JOURNALOF NEUROPHYSIOLOGY Vol. 7 1, No. 4, April 1994. Printed in U.S.A. A Control Strategy for the Execution of Explosive Movements From Varying Starting Positions ARTHUR J. VAN SOEST, MAARTEN F. BOBBERT, AND GERRIT JAN VAN INGEN SCHENAU Faculty of Human MovementSciences,Vrije Universiteit,NL 1081 BT Amsterdam, The Netherlands exerted on the ground be properly controlled (van Ingen Schenau et al. 1992; Jacobs and van Ingen Schenau 1992). 1. Humans can execute explosive movements such asjumping Obviously, successful execution of extremely fast moveand hitting an object irrespective of the starting position from skeletal system subwhich these movements have to be initiated; in fact, variability of ments of an inverted pendulum-like kinematic parameters has been shown to decreasein the course of jected to strong dynamic coupling between segments poses the movement. extreme demands on the control system. Thus it is rather 2. We address the question of whether it is necessaryto adapt surprising that quite successful performance of these tasks the stimulation pattern of the muscles to such variations in start- is seen in virtually all humans. In expert performers kineing position or whether the stabilizing effect of intrinsic muscle matic patterns have been shown to be highly consistent for properties is such that one single muscle stimulation pattern might tasks such as vertical jumping (Bobbert and van Ingen be used for a wide range of starting positions. Schenau 1988), the forehand drive in table tennis 3. Specifically, we address this question for maximum-height human vertical squat jumping, using an approach based on mathe- (Bootsma and van Wieringen 1990), and cricket batting matical modeling and computer simulation. The stimulation pat- ( McCleod 1987). Perhaps the most surprising finding is tern of the muscles is the input of the model and the resulting that execution does not seem to be hampered when the movement has to be started from widely different starting movement is the output. 4. The optimal stimulation pattern for a starting position in the positions. For example, the adequacy of performance of middle of the range of starting positions considered does not lead squat jumps (i.e., maximally high vertical jumps starting to adequate performance for other starting positions in that range. from a static squatted position) does not depend to any 5. However, a muscle stimulation pattern can be found that degree on the initial position (M. F. Bobbert, K. G. M. does result in close to optimal achievement for a wide range of Gerritsen, M. C. A. Litjens, and A. J. van Soest, unpubstarting positions. This muscle stimulation pattern, which is not optimal for any specific starting position, may be considered as lished data). A similar conclusion was drawn by Bootsma and van Wieringen ( 1990) in their study of the table tennis “control that works” as opposed to “optimal control.” 6. The latter muscle stimulation pattern also leads to adequate forehand drive. These findings imply that variability of imbehavior for “new” starting positions both within and outside the portant movement parameters decreases as the instant of time on which achievement depends (i.e., takeoff in jumprange considered. ing; ball contact in table tennis) is approached. The combination of high task demands and observed sucINTRODUCTION cessful performance leads to the general question of which control strategy is employed by performers of such tasks. A In animals, skillful execution of explosive movements such as jumping, sprint running, throwing, kicking, or hit- communis opinio exists with respect to the fact that the ting an object is of paramount importance both in fleeing control of explosive movements is to a certain extent preprogrammed (Schmidt 1982). However, the opinions conbehavior and in prey catching. In humans these movements cerning two more specific questions are less unequivocal. are prominent primarily in sport situations. Roughly stated, the purpose of these tasks is to maximize the final The first of these concerns the way in which disturbances that take place during the unfolding of the movement are velocity of the end point of a skeletal chain. Joint angular velocities reach values as high as 1,000” /s in human verti- counteracted. The second question is how the observed cal jumping (Bobbert and van Ingen Schenau 1988)) which large variations in starting position are dealt with. Adaptation of the motor program on the basis of propriois one of the slower explosive movements. The strong dyceptive feedback is the first mechanism that springs to mind namic coupling between the segments that results from in relation to the first question: it is imaginable that the such velocities complicates the control of these movements muscle stimulation pattern is significantly (Hollerbach and Flash 1982). A related char- default preprogrammed adapted when information on a disturbance of the moveacteristic is the short duration of the movement; movement times for these tasks in humans vary from -40 ms for Mu- ment becomes available on the basis of proprioceptive feedhammed Ali’s left jab (Schmidt 1982) to -300 ms for the back.. However, it is well established that in the lower expushoff phase in vertical jumping (Bobbert and van Ingen tremity the fastest neural feedback loop, i.e., the myotatic Schenau 1988). Finally, the skeletal system is mechanically reflex, involves a latency of -45 ms between stimulus and analogous to an inverted pendulum in cases where the end first change in electromyogram (EMG) and that the gain of point moves against gravity. Vertical jumping is a clear ex- this reflex at high velocities is low (Gottlieb and Agarwal 1979). In the upper extremity this latency is somewhat ample in this respect. Preventing disintegration of the movement in such cases requires that the direction of the force shorter, i.e., -30 ms (Gielen and Houk 1984). FurtherSUMMARY 1390 AND CONCLUSIONS 0022-3077/94 $3.00 Copyright 0 1994 The American Physiological Society CONTROL OF EXPLOSIVE more, although a certain degree of interjoint coordination exists with respect to this reflex (Lacquaniti and Soechting 1986; Smeets and Erkelens 199 1), it is unlikely that these effects can be modulated to such an extent, that the timevarying dynamic coupling between segments can be counteracted adequately. In contrast, the “functional stretch reflex” (Melville Jones and Watt 197 1) can be modulated to a large extent; however, this reflex is characterized by a much longer latency, typically - 120 ms. Also, it must be considered that muscle force lags behind EMG by -5O100 ms (Vos et al. 1990); these forces determine acceleration, which has to be integrated over time once or twice to obtain changes in velocity/position, which again takes some time. Because all these processes occur serially all latencies involved have to be added, leading to a total latency of - lOO- 150 ms at the minimum for a feedback loop that can be modulated only to a limited extent. Comparing these latencies to the movement times involved it is inevitable to conclude that adaptation of the stimulation pattern of muscles on the basis of neural feedback cannot be effective in counteracting disturbances occurring during explosive movements. A second mechanism that has been suggested to reduce the effects of disturbances occurring during movement concerns the stabilizing effect of muscle properties. The contribution of the force-length-velocity relation of muscle to the counteracting of externally imposed disturbances has been known for quite some time (e.g., Grillner 1972). In fact, the intrinsic stiffness resulting from the force-length relationship of muscles below optimum length is at the heart of the equilibrium point theory of Bizzi et al. (e.g., Bizzi et al. 1992). Although this intrinsic stiffness