The Influence of Intrinsic Muscle Properties on Musculoskeletal System Stability: A Modelling Study

The objective of this study is to investigate how the intrinsic mechanical properties of muscles will affect the musculoskeletal system stability. A typical musculoskeletal joint driven by a pair of antagonist muscles confined only in the sigittal plane was constructed. The dynamic characteristics of the flexor and extensor muscles induced by neural inputs were represented by three dynamic processes: neural excitation, muscle activation and muscle contraction dynamics. The muscle contraction mechanics was described using a modified Hill's model with a Contractile Element (CE), a parallel elastic element and a serial elastic element. Additionally, the change of muscle Physiological Cross-Sectional Area (PCSA) and pennation angle during muscle contraction were also considered. A set of dynamic simulations were conducted by applying an external impulsive force at the distal part of the musculoskeletal system. Sensitivity analysis was conducted to investigate the effect of the CE's force-length relationship, the CE's force-velocity relationship, the force-length relationship of the serial elastic element, the parallel elastic element and the pennation angle on the system stability. The results show that the muscles with full intrinsic mechanical properties are sufficient to stabilize the system subject to an impulsive force perturbation without reflexive changes in activations. To guarantee a self-stabilizing ability, a proper CE's force-velocity relationship, the existence of a series elastic element and a sufficient muscle co-contraction level are necessary. This study would provide insight into the intrinsic design and function of the musculoskeletal system, and also give implications for the design of bionic actuators, biomimetic robotics and prosthetic devices.     

Dependence between the parameters of dynamic and stationary segments of the EMG activity in the elbow joint muscles and the joint angle during realization of targeted movements in cats

In experiments on cats we studied the pattern of EMG activity recorded from the flexors and extensors of the elbow joint and related to realization of flexor targeted operant movements of the forearm. The levels of stationary EMG activity generated by the flexors at a stabilized equilibrium position of the joint demonstrated no correlation with the values of joint angles. We suppose that this feature depends on manifestation of the hysteresis effects of muscle contraction. A target position was attained mostly due to the dynamic phases of muscle activity. The respective patterns of the movement-related activity of synergic muscles significantly differed; separate components related to leaving an equilibrium state with a certain acceleration and attaining a presettled equilibrium joint angle could be differentiated in this activity. Final positions of the forearm could be significantly different at equal levels of the stationary EMG activity generated during stabilization of these positions; they depended on specificities in the time course of dynamic phase of the activity (in particular, on the time of activity decay to a stationary level). We conclude that the movement of a limb link from one equilibrium position to another is mostly controlled via coordination of the dynamic phase of muscle activity.     

Agonist muscle activity and antagonist muscle co-activity levels during standardized isotonic and isokinetic knee extensions

This study aimed to analyze the effects of the contraction mode (isotonic vs. isokinetic concentric conditions), the joint angle and the investigated muscle on agonist muscle activity and antagonist muscle co-activity during standardized knee extensions. Twelve healthy adult subjects performed three sets of isotonic knee extensions at 40% of their maximal voluntary isometric torque followed by three sets of maximal isokinetic knee extensions on an isokinetic dynamometer. For each set, the mean angular velocity and the total external amount of work performed were standardized during the two contraction modes. Surface electromyographic activity of vastus lateralis (VL), vastus medialis (VM), rectus femoris (RF), semitendinosus (ST) and biceps femoris (BF) muscles was recorded. Root mean square values were then calculated for each 10° between 85° and 45° of knee extension (0° = horizontal position). Results show that agonist muscle activity and antagonist muscle co-activity levels are significantly greater in isotonic mode compared to isokinetic mode. Quadriceps activity and hamstrings co-activity are significantly lower at knee extended position in both contraction modes. Considering agonist muscles, VL reveals a specific pattern of activity compared to VM and RF; whereas considering hamstring muscles, BF shows a significantly higher co-activity than ST in both contraction modes. Results of this study confirmed our hypothesis that higher quadriceps activity is required during isotonic movements compared to isokinetic movements leading to a higher hamstrings co-activity.     

Interlimb interactions during cyclic in-phase and antiphase movements of arms and legs and their dependence on afferent influences

Coordinated arm and leg movements imply neural interactions between the rhythmic generators of the upper and lower extremities. In ten healthy subjects in the lying position, activity of the muscles of the upper and lower extremities was recorded during separate and joint cyclic movements of the arms and legs with different phase relationships between the movements of the limbs and under various conditions of the motor task. Antiphase active arm movements were characterized by higher muscle activity than during the inphase mode. The muscle activity during passive arm movements imposed by the experimentalist was significantly lower than muscle activity during passive arm movements imposed by the other arm. When loading one arm, the muscle activity in the other, passively moving, arm increased independently from the synergy of arm movements. During a motor task implementing joint antiphase movements of both upper and lower extremities, compared to a motor task implementing their joint in-phase movements, we observed a significant increase in activity in the biceps brahii muscle, the tibialis anterior muscle, and the biceps femoris muscle. Loading of arms in these motor tasks has been accompanied by increased activity in some leg muscles. An increase in the frequency of rhythmic movements resulted in a significant growth of the muscle activity of the arms and legs during their cooperative movements with a greater rate of rise in the flexor muscle activity of the arms and legs during joint antiphase movements. Thus, both the spatial organization of movements and the type of afferent influences are significant factors of interlimb interactions, which, in turn, determine the type of neural interconnections that are involved in movement regulation.     

