunk extensor group were likely due to the reduced hip extensor efforts during landing. Overall these findings support our overall hypothesis that both trunk mass and trunk position affect lower extremity biomechanics during vertically oriented landing tasks.     

How honeybees make grazing landings on flat surfaces

 Freely flying bees were filmed as they landed on a flat, horizontal surface, to investigate the underlying visuomotor control strategies. The results reveal that (1) landing bees approach the surface at a relatively shallow descent angle; (2) they tend to hold the angular velocity of the image of the surface constant as they approach it; and (3) the instantaneous speed of descent is proportional to the instantaneous forward speed. These characteristics reflect a surprisingly simple and effective strategy for achieving a smooth landing, by which the forward and descent speeds are automatically reduced as the surface is approached and are both close to zero at touchdown. No explicit knowledge of flight speed or height above the ground is necessary. A model of the control scheme is developed and its predictions are verified. It is also shown that, during landing, the bee decelerates continuously and in such a way as to keep the projected time to touchdown constant as the surface is approached. The feasibility of this landing strategy is demonstrated by implementation in a robotic gantry equipped with vision. Received: 21 June 1999 / Accepted in revised form: 20 March 2000     

Innate Flower Recognition in Bumblebees (Bombus terrestris,B. lucorum; Apidae): Optical Signals from Stamens as Landing Reaction Releasers

Innate behavioural reactions in bumblebees were observed in flower dummy experiments. In a previous paper, it was shown that the approach of flower-naive, untrained bumblebees toward flower dummies is released by colour signals, i.e. a gradient of spectral purity. The bumblebees are directed toward that part of a flower dummy which exhibits the highest spectral purity (Lunau 1990). In the present work it is shown that the bumblebees, while still in flight, make contact with this part using their antennal tips. The antennae reaction is followed by a landing reaction which is released by olfactory and optical signals. The specific optical signals of the guide area which release the landing reaction were investigated. Since in the flower dummies which were used, the guide had the highest spectral purity, most of the antennae reactions were executed at the guide. The antennae reaction allows for a precise orientation and may involve the stimulation of a specialized area of ommatidia. Minute changes in optical guide parameters could thus be detected. The following features of stamens which most effectively released the landing reaction were: a minimum of 2 theca dummies; an oval shape of theca dummies which face each other along the longitudinal axis (both these characteristics are found in genuine stamens); 4 mm distance between theca dummies (this coincides with the distance between the antennal tips of bumblebees); two equally sized theca dummies each with an area of 7 mm² (this size of theca dummies was preferred over larger or smaller ones; the latter parameter was not changed when the corolla size was varied). Colour parameters are not involved in releasing the landing reaction. Clearly, bumblebees possess an innate releasing mechanism (IRM) that responds to optical signals of anthers which release the landing reaction. The experimental results are in accordance with conclusions drawn from the comparative studies of Osche (1979, 1983a, 1983b) that many flower guides can be interpreted as stamen mimics. Therefore stamens as well as stamen mimics represent a highly standardized signal for the insect pollinators and they are well suited for innate orientation at flowers.     

In vivo measurement of age-related stiffening in the crystalline lens by Brillouin optical microscopy

The biophysical and biomechanical properties of the crystalline lens (e.g., viscoelasticity) have long been implicated in accommodation and vision problems, such as presbyopia and cataracts. However, it has been difficult to measure such parameters noninvasively. Here, we used in vivo Brillouin optical microscopy to characterize material acoustic properties at GHz frequency and measure the longitudinal elastic moduli of lenses. We obtained three-dimensional elasticity maps of the lenses in live mice, which showed biomechanical heterogeneity in the cortex and nucleus of the lens with high spatial resolution. An in vivo longitudinal study of mice over a period of 2 months revealed a marked age-related stiffening of the lens nucleus. We found remarkably good correlation (log-log linear) between the Brillouin elastic modulus and the Young's modulus measured by conventional mechanical techniques at low frequencies (~1 Hz). Our results suggest that Brillouin microscopy is potentially useful for basic and animal research and clinical ophthalmology.     