is smaller than that resulting from the myotatic reflex (Carter et al. 1990; Nichols and Houk 1976)) it has the advantage of zero time delay (Hogan 1990; McMahon 1984) : an adaptation of muscle force occurs instantaneously when muscle length and/ or velocity are disturbed. This zero time delay might be an especially important property in fast movements, because in these movements the role of neural feedback is limited. Recently we investigated the potential contribution of the force-length-velocity relation of muscle in counteracting disturbances occurring during movement in human squat jumping (van Soest and Bobbert 1993). Using a modeling and simulation approach it was shown that these muscle properties in fact reduce the effect of disturbances to such a degree that an adaptation of the muscle stimulation pattern is no longer needed. Because of muscle properties the control system is relieved of the burden of adapting the muscle stimulation pattern in case of disturbances. Having identified the mechanism most likely to be responsible for handling of disturbances occurring during movement, we now turn to the question of how the observed large variations in starting position are dealt with. Obviously these variations may be considered as disturbances of a “standard” starting position. As such, this question may be considered as a special case of the question discussed above. However, explosive movements are typically started from a static position that was maintained for quite some time. Consequently, neural feedback of information concerning this position is definitely possible. Thus the question of how variations in initial position are han- MOVEMENTS 1391 dled is usually replaced by the more specific question of how the control system succeeds in generating an adequate muscle stimulation pattern on the basis of “knowledge” of the initial position. Several propositions have been made with respect to this more specific question. On the one hand, it has been suggested that an adequate “specific motor program” is either retrieved from storage directly or created by setting a number of parameters in a stored “general motor program” (Schmidt 1982). On the other hand, it has been suggested (Cooke and Diggles 1984; van der Meulen et al. 1990) that adequate control signals are generated “on-line” on the basis of an efference copy (von Holst and Mittelstaedt 1950). A characteristic shared by all these suggestions is that they place heavy demands on the capabilities of the control system. In the present study we attempt to formulate a less involved control strategy for handling large variations in starting position. The alternative strategy proposed is based on the finding that muscle properties reduce the effect of disturbances adequately. Generalizing this finding, we hypothesize that a single muscle stimulation pattern exists that yields adequate performance for a wide range of starting positions. Direct experimental confirmation of any of the control strategy hypotheses mentioned is difficult. As a start, we feel it is justified to test our hypothesis using a modeling and simulation approach. The results of this test are described in this study. After showing that the salient features of the “real” system are adequately represented in the model, it will be shown that it is indeed possible to find a single muscle stimulation pattern that results in successful performance of human squat jumps initiated from a wide range of static starting positions. Once more, we stress that a control strategy based on using such a stimulation pattern would imply an enormous simplification of the control problem associated with handling variations in starting position when compared with the specific motor program and “efference copy” hypotheses. After showing in this study that a simple control strategy along the lines of our hypothesis might be used, the next step is to investigate whether such a strategy is actually used in reality. As a first step in this direction we will show that available experimental data can be explained by the hypothesis presented in this study. METHODS Outline of the study The central question addressed in this study is whether successful performance of squat jumps initiated from widely varying static starting positions might be achieved by using a single suitably selected muscle stimulation pattern. This question is investigated using a modeling and simulation approach. After supplying a concise description of the task studied we will describe the model of the musculoskeletal system. In the simulations a simple representation of the activation of alpha-motoneurons as a function of time (henceforth called STIM pattern) and the initial position constitute the input to the model; the resulting movement constitutes the output of the model. To evaluate whether the selected STIM pattern (see below) yields successful performance when used in a specific starting position, the resulting jump height must be compared against the maximal jump height for that starting position. Thus for any starting position the optimal STIM pattern and the corresponding optimal movement and jump height must 1392 A. J. VAN SOEST, M. F. BOBBERT, be used as a reference. The details of the numerical procedure involved in the optimization of the STIM pattern will be described. Although a simulation approach is used in this study, it is the real system that we are ultimately interested in. Therefore it must be shown that the salient features of the real system are adequately represented in the model. To this aim we made a comparison between the performance of well-trained human subjects and optimal performance of the model. The assumptions underlying this comparison are that (I) the optimization criterion used in the simulations, i.e., height reached by the body’s center of mass, is the one also used by our subjects and (2) that well-trained subjects perform more or lessoptimally. A short description of the experimental procedures is given. The general approach used in the simulation experiments aimed at finding one STIM pattern that yields successfulperformance for a range of starting positions can be summarized as follows: 1) select a STIM pattern; 2) apply this STIM pattern to a number of static starting positions; and 3) compare the resulting movement patterns and jump heights to the optimal movement patterns and jump heights pertaining to these starting positions. Two simulation experiments were conducted, differing in the STIM patterns used. The STIM pattern used in simulation experiment I was an obvious choice: the optimal STIM pattern pertaining to the standard (middle) starting position. In simulation experiment II no STIM pattern was selected a priori. Rather, we attempted to find a STIM pattern that results in successfulperformance for all starting positions considered. In addition, we investigated how successfulperformance was when the resulting STIM pattern was applied to “new” starting positions, i.e., starting positions that were not used in the derivation of this STIM pattern. Descriptionof the task The type of human vertical jumping studied is the maximumheight squat jump, i.e., a jump starting from a static squatted position aimed at reaching a maximal height of the body’s center of mass. Becausejump height can be calculated from the position and velocity at takeoff, only the pushoff phase, which starts when the body’s center of massstarts moving upward and ends when the feet lose contact with the ground, is considered. Successof jumps is not strictly defined in this study. Becausethe aim of the task is to jump as high as possible,jump height is the most important determinant of success;however, the movement pattern and specifically the position of the segments and the direction of the velocity of the body’s center of mass at takeoff are taken into account as well. Model of the musculoskeletalsystem The model used in our simulations of human vertical jumping has been extensively described elsewhere (van Soestet al. 1993). It consists of a submodel describing the skeletal structure and a submodel describing the behavior of the muscles. The skeletal submodel is two-dimensional and consists of four rigid segments representing feet, lower legs, upper legs, and headarms-trunk. These segments are connected in frictionless hinge joints representing hip, knee, and ankle joints. At the distal end of the foot segment the skeletal model is connected to the rigid ground by a fourth hinge joint that can be considered as a representation of the metatarsophalangeal joint. Possibleground contact at the heel is assumed to be elastic. Acceleration-determining forces are the gravitational forces, the force at the heel in case of ground contact, and the moments acting at the joints that represent the net action of the muscles. The dynamic equations of motion of the skeletal model were derived using SPACAR, a software subroutine package developed at Delft University of Technology (van Soestet al. 1992; van der Werff 1977). These equations allow calculation AND G. J. VAN INGEN SCHENAU of the acceleration of the skeletal model as a function of position, velocity, and the forces mentioned above. Segment lengths for the skeletal model as well as initial position were directly based on the mean values of the experimental subjects. The initial position based on experimental data will be referred to as the standard initial position. Segment masses, locations of segment mass centers, and segmental moments of inertia were estimated from segment lengths and body mass of our subjects, in conjunction with cadaver data reported by Clauser et al. ( 1969). The muscular submodel consists of six “muscles” representing the major muscle groups that contribute to extension of the lower extremity, i.e., gluteal muscles, hamstrings, vasti, rectus femoris, soleus, and gastrocnemius. A Hill-type muscle model was used to represent these muscles. It consists of a contractile element, a series elastic element, and a parallel elastic element. Behavior of the elastic elements is governed by nonlinear force-length relationships. Behavior of the contractile element is more complex: contractile element contraction velocity depends on active state, contractile element length, and force. Force is directly related to the length of the serieselastic element. This length can be calculated at any instant from the position of the skeleton and contractile element length (which are used as state variables) because muscletendon complex length is directly related to position of the skeleton. Active state is related to muscle stimulation STIM, the independent neural input of the model, by first-order dynamics as described by Hatze ( 198 1). STIM ranges between 0 and 1 and is a one-dimensional representation of the effects of recruitment and firing rate of alpha-motoneurons. Wherever possible, parameter values for the muscles were derived on the basis of morphometric data. Most importantly, contractile element optimal lengths were based on sarcomere numbers; moment arms of the muscles were based on muscle length versusjoint angle measurements; relative values for maximal isometric forces were based on muscle cross-sectionalareas. All these measurements were made on the same group of cadavers. Sarcomere numbers and moment arms were scaled in proportion to segment lengths; absolute values of muscle forces were set in such a way that realistic maximal isometric moments were obtained. The force-velocity relation was based on Hill’s classicalequation (Hill, 1938) where FcEis contractile element force, FyIAxis maximal isometric force, V& is contractile element contraction velocity, and LCE(OPT) is contractile element optimal length. The dimensionless parameters dFnfi*x and WCE(OPT)were set to 0.41 and 5.2, respectively, for all muscles. The stretch of the serieselastic element at maximal isometric force was set at 0.04 times its slack length for all muscles. A detailed discussion of the structure of the muscle model and the strategy followed in the estimation of parameter values has been presented elsewhere (van Soest and Bobbert 1993; van Soestet al. 1993). In total, the model is mathematically described by a set of 20 coupled nonlinear first-order ordinary differential equations. Given the initial state and given the independent control signals (i.e., STIM) as a function of time, the resulting movement can be calculated by numerical integration. A variable-order variablestep sizeAdams-Bashford predictor Adams-Moulton corrector integration algorithm was used (Shampine and Gordon 1975). Optimization of STIM pattern The problem is to find the STIM( t) pattern that leads to a maximal vertical jump, i.e., to a maximal height of the body’s center of mass. This dynamic optimization problem was studied in its full complexity by Pandy et al. ( 1990). Partly on the basis of their results a more restricted form of dynamic optimization is used in CONTROL OF EXPLOSIVE this study. Specifically, the following restrictions were imposed on STIM: first, the initial STIM level was set in such a way that the static squatting starting position was maintained. Second, STIM was allowed to take on either this initial value or the maximal value of 1.0. Third, STIM was allowed to switch to its maximal value just once and thereafter remained maximal until takeoff. Under these restrictions, STIM ( t ) of each of the six muscle groups is described by a single parameter: the instant at which STIM switches from initial value to maximal value. The optimization problem is thus reduced to finding the combination of six switching times that results in maximal jump height. Although this is still a computationally demanding problem, it can be solved using standard algorithms. NAG subroutine E04UCF, a sequential quadratic programming algorithm, was used (NAG Fortran Library Manual Mark 13, Numeric Algorithms Group, Oxford, UK). In the following, the optimal STIM pattern pertaining to the initial position derived from experimental data will be referred to as the standard STIM pattern and similarly the resulting movement will be referred to as the standard movement. Severe constraints were imposed on STIM in this study to keep calculation time manageable and to ensure that a well-defined optimum could be found. To investigate the extent to which these constraints influenced performance, optimizations were performed where each muscle was given more “freedom:” each was allowed to switch on two times and to be deactivated in between. Although this enhanced jump height, the enhancement was only ~2 mm. Furthermore, numerous local optima appeared to exist. It is concluded that by reducing the dimension of the control space to 6, as is done in this study, the optimization problem becomes more well-behaved and the reduction in jump height is small. Experimental procedures Six elite male volleyball players performed a number of jumps starting from a freely chosen static squatting position. Subjects were instructed to jump as high as possible, to make no countermovement before pushoff, and to keep their hands on their backs. Trials in which these instructions were violated were discarded. The highest successful jump of each subject was selected for analysis. During jumping, kinematic data were gathered using a lOO-Hz VICON video registration and analysis system and ground reaction force was measured using a force platform (Kistler 928 1B) . These data were used for an inverse dynamic analysis using standard procedures ( Bobbert et al. 1993 ) . Surface EMG was recorded simultaneously using a Biomess 80 telemetric system. After amplification and analog band-pass filtering (20-200Hz), EMG data were sampled at 1,000 Hz and stored in digital form. Simulation experiment I: locally optimal STIMpattern As stated above, the hypothesis tested here is simply that the optimal STIM pattern pertaining to a “preferred” static starting position is always used and that this results in successful performance for a wide range of starting positions around the preferred one. To test this hypothesis the following steps were taken. First it was decided to treat the experimentally derived standard starting position asthe preferred starting position and to treat the standard STIM pattern correspondingly. Next two additional starting positions were defined. All three starting positions are shown in Fig. 1 (positions 1, 3, and 5). In all three, the horizontal position of the body’s center of mass and the point of ground contact at the metatarsophalangeal joint are equal. The two additional starting positions differ from the standard one in the vertical position of the body’s center of mass by amounts of to.1 m. As for the standard starting position, initial state of the muscles was calculated from 1393 MOVEMENTS 1 2 3 4 5 6 FIG. 1. Static initial positions used in the simulation experiments. Dot: position of body’s center of mass. Position 3 was derived from experimental data. See text for details. the demand of static equilibrium. Next the standard STIM pattern was applied to the additional two starting positions. As explained earlier, for comparison the optimal movement pattern and optimal jump height for the additional starting positions had to be obtained as well. This was done using the optimization approach described above. Finally, for each of the additional starting positions, the performance resulting from application of the standard STIM pattern was compared with the optimal performance for that starting position. Simulation experiment II: globally optimal STIM pattern As indicated earlier, no STIM pattern is selected a priori in this simulation experiment. Rather, the question asked here is whether a single STIM pattern can be found that resultsin successfulperformance for a number of starting positions. Note that if such a pattern can be found it is not likely to be optimal for any specific starting position! Despite the stringent constraints imposed on STIM in this study, the number of possible STIM patterns is still extremely large. As a consequence it is not possible to find the STIM pattern we are looking for by checking all possible STIM patterns. Instead the following approach was adopted. First, it was decided to use the three starting positions used in simulation experiment I (Fig. 1, positions 1,3, and 5 ) asthe “learning set” in the determination of the globally optimal STIM pattern. Next we decided to consider averagejump height for these three starting positions as the measure of successof any specific STIM pattern. In other words, the best STIM pattern was defined to be the one that yields the highest average jump height when applied to all three starting positions. Obviously, finding this STIM pattern constitutes an optimization problem. In fact, this optimization problem is similar to those encountered earlier, and consequently the numerical optimization method described earlier was fully applicable. Application of this optimization procedure resulted in a STIM pattern that will be called the “globally optimal STIM pattern.” Next, to evaluate just how successfulthe performance resulting from application of this globally optimal STIM pattern to all three starting positions was, this performance was compared with optimal performance for these three static starting positions, as obtained earlier. Finally we investigated whether the globally optimal STIM pattern resulted in successful performance when applied to starting positions that were not included in the learning set. To this aim, three additional starting positions were defined, two of which fell within the range covered by the learning set and one of which fell outside that range. The globally optimal STIM pattern was applied to these three additional starting positions and resulting performance was compared to the relevant optimal performance. These additional initial positions are shown in Fig. 1, positions 2, 4, and 6. Positions 2 and 4 are midway (in terms of vertical position of the body’s center of mass) between the three A. J. 1394 VAN SOEST, M. F. BOBBERT, AND G. J. VAN INGEN SCHENAU A GLU HAM VAS REC SOL GAS T = 0.0 T = -0.310 -0 1 35 -0.3 1 I I I I -0.25 -0.2 -0.15 -0.1 -0.05 0 TIME [s] . cccc T = -0.370 I c T = 0.0 , -0.9 I -0.8 1 -0.7 1 -0.6 1 -0.5 1 -0.4 , -0.3 I -0.2 1 -0.1 0 TIME [s] 2. A : stick-figure representation of the pushoff movement of the model resulting in maximal jump height. Velocity of the body’s center of mass is represented by a vector that originates from the body’s center of mass. B: stick-figure representation of the mean movement of well-trained experimental subjects performing a maximal squat jump ( yt = 6). C: muscle stimulation as a function of time for gluteal muscles (GLU), hamstrings ( HAM), vasti (VAS), rectus femoris (REC), soleus ( SOL), and gastrocnemius (GAS) muscles. Drawn lines: period of maximal activation of alpha-motoneurons as a function of time (STIM). Top line of each pair concerns the optimal STIM pattern for starting position 3; bottom line concerns the globally optimal STIM pattern. D: typical example of surface electromyogram (EMG) patterns and vertical ground reaction force (top tracing) obtained during pushoff. ST, semitendinosus; BF, biceps femoris, caput longum. Vertical line: start of the pushoff phase. Note the difference in scaling of the time axes between C and D. In all figures, time is expressed relative to takeoff. FIG. starting positions used earlier (i.e., positions 1, 3, and 5 in Fig. 1). Vertical position of the body’s center of mass in position 6 is 5 cm higher than in position 5. RESULTS Comparisonof experimentaldata and simulation data Before showing results pertaining to the main question of this study it must be shown that the salient features of the real system are adequately represented in the model used. To this aim, simulation data pertaining to the optimal (i.e., maximum-height) jump are compared with experimental data. In Fig. 2, A and B, stick-figure representations of the movement are given both for our experimental subjects ( Fig. 2 B) and for the optimal simulation data ( Fig. 24. Note that the initial position for the model was set to equal that of our experimental data; the correspondence in this respect is imposed. Comparing these stick diagrams a number of general observations can be made. First, the general kinematic patterns are similar: a continuous extension occurs at all joints except at the ankle joint in the first part of pushoff. Second, a near-perfect alignment of body’s center of mass with the point of ground contact is observed. Third, CONTROL TABLE 1. Experiment Simulation OF EXPLOSIVE MOVEMENTS 1395 Comparison of experimental and simulation data of maximum-height squat jumps H, m Y cm9 m @a,rad 0.447 0.098 0.048 0.52 0.60 0.390 +k, rad 0.12 0.40 (bh, l-ad 0.09 0.5 1 Ycm, m/s 2.67 2.59 dir,, rad w, J 0.0525 0.0430 612 657 All jumps are from starting position 3 (see Fig. 1). H, jump height relative to upright standing; y,,, height of body’s center of mass at takeoff, relative to upright standing; &, &, +h, ankle, knee, hip, angle at takeoff, relative to full extension; pcrn,vertical velocity of body’s center of mass at take-off; dir,, absolute angle of the velocity vector at takeoff, relative to the vertical; W, total mechanical work done by the muscles. Experimental data are averaged over subjects (n = 6). See text for details. the velocity of the body’s center of mass at takeoff is almost perfectly vertical. In Fig. 2C (top line of each pair) the