Comparative efficiency of different regimens of locomotor training in prolonged space flights as estimated from the data on biomechanical and electromyographic parameters of walking

Biomechanical and electromyographic characteristics of locomotion were studied before and after a space flight on days 3, 7, and 10 after landing in 18 participants of prolonged space missions on board the International Space Station. It has been shown that microgravity causes significant changes in biomechanical and electromyographic characteristics of walking, such as a decrease in the amplitude of angular displacement in leg joints, a decrease in the double step length, and an increase in the electromyographic costs of locomotion. It has been also shown that interval locomotor physical training, such as alternation of running and walking, in prolonged space flights prevents an increase in the physiological costs of locomototions after a space flight and provides more efficient maintenance of the neuromuscular system’s performance after a flight. Cosmonauts who performed interval locomotor training had fewer changes in biomechanical and electromyographic characteristics of walking.     

Aerial maneuvers of leaping lemurs: The physics of whole-body rotations while airborne

Several prosimian species begin a leap from a vertical support with their back toward the landing target. To reorient themselves from this dorsally facing, head-first lift-off to a ventrally facing, feet-first landing, the animals combine an initial twist with a partial backward somersault. Cinefilms of a captive colony of ringtailed lemurs (Lemur catta) revealed that during leaps from vertical poles to horizontal supports, the backward somersaulting rotations were often initiated while the animals were airborne. How could these prosimians initiate rotations in the absence of externally applied forces without violating angular momentum conservation? The problem was approached through vector analysis to demonstrate angular momentum (H) changes about the three principal (symmetrical) axes of rotation for a series of critical body positions that were extracted from the filmed sequences. One L. catta specimen was segmented to provide the dimensions and weights necessary for modeling the various body positions. These data were also used to calculate moments of inertia about the three principal axes in order to predict if rotations about these axes were stable or metastable. Lemurs, like any projectile, must conserve the total angular momentum (H_T) established at lift-off. H_T, however, is a vector quantity that is the resultant of component vectors about the three principal axes. Thus, H about the individual axes may change as long as H_T remains constant. Strategically timed tail movements tilted the body, thereby changing the H value about the head-to-toe (twisting) axis. To conserve H_T, also aligned along the twisting axis, angular momentum transferred to the somersaulting axis. Owing to the direction of tail-throw, the initiated rotations were partial backward somersaults that brought the hindlimbs forward for landing. This strategy for initiating specific rotations parallels that practiced by human springboard divers.     

The effect of falling height on muscle activity and foot motion during landings

The aims of this study were: (a) to examine the effect of falling height on the kinematics of the tibiotalar, talonavicular and calcaneocuboid joints and (b) to study the influence of falling height on the muscle activity of the leg during landings. Six female gymnasts (height: 1.63±0.04 m, weight: 58.21±3.46 kg) participated in this study. All six gymnasts carried out barefoot landings, falling from 1.0, 1.5 and 2.0 m height onto a mat. Three genlocked digital high speed video cameras (250 Hz) captured the motion of the left shank and foot. Surface electromyography (EMG) was used to measure muscle activity (1000 Hz) from five muscles (gastrocnemius medialis, tibialis anterior, peroneus longus, vastus lateralis and hamstrings) of the left leg. The kinematics of the tibiotalar, talonavicular and calcaneocuboid joints were studied. The lower-leg and the foot were modelled by means of a multi-body system, comprising seven rigid bodies. The falling height does not show any influence on the kinematics neither of the tibiotalar nor of the talonavicular joints during landing. The eversion at the calcaneocuboid joint increases with increasing falling height. When augmenting falling height, the myoelectric activity of the muscles of the lower limb increases as well during the pre-activation phase as during the landing itself. The muscles of the lower extremities are capable of stabilizing the tibiotalar and the talonavicular joints actively, restricting their maximal motion by means of a higher activation before and after touchdown. Maximal eversion at the calcaneocuboid joint increases about 52% when landing from 2.0 m.     