optimal STIM pattern is presented. In Fig. 2D, surface EMG tracings for a maximum-height squat jump of a typical subject are shown together with the vertical ground reaction force. Taking surface EMG as a measure of the muscle activation level, two observations can be made when comparing Fig. 2, C and D. First, the EMG data support the assumption that, once activated, muscles remain active until takeoff, although maximal activity is not maintained in all muscles. Second, the notion of a proximal-distal sequence in muscle activation (e.g., Bobber-t and van Ingen Schenau 1988) is confirmed by both the EMG data and the optimal STIM pattern. To allow a more quantitative comparison between experiment and simulation, a number of parameters related to achievement in vertical jumping are presented in Table 1. The first thing to note from this table is that jump height for the model is 13% lower than that observed experimentally. As analyzed in detail in van Soest et al. ( 1993 ), this is because of the fact that the upper body is represented by a single rigid segment in the model, as a result of which trunk extension cannot contribute to jump height. This trunk extension by itself results in raising the body’s center of mass at takeoff; additionally, the inertial forces associated with trunk extension postpone takeoff, as a result of which the joints of the lower extremities are closer to full extension in reality than in simulation (see Table 1). Thus the difference in jump height is largely caused by a higher position of the body’s center of mass at takeoff; the difference in vertical velocity of the body’s center of mass is relatively small. For both experiment and simulation, the direction of the velocity vector at takeoff is within 3O of the vertical; thus very little work is invested in horizontal kinetic energy. Finally, the mechanical work (i.e., sum over all joints of the integrals of net joint moments over joint angle) done by the muscles during pushoff is compared. Given the difference in jump height, it is surprising that the total work is higher in simulation. The unexpectedly low value of the experimentally observed muscle work is, again, due to trunk deformation: in movement registration and inverse dynamic analysis the trunk was assumed to be rigid. Thus in the experimental value work done by the upper body muscles is neglected, which explains the relatively low work value calculated from the experimental data. Additional data on this comparison of experiment and simulation have been presented in van Soest et al. ( 1993). It is concluded that although simplifications in the musculoskeletal model result in certain differences the degree of correspondence between experimental and simulation data is such that the model can be used with confidence to investigate the question addressed in this study. Simulation experiment I: locally optimal STIMpattern In this simulation experiment we tested the hypothesis that the optimal STIM pattern pertaining to a specific starting position is always used and that this strategy results in successful performance for a wide range of starting positions around that specific one. To test this hypothesis we applied the optimal STIM pattern pertaining to the experimentally derived standard starting position (standard STIM) to two additional starting positions (Fig. 1, positions 1 and 5). This resulted in movements that are presented in the form of stick diagrams in Fig. 3, D and E, along with corresponding jump heights relative to upright standing. For comparison, the optimal performance pertaining to these starting positions is presented in Fig. 3, A and C. In addition, values of a number of parameters related to achievement in vertical jumping are presented in Table 2. Both for the optimal jumps and for the jumps resulting from applying standard STIM to starting positions 1, 3, and 5, average values are presented in Table 2. Comparison of Fig. 3, A and Cand D and E indicates that application of the standard STIM pattern to widely varying starting positions results in movements that differ considerably from the optimal movements for these starting positions. For example, in starting position 1 hyperextension of the hip is present at takeoff, which clearly does not occur in either the optimal jump for that starting position or in reality. Even more importantly, jump height is seriously affected ( see also Table 2 ) : averaged over starting positions 1, 3, and 5 jump height is 0.115 m less, which in a relative sense amounts to as much as 30%. This deterioration of performance is for the major part because of a lower vertical velocity at takeoff and to a lesser extent because of a lower position of the body’s center of mass at takeoff. In turn, this lower velocity is caused by a lower amount of energy delivered by the muscles during pushoff: whereas the averaged value for the work done equals 652 J for the optimal performance, it equals 558 J on average in cases where standard STIM is applied to each of the starting positions 1, 3, and 5. Although the direction of velocity at takeoff diverges more from the vertical when applying standard STIM than when applying optimal STIM (0.0992 vs. 0.0405 rad), the importance of this parameter in terms of work wasted in the form of horizontal kinetic energy is small. In conclusion, the hypothesis that application of standard STIM to a wide range of starting positions results in A. J. T = -0.356 A VAN SOEST, M. F. BOBBERT, T = -0.310 AND G. J. B VAN INGEN SCHENAU T = -0.256 H = 0.215 H = 0.208 T = -0.386 T = -0.256 D H = 0.352 H = 0.423 T = -0.374 F C T = -0.306 T = -0.262 U H FIG. 3. Stick-figure representation of pushoff movements obtained in simulation experiments I and II concerning starting positions 1 (A, D, and F), 3 (B and G), and 5 (C, E, and H). A-C: optimal movement for these starting positions, used as reference. D and E: movements obtained in simulation experiment I. F-H: movements obtained in simulation experiment II. H: resulting jump height. For further details see legend to Fig. 2 and main text. close to optimal achievement was refuted by the results of this simulation experiment. Simulation experimentII: globally optimal STIMpattern In this simulation experiment we attempted to find one single STIM pattern that results in successful performance for a number of static starting positions. Second we investigated whether this STIM pattern results in successful performance for new starting positions. To find the globally optimal STIM pattern we performed optimization with respect to average jump height over the initial positions 1,3, and 5 as indicated in Fig. 1. The resulting STIM pattern is depicted in Fig. 2C, together with the optimal STIM pattern for starting position 3. For the gluteal muscles, the hamstrings, and the vasti group, the time of onset of maximal stimulation differs by < 10 ms. Rectus femoris is activated 34 ms earlier in the globally optimal STIM pattern. The most significant difference concerns the ankle plantar flexors soleus and gastrocnemius. In both cases they are the last activated muscle group. In the optimal STIM pattern, the biarticular gastrocnemius was activated 84 ms after the monoarticular soleus; in the globally optimal STIM pattern this sequence was reversed. This reversal can be speculated to have a relatively small effect on the (plantar flexing) ankle joint moment; the main consequence is that a direct coupling between ankle and knee joints is created earlier in the pushoff. For an extensive discussion of the relation between subtle changes in STIM and the resulting movement the reader is referred to Bobbert and van Soest ( 1994). The movement resulting from applying the resulting globally optimal STIM pattern to all three initial positions is presented in Fig. 3, F-H, along with corresponding jump heights and pushoff phase durations. Note that application CONTROL TABLE 2. OF EXPLOSIVE MOVEMENTS 1397 Values of numerical parameters related to successof simulated vertical jumps Optimal STIM Avg-opt STIM Standard STIM K m Y cm7 l-n 0.386 0.365 0.27 1 0.048 0.045 0.017 &I, rad 0.61 0.57 0.93 hc, rad 0.39 0.39 0.14 4h, I-ad 0.47 0.47 0.13 iQcrn3 m/s 2.57 2.49 2.21 dir,, rad 0.0405 0.0343 0.0992 w J 652 638 558 Val ues are averaged over starting positions 1,3 and 5 (see Fig. 1). Optimal STIM, optimal jumps for each starting position; Standard STIM, jumps that result from applying the optimal STIM for starting position 3; Avg-opt STIM, jumps that result from applying the single globally optimal STIM pattern. For other abbreviations see Table 1. of the same STIM pattern to different starting positions leads to pushoff movements with durations varying between 0.262 and 0.374 s. Similarly, as shown in Fig. 4, the net joint moments that result from application of the same STIM pattern to different starting positions vary largely. However, from Fig. 5 it is seen that application of this single STIM pattern results in decreasing variability in joint angles as the instant of takeoff is approached. The decreasing variability in these joint angle-time plots can be quantified by looking at their SDS. Specifically, the SD of the joint angles for starting positions 1, 3, and 5, summed over hip, knee, and ankle joints, decreased from 0.66 radians at the start of the pushoff phase to 0.49 radians at takeoff. Thus in a relative sense this SD decreases by 26% during pushoff. Numerical values of parameters related to achievement are presented in Table 2, again averaged over starting positions 1, 3, and 5. In comparison with the application of standard STIM to each of these starting positions, results are much closer to the optimal values here. Jump height resulting from application of this single STIM pattern to the three starting positions is on average only 0.02 m less than optimal, which amounts to 5% in a relative sense. Joint angles at takeoff are virtually identical to the optimal values; as a result, the same is true for the position of the body’s center of mass. Thus the lower jump height is due to a lower velocity at takeoff, which in turn is due to a slightly lower amount of work done by the muscles. Finally, the direction of the velocity at takeoff diverges from the vertical by an amount comparable with that found for the optimal jumps (0.0343 vs. 0.0405 rad). It is concluded that it is indeed possible to find one single STIM pattern that results in close to optimal performance for a number of static initial positions. To test the second part of the idea, concerning generalization to new starting positions, three extra static starting positions were defined, two of which (2 and 4) fall inside the range delimited by starting positions 1, 3, and 5 and one of which (6) falls outside that range (see Fig. 1). Note that starting positions 2,4, and 6 were not involved in the optimization procedure that yielded the globally optimal STIM pattern. Application of the globally optimal STIM pattern to these starting positions results in movement patterns presented in Fig. 6, D-F. For comparison optimal movement patterns for these starting positions are shown in Fig. 6, A-C. Numerical values of relevant parameters are presented in Table 3 for interpolated (2 and 4) and extrapolated (6) starting positions separately. For these extra starting positions jump height resulting from application of the globally optimal STIM pattern is -0.04 m. less than optimal. As would be expected, this difference is larger than that for the starting positions used in finding the globally optimal STIM pattern, which was ~0.02 m (see Table 2). Yet the difference is still 5 10% from optimal. As a consequence, the same holds true for the variables that determine jump height, i.e., height of the body’s center of mass and body’s center of mass vertical velocity at takeoff (see Table 3). As can be seen from Fig. 6 and Table 3, a similar trend is also found for joint angles at takeoff and the verticality of the jump: the differences from optimal values are small, but larger than those for the initial positions involved in the derivation of the globally optimal STIM pattern. Note that the “interpolated” starting positions (2 and 4) do not result in performance that is clearly closer to optimal than the performance obtained when starting from position 6 that was outside the range used in finding the globally optimal STIM pattern. In conclusion, performance resulting from application of the globally optimal STIM pattern to new starting positions is 5 10% from optimal. This is true not only for starting positions lying in the range used in the derivation of the globally optimal STIM pattern but also for a starting position outside that range. DISCUSSION In this study we generated a parsimonious explanation for the observation that humans are able to perform explosive movement tasks from various starting positions. According to the explanation this remarkable ability might be achieved on the basis of an equally remarkable control strategy: using one and the same muscle stimulation pattern for all initial positions! Support for this explanation was gathered on the basis of simulation experiments. The idea that such a strategy might work was arrived at on the basis of simulation results described elsewhere (van Soest and Bobbert 1993) that dealt with the question how disturbances occurring during the execution of explosive movements are handled. In that study it was shown that intrinsic muscle properties counteract such disturbances to such a degree that adaptation of the muscle stimulation pattern is not necessary. A possibility for experimental verification of this finding lies in direct measurement of joint stiffness and damping during movement. Such measurements were recently undertaken by Bennett et al. ( 1992) during slow, low-force arm movements. Because stiffness and damping during such movements are primarily determined by neural feedback-based adaptation of muscle activation, as opposed to by intrinsic muscle properties, the results of Bennett et al. ( 1992) can unfortunately not be generalized to the fast, high-force explosive movements discussed in this study. Two STIM patterns that possibly result in successful performance when applied to a range of starting positions were 1398 4OOb 1 . SOEST, M. F. BOBBERT, . 350- r . I I HIP JOINT E z - VANVAN A. J. AND G. J. SCHENAU 3- 9 *e.. ’ INGEN . 2.8- I I . HIP JOINT / \ L 3009 5 w z 250- ii Iz 07 100 . I- W t 01 -0.4 1 I I n I . 1 -0.35 -0.3 -0.25 -0.2 -0.15 -0.1 -0.05 TIME 350 . . Jo.4 1 1 250- 2 O z 200- 5 e 150- 2 !g loo- 1 1 I 3 -0.35 -0.3 -0.25 ". 1 . I I -0.1 -0.05 2.4 -0.1 . -0.05 1 1 1 1 0I 1 1 [s] 1 / 1 i Jo.4 1 0 -0.35 1 -0.3 1 -0.25 I -0.2 I TIME I I I -0.15 -0.1 1 -0.05 1 0I -0.15 -0.1 . -0.05 I 0 [s] 1 1-1 2.6 ANKLE JOINT 250- Iz 200- 02 1509 W 2 -0.15 1 KNEE JOINT [s] ANKLE JOINT 'E' zd -0.2 1 2.6 E s - -0.15 -0.2 TIME 300 . 2.81-l I \ 504 -0.4 -0.25 I TIME . 3009 l5 -0.3 1 [s] KNEE JOINT 'E' zl -0.35 1 0 I- FIG. z 0 -.