Spatial Scaling of Environmental Variables Improves Species-Habitat Models of Fishes in a Small,Sand-Bed Lowland River

Habitat suitability and the distinct mobility of species depict fundamental keys for explaining and understanding the distribution of river fishes. In recent years, comprehensive data on river hydromorphology has been mapped at spatial scales down to 100 m, potentially serving high resolution species-habitat models, e.g., for fish. However, the relative importance of specific hydromorphological and in-stream habitat variables and their spatial scales of influence is poorly understood. Applying boosted regression trees, we developed species-habitat models for 13 fish species in a sand-bed lowland river based on river morphological and in-stream habitat data. First, we calculated mean values for the predictor variables in five distance classes (from the sampling site up to 4000 m up- and downstream) to identify the spatial scale that best predicts the presence of fish species. Second, we compared the suitability of measured variables and assessment scores related to natural reference conditions. Third, we identified variables which best explained the presence of fish species. The mean model quality (AUC = 0.78, area under the receiver operating characteristic curve) significantly increased when information on the habitat conditions up- and downstream of a sampling site (maximum AUC at 2500 m distance class, +0.049) and topological variables (e.g., stream order) were included (AUC = +0.014). Both measured and assessed variables were similarly well suited to predict species’ presence. Stream order variables and measured cross section features (e.g., width, depth, velocity) were best-suited predictors. In addition, measured channel-bed characteristics (e.g., substrate types) and assessed longitudinal channel features (e.g., naturalness of river planform) were also good predictors. These findings demonstrate (i) the applicability of high resolution river morphological and instream-habitat data (measured and assessed variables) to predict fish presence, (ii) the importance of considering habitat at spatial scales larger than the sampling site, and (iii) that the importance of (river morphological) habitat characteristics differs depending on the spatial scale.     

A virtual environment for learning to view during aerial movements

Training the gymnasts to view the landing area when learning aerial skills may lead to more consistent landings but can be problematic and potentially dangerous. A virtual environment allowing gymnasts to get introduced to viewing techniques safely is presented. The system is based on existing simulation models and visualisation software, and is implemented using client–server technology to allow reuse with new simulation models in the future.     

Basic organization of operant behavior as revealed in Drosophila flight orientation

Summary Operant behavior is studied in tethered Drosophila flies using visual motion, heat or odour as operandum and yaw torque, thrust or direction of flight as operans in various combinations (Fig. 1). On the basis of these results a conceptual framework of operant behavior is proposed: (1) It requires a goal (desired state) of which the actual state deviates. (2) To attain the goal a range of motor programs is activated (initiating activity, see Fig. 7). (3) Efference copies of the motor programs are compared to the sensory input referring to the deviation from the desired state (e.g. by cross-correlation). (4) In case of a significant coincidence the respective motor program is used to modify the sensory input in the direction towards the goal. (5) Consistent control of a sensory stimulus by a behavior may lead to a more permanent behavioral change (conditioning). In this scheme operant activity (1–4) and operant conditioning (1–5) are distinguished.Abbreviations ALU arbitrary length unit - d horizontal angular width of visual pattern - IR infrared - SEM standard error of the means - T yaw torque - Th thrust - performance index - horizontal angle between visual pattern position and longitudinal body axis of the fly - vertical angular extension of visual pattern     

The influence of simulation model complexity on the estimation of internal loading in gymnastics landings

Evaluating landing technique using a computer simulation model of a gymnast and landing mat could be a useful tool when attempting to assess injury risk. The aims of this study were: (1) to investigate whether a subject-specific torque-driven or a subject-specific muscle-driven model of a gymnast is better at matching experimental ground reaction forces and kinematics during gymnastics landings, (2) to calculate their respective simulation run times and (3) to determine what level of model complexity is required to assess injury risk. A subject-specific planar seven-link wobbling mass model of a gymnast and a multi-layer model of a landing mat were developed for this study. Subject-specific strength parameters were determined which defined the maximum voluntary torque/angle/angular velocity relationship about each joint. This relationship was also used to produce subject-specific 'lumped' muscle models for each joint. Kinetic and kinematic data were obtained during landings from backward and forward rotating gymnastics vaults. Both torque-driven and muscle-driven models were capable of producing simulated landings that matched the actual performances (with overall percentage differences between 10.1% and 18.2%). The torque-driven model underestimated the internal loading on joints and bones, resulting in joint reaction forces that were less than 50% of those calculated using the muscle-driven model. Simulation time increased from approximately 3 min (torque driven) to more than 10 min (muscle driven) as model complexity increased. The selection of a simulation model for assessing injury risk must consider the need for determining realistic internal forces as the priority despite increases in simulation run time.     