-.0-w loo, Ot -0.4 1 -0.35 - - . -0.25 -0.3 I -0.2 TIME . -0.15 -0.1 . -0.05 [s] 1.21 -0.4 1 -0.35 -0.3 . -0.25 I -0.2 4. Net joint moments versus time (relative to takeoff) for nip (top), knee (middZe), and ankle (bottom) joints, resulting from applying the globally optimal STIM pattern obtained in simulation experiment II to starting positions 1 ( -), 3 ( - l ), and 5 (- - -). See text for details. TIME [s] 5. Joint angles vs. time (relative to takeoff) for hip (top), knee (middle), and ankle (bottom) joints, resulting from applying the globally optimal STIM pattern obtained in simulation experiment II to starting positions 1 ( -), 3 ( - l ), and 5 (- - -). See text for details. investigated. These STIM patterns represent two ends of a continuum. The first of these is the optimal STIM pattern for a “middle” starting position. To arrive at such a pattern, humans would have to “tune” the stored muscle stimulation pattern in such a way that it yields optimal performance when applied to a preferred starting position. Evi- dently application of this muscle stimulation pattern to other starting positions will result in suboptimal performance; because of the effect of muscle properties, however, performance will be successful as long as starting positions are sufficiently close to the preferred starting position. The second STIM pattern is optimal with respect to average l FIG. l CONTROL OF EXPLOSIVE MOVEMENTS 1399 H = 0.306 j i T = -0.336 T = -0.250 A H = 0.361 T = -0.202 B C H = 0.273 H = 0.323 1 T = -0.336 T = -0.268 D T = -0.234 E F 6. Stick-figure representation of pushoff movements concerning starting positions 2 (A and D), 4 (B and E), and 6 ( C and F) . A-C: optimal movement for these starting positions, used as reference. D-F: movements resulting from applying the globally optimal STIM pattern obtained in simulation experiment II to these “new” starting positions. For further details see legend to Fig. 3 and main text. FIG. jump height achieved. To arrive at this pattern humans would have to tune the stored muscle stimulation pattern in such a way that reasonably successful performance is obtained for a wide range of starting positions. In other words, such a strategy is aimed at establishing “control that works” (Gottlieb et al. 1990; Hogan and Winters 1990) as opposed to optimal control. In simulation experiment I we found that the locally optimal STIM pattern does not result in successful performance for a wide range of starting positions. Thus this pattern is not the one we are looking for in this study. It seems that the price that has to be paid for local optimality is that the range of starting positions for which successful performance is obtained is relatively narrow. Although it is conceivable that expert performers are willing to narrow the range of allowed starting positions in return for higher achievement, such a strategy is probably not favored by most people. In simulation experiment II we found that the globally TABLE 3. optimal STIM pattern results in successful performance for a wide range of starting positions. Thus our hypothesis was confirmed in this experiment: a single control pattern that works for a wide range of starting positions can indeed be found. In simulation experiment II it was further shown that the STIM pattern that works works for new starting positions as well. Thus the novelty problem, traditionally one of the weak points of motor program theories, does not apply to the theory presented here. In conclusion, the results of simulation experiment II convincingly show that in return for an insignificant loss ofjump height an enormous simplification of the control problem can be obtained: the control problem is reduced to learning, storing, and retrieving just one muscle stimulation pattern. Having shown that the proposed control strategy works in a simulation context, two questions must be considered: I) would such a strategy work in reality and 2) is it actually used in reality. An affirmative answer to the first question can be given only if we are confident that the model ade- Values of numerical parameters related to successof simulated vertical jumps K Positions 2 and 4 Optimal STIM Avg-opt STIM Position 6 Optimal STIM Avg-opt STIM l-n Y cm9 In (b,, rad AC, I-ad 0.386 0.342 0.050 0.044 0.64 0.38 0.38 0.48 0.43 0.67 0.306 0.273 0.058 0.05 1 0.60 0.72 0.35 0.21 0.34 0.25 Values concern starting positions 2 and 4 (interpolation) and 6 (extrapolation) h, rad dir,, rad w9 J 2.57 2.42 0.0463 0.0406 642 624 2.21 2.09 0.0153 0.0104 445 412 iQcm9 m/s (see Fig. 1). For abbreviations see Table 1. 1400 A. J. VAN SOEST, M. F. BOBBERT, quately represents the salient features of the real system. This confidence was gained on the basis of a comparison of experimental data concerning expert performance and simulation data concerning optimal model performance. In other words it is likely that the strategy suggested would work in reality. Thus the question remains whether it is actually used. At present experiments are being conducted that address this question. A tentative theory The theory emanating from these simulation experiments can be summarized as follows. Execution of explosive movement tasks is governed largely by open loop control. For specific groups of tasks, e.g., “explosive leg extensions,” a muscle stimulation pattern is stored in some at present unspecified form within the CNS. Any time such a task has to be performed the muscle stimulation pattern is called on. Disturbances occurring during the unfolding of the movement cannot be corrected fully on the basis of neural feedback because of stringent time constraints. Instead such disturbances are initially counteracted by the viscoelastic behavior resulting from the force-length-velocity relation of muscle. The stored pattern is molded by experience. Typically this results in control that works, i.e., in a muscle stimulation pattern that yields successful performance for a wide range of starting positions. Even for starting positions that were never encountered before, reasonably successful performance is achieved. Alternative strategies for coping with variations in starting posit ion As mentioned in the INTRODUCTION, the strategy proposed in this study to cope with variations in starting position is not the only possible one that might be successful. At least two alternative strategies can be mentioned and will be shortly discussed here. In the first alternative strategy, which is based on efference copy theory (von Holst and Mittelstaedt 1950), it is assumed that the muscle stimulation pattern is formed as the movement unfolds. The efference copy theory was formulated to explain observations like the following. When an animal’s motions are restricted to movement of the eyes it is obvious that identical changes in the retinal image can result from either movement of the eye or movement of the world. Yet from the animal’s behavior it is concluded that it is able to distinguish between these possibilities. According to efference copy theory, this is achieved by sending a copy of the “motor command” to an “internal representation.” There the eye movement that is going to result from this command is “calculated” and this movement is “subtracted” from the movement of the retinal image. This subtraction results in the movement of the world, which solves the animal’s problem. The central aspect of the theory is the internal representation, in which the movement that will result from the motor commands is calculated. Theories in which the use of such an internal representation of the effector system is assumed are called efference copy theories. The efference copy theory pertaining to the control of explosive movements states that the expected movement calculated in the internal representation is used in the genera- AND G. J. VAN INGEN SCHENAU tion of the subsequent motor commands. The advantage of the use of such an internal representation over peripheral feedback is, of course, that delays occurring in the internal representation are much smaller. Although the theory in its general form is elegant, its application to the problem of movement control requires that the internal representation be highly accurate. For a mechanically simple system such as the eye, existence of a sufficiently accurate representation is conceivable, and efference copy theory gives rise to an elegant explanation. By contrast, a sufficiently accurate representation of the motor system of the extremities would have to be extremely complicated, much more complicated