Foot structure and knee joint kinetics during walking with and without wedged footwear insoles

The relationship between static foot structure characteristics and knee joint biomechanics during walking, or the biomechanical response to wedged insoles are currently unknown. In this study, 3D foot scanning, dual X-ray absorptiometry and gait analysis methods were used to determine structural parameters of the foot and assess their relation to knee joint loading and biomechanical response to wedged insoles in 30 patients with knee osteoarthritis. In multiple linear regression models, foot fat content, height of the medial longitudinal arch and static hind foot angle were not associated with the magnitude of the knee adduction moment (R² = 0.24, p = 0.060), knee adduction angular impulse (R² = 0.21, p = 0.099) or 3D resultant knee moment (R² = 0.23, p = 0.073) during gait. Furthermore, these foot structure parameters were not associated with the patients’ biomechanical response to medial or lateral wedge footwear insoles (all p < 0.01). These findings suggest that static foot structure is not associated with gait mechanics at the knee, and that static foot structure alone cannot be utilized to predict an individual’s biomechanical response to wedged footwear insoles in patients with knee osteoarthritis.     

Stem biomechanics of three columnar cacti from the Sonoran Desert

The allometric relationship of stem length L with respect to mean stem diameter D was determined for 80 shoots of each of three columnar cactus species (Stenocereus thurberi, Lophocereus schottii, and S. gummosus) to determine whether this relationship accords with that predicted by each of three contending models purporting to describe the mechanical architecture of vertical shoots (i.e., geometric, stress, and elastic similitude, which predict L proportional to D(alpha), with alpha = 1/1, 1/2, and 2/3, respectively). In addition, anatomical, physical, and biomechanical stem properties were measured to determine how the stems of these three species maintain their elastic stability as they increase in size. Reduced major axis regression of L with respect to D showed that alpha = 2.82 ± 0.14 for S. thurberi, 2.32 ± 0.19 for L. schottii, and 4.21 ± 0.31 for S. gummosus. Thus, the scaling exponents for the allometry of L differed significantly from that predicted by each of the three biomechanical models. In contrast, these exponents were similar to that for the allometry previously reported for saguaro. Analyses of biomechanical data derived from bending tests performed on 30 stems selected from each of the three species indicated that the bulk stem tissue stiffness was roughly proportional to L2, while stem flexural rigidity (i.e., the ability to resist a bending force) scaled roughly as L3. Stem length was significantly and positively correlated with the volume fraction of wood, while regression analysis of the pooled data from the three species (i.e., 90 stems) indicated that bulk tissue stiffness scaled roughly as the 5/3-power of the volume fraction of wood in stems. These data were interpreted to indicate that wood served as the major stiffening agent in stems and that this tissue accumulates at a sufficient rate to afford unusually high scaling exponents tot stem length with respect to stem diameter (i.e., disproportionately large increments of stem length with respect to increments in stem diameter). Nevertheless, the safety factor against the elastic failure of stems (computed on the basis of the critical buckling height divided by actual stem length) decreased with increasing stem size tot each species, even though each species maintained an average safety factor equal to two. We speculate that the apparent upper limit to plant height calculated for each species may serve as a biomechanical mechanism for vegetative propagation and the establishment of dense plant colonies by means of extreme stem flexure and ultimate breakage, especially for S. gummosus.     

Heavy-ion anisotropy measured by ALTEA in the International Space Station

The uneven shielding of the International Space Station from the vessel hull, racks and experiments produces a modulation of the internal radiation environment. A detailed knowledge of this environment, and therefore of the Station's shielding effectiveness, is mandatory for an accurate assessment of radiation risk. We present here the first 3D measurements of the Station's radiation environment, discriminating particle trajectories and LET, made possible using the detection capability of the ALTEA-space detector. We provide evidence for a strong (factor ≈ 3) anisotropy in the inner integral LET for high-LET particles (LET > 50?keV/μm) showing a minimum along the longitudinal station axis (most shielded) and a maximum normal to it. Integrating over all measured LETs, the anisotropy is strongly reduced, showing that unstopped light ions plus the fragments produced by heavier ions approximately maintain flux/LET isotropy. This suggests that, while changing the quality of radiation, the extra shielding along the station main axis is not producing a benefit in terms of total LET. These features should be taken into account (1) when measuring radiation with detectors that cannot distinguish the direction of the impinging radiation or that are unidirectional, (2) when planning radiation biology experiments on the ISS, and (3) when simulating the space radiation environment for experiments on the ground. A novel analysis technique that fully exploits the ability to retrieve the angular distribution of the radiation is also presented as well as the angular particle flux and LET characteristic of three geomagnetic zones measured during 2009 by the ALTEA-space detector. This technique is applied to the ALTEA-space detector, but a wider applicability to other detectors is suggested.     

Mechanical demand and multijoint control during landing depend on orientation of the body segments relative to the reaction force.

The purpose of this study was to determine how diverse momentum conditions and anatomical orientation at contact influences mechanical loading and multijoint control of the reaction force during landings. Male collegiate gymnasts (n=6) performed competition style landings (n=3) of drop jumps, front saltos, and back saltos from a platform (0.72 m) onto landing mats (0.12 m). Kinematics (200 fps), reaction forces (800 Hz) and muscle activation patterns (surface EMG, 1600 Hz) of seven lower extremity muscles were collected simultaneously. Between-task differences in segment orientation relative to the reaction force contributed to significant between-task differences in knee and hip net joint moments (NJM) during the impact phase. During the stabilization phase, ankle, knee, and hip NJMs acted to control joint flexion. Between-task differences in muscle activation patterns indicated that gymnasts scaled biarticular muscle activation to accommodate for between-task differences in NJM after contact. Activation of muscles on both sides of the joint suggests that impedance like control was used to stabilize the joints and satisfy the mechanical demand imposed on the lower extremity. Between-subject differences in the set of muscles used to control total body center of mass (TBCM) trajectory and achieve lower extremity NJMs suggests that control of multijoint movements involving impact needs to incorporate mechanical objectives at both the total body and local level. The functional consequences of such a control structure may prove to be an asset to gymnasts, particularly when required to perform a variety of landing tasks under a variety of environmental constraints.     

Orientation of spin-labeled light chain 2 of myosin heads in muscle fibers

Electron paramagnetic resonance (e.p.r.) spectroscopy has been used to monitor the orientation of spin labels attached rigidly to a reactive SH residue on the light chain 2 (LC2) of myosin heads in muscle fibers. e.p.r. spectra from spin-labeled myosin subfragment-1 (S1), allowed to diffuse into unlabeled rigor (ATP-free) fibers, were roughly approximated by a narrow angular distribution of spin labels centered at 66 degrees relative to the fiber axis, indicating a uniform orientation of S1 bound to actin. On the other hand, spectra from spin-labeled heavy meromyosin (HMM) were roughly approximated by two narrow angular distributions centered at 42 degrees and 66 degrees, suggesting that the LC2 domains of the two HMM heads have different orientations. In contrast to S1 or HMM, the spectra from rigor fibers, in which LC2 of endogenous myosin heads was labeled, showed a random orientation which may be due to distortion imposed by the structure of the filament lattice and the mismatch of the helical periodicities of the thick and thin filaments. However, spectra from the fibers in the presence of ATP analog 5'-adenylyl imidodiphosphate (AMPPNP) were approximated by two narrow angular distributions similar to those obtained with HMM. Thus, AMPPNP may cause the LC2 domain to be less flexible and/or the S2 portion to be more flexible, so as to release the distortion of the LC2 domain and make it return to its natural position. At high ionic strength, AMPPNP disoriented the spin labels as ATP did under relaxing conditions, suggesting that the myosin head is detached from and/or weakly (flexibly) attached to a thin filament.