than the simulation model used in this study, in fact. In our view, explanations of the control of explosive movements based on the assumption that such a complex internal representation is present in the CNS are not attractive. The second alternative strategy is based on a priori formation of a motor program. In fact, several versions of such a strategy can be distinguished. According to the first a single muscle stimulation pattern is stored. This stored pattern is the optimal one for a standard starting position. Because of muscle properties this pattern will lead to acceptable movements for a small range of starting positions around the standard one. To execute the task from any starting position a relatively slow movement is initiated in the direction of the standard starting position under feedback control; as soon as the small range mentioned is reached, the optimal muscle stimulation pattern is triggered. In fact, such a strategy was used in an early attempt at deriving the optimal control for a simplified model of vertical jumping (Levine et al. 1983). However, two arguments may be raised against such a strategy. In the first place, experimental data (Bootsma and van Wieringen 1990; M. F. Bobber-t, K. G. M. Gerritsen, M. C. A. Litjens, and A. J. van Soest, unpublished data) do not yield any evidence that such a strategy is used. Second, such a strategy would result in increases in movement time that are unacceptable in tasks such as hitting an approaching object, in which timing aspects are imposed externally. According to the second version of motor program theory, a specific muscle stimulation pattern is stored for every starting position ( Atkeson 1989). This version is not attractive, because it raises more questions than it answers: the storage space required is virtually infinite; it cannot be imagined how the correct program can be retrieved fast enough; from the theory it is absolutely unclear how a control pattern is generated when a new starting position is encountered (the “novelty problem”). Finally, according to the third version of motor program theory, a general motor program is stored in which a few parameters are set to obtain a specific motor program (e.g., Carter and Shapiro 1984). In the context of the problem considered in this study, this would imply that the specific motor program is formed on the basis of feedback of the initial position. This version cannot be dismissed at present; however, the relations between starting position and values for the parameters to be set are probably not simple. To our knowledge such relations have never been described for any specific task. In conclusion, alternative control strategies for dealing with variations in initial position that are mentioned in liter- CONTROL OF EXPLOSIVE ature are less parsimonious than the theory outlined in this study. Furthermore, the theory presented in this study is supported by concrete simulation results, which is not the case for the theories discussed above. Possibleneurophysiologicalimplementationof the ideas presented An intriguing question is how the muscle stimulation pattern is formed in the CNS. Recent developments in the field of neural networks provide suggestions for ways in which time-varying patterns may be stored and retrieved (Coolen and Gielen 1988; Reiss and Taylor 199 1) . These authors describe relatively simple and biologically not too implausible neural networks that are able to regenerate a sequence of patterns without actually storing the patterns themselves. Such a process of sequence regeneration is triggered by bringing the network into a specific initial state. It is the strength of the connections between the neurons involved that determines what pattern is produced. As a consequence, by changing the strength of the neuronal connections involved, changes to these patterns can be made. It is imaginable that the regenerated patterns represent the default alpha-motoneuron activity. Alternatively these patterns might represent the control signals at a more global level of description, in which case the details must be filled in at a lower level. In the case of positioning tasks this global level might well be thought of as a virtual trajectory of equilibrium points (e.g., Bizzi et al. 1992; Feldman 1986). In explosive tasks an equilibrium point representation probably does not simplify the control very much, because an adequate equilibrium point trajectory is expected to be unrelated to the actual movement because of the demand of increasing velocity; however, other global representations may be conceivable. In any case, these regenerated patterns would, as far as possible together with feedback signals, finally lead to activation of alpha-motoneurons. Thus it can be imagined that the central command “JUMP!” consists of little more than a set of signals that triggers a network capable of reproducing the adequate muscle stimulation pattern. Some speculationson experimentaldata on explosive human movements Experiments directly addressing the theory proposed in this study have not yet been undertaken. In fact, studies dealing with any aspect of explosive human movements are scarce. For some of the results of the studies that have been performed, we will discuss to what extent they might be explained from our theory. An interesting set of experimental data was recently gathered by Jacobs et al. ( 1994), who analyzed the sprint start as well as maximal vertical squat jumps of elite sprint runners. Obviously both tasks require explosive leg extension. It was found that these subjects’ performance in the pushoff in vertical jumping was very poor, in contrast with their performance in the pushoff in sprint running. In the light of the ideas presented this finding can be explained as follows. By performing numerous sprint pushoffs sprint runners tune the neuronal network devoted to generating the control pattern for explosive leg extensions to this specific situa- MOVEMENTS 1401 tion. In other words, they end up with a muscle stimulation pattern that is optimal for the sprinting situation. As was concluded from the results of simulation experiment I, however, the price that has to be paid for obtaining optimal performance in sprint running is a reduction in performance of related tasks that use the same muscle stimulation pattern, such as vertical jumping. Another group of interesting findings concerns hitting tasks where not only is a high velocity important, but high demands are imposed on final spatiotemporal accuracy as well. Examples of such tasks are the attacking forehand drive in table tennis (Bootsma and van Wieringen 1990) and batting in cricket ( McLeod 1987). Expert performance on all such tasks is highly consistent, meaning that intraindividual variability is small over the entire movement. However, indications exist (Bootsma and van Wieringen 1990) that the remaining variability in fact decreases as the instant of ball contact is approached. This finding cannot be explained on the basis of feedback, because the movement time is well below 200 ms for this task. How then is this reduction in variability achieved? Surprisingly, Bootsma and van Wieringen devote no attention to this question. In our view a reduction in variability as the instant of ball contact is approached may result from using one single muscle stimulation pattern for all starting positions considered. This view is based primarily on the results presented in Fig. 5, which show for human squat jumping that when the globally optimal STIM pattern is used variability in joint angles decreases as the instant of takeoff is approached. The observed decrease in this case is the more surprising because the optimization criterion (average jump height) was unrelated to kinematic variability. 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