Muscle activity in rapid multi-degree-of-freedom elbow movements: solutions from a musculoskeletal model

The activity of certain muscles that cross the elbow joint complex (EJC) are affected by forearm position and forearm movement during elbow flexion/extension. To investigate whether these changes are based on the musculoskeletal geometry of the joint, a three-dimensional musculotendinoskeletal computer model of the EJC was used to estimate individual muscle activity in multi-degree-of-freedom (df) rapid (ballistic) elbow movements. It is hypothesized that this model could reproduce the major features of elbow muscle activity during multi-df elbow movements using dynamic optimal control theory, given a minimum-time performance criterion. Results from the model are presented and verified with experimental kinematic and electromyographic data from movements that involved both one-df elbow flexion/extension and two-df flexion/extension with forearm pronation/supination. The model demonstrated how the activity of particular muscles is affected by both forearm position and movement, as measured in these experiments and as previously reported by others. These changes were most evident in the flexor muscles and least evident in the extensor muscles. The model also indicated that, for specific one- and two-df movements, activating a muscle that is antagonistic or noncontributory to the movement could reduce the movement time. The major features of muscle activity in multi-df elbow movements appear to be highly dependent on the joint's musculoskeletal geometry and are not strictly based on neural influences or neuroanatomical substrates. Received: 9 May 1997 / Accepted in revised form: 8 December 1998     

Hysteresis aftereffects in human single-joint voluntary movements

The single-joint voluntary plantar flexion in the ankle joint of humans was tested with an external load perturbation consisting of two opposite sinus half-waves. Pronounced manifestations of hysteresis were found in the dependence of the joint angle on the external load torque. In particular, the hysteresis displayed itself as an increase in joint stiffness following changes in the direction of movement. It led to the ambiguity of the equilibrium values of the joint angle. With goal-directed voluntary single-joint flexion and extension movements under isotonic conditions due to the corresponding changes in activation of flexor muscles only (without the activation of extensors), the hysteresis manifested itself as the uncertainty of the joint-angle dependence on the efferent activity coming to flexors during movement phases with varying prehistory. The importance of sensory information for the mechanism compensating hysteresis effects was demonstrated. The possible ways of regulation of efferent activity of the motoneuronal pools generating central motor commands in the presence of hysteresis of muscle contraction are discussed.Neirofiziologiya/Neurophysiology, Vol. 26, No. 2, pp. 83–90, March–April, 1994.     

Roles of dynamic and stationary components of a motor command in establishing the equilibrium state at simplest targeted movements

We studied in humans interrelations between the kinematic characteristics of targeted movements of the arm and current levels of EMG of the muscles providing these movements; the movements were relatively slow, and the attained joint angle was held for a time. The EMG level was considered a correlate of the level of integral motor commands (efferent activity of the respective motoneuronal pools). Application of low-amplitude non-inertial loadings, directed against the forces developed by one or another muscle group, allowed us to provide realization of targeted movements exclusively by the activity of this muscle group, without Involvement of the antagonists. It was demonstrated that the target equilibrium joint angle is reached synchronously with the dynamic phase of EMG activity, before the latter reaches a stationary level. The structure of the dynamic EMG phase itself is complex; in most cases it is split into several components. The dependence between the joint angle and amplitude of the EMG stationary phase is rather complex and variable, and usually it is difficult to predict the characteristics of this phase based on simple biomechanical considerations. There are proofs that at the performance of the studied movements and maintaining a target position there are some components in the mechanical muscle activity, which are not controlled by the motor commands. Thus, the stationary level of a motor command represents only one of several factors responsible for attaining and maintaining a target equilibrium position. Establishing this position is provided, first of all, by interaction of dynamic components of the motor commands to different muscles. Our results show that the attempts to interpret the processes of control of targeted movements on the basis of modifications of the equilibrium point hypothesis are inadequate; these data are in better compliance with the concept of impulse-temporal control; at their interpretation it is also necessary to take more thoroughly into account nonlinear properties of the muscle reactions.     

The investigation of control mechanisms of stepping rhythm in human in the air-stepping conditions during passive and voluntary leg movements

In unloading condition the degree of activation of the central stepping program was investigated during passive leg movements in healthy subjects, as well as the excitability of spinal motoneurons during passive and voluntary stepping movement. Passive stepping movements with characteristics maximally approximated to those during voluntary stepping were accomplished by experimenter. The comparison of the muscle activity bursts during voluntary and imposed movements was made. In addition to that the influence of artificially created loading onto the foot to the leg movement characteristics was analyzed. Spinal motoneuron excitability was estimated by means of evaluation of amplitude modulation of the soleus H-reflex. The changes of H-reflexes under the fixation of knee or hip joints were also studied. In majority of subjects the passive movements were accompanied by bursts of EMG activity of hip muscles (and sometimes of knee muscles), which timing during step cycle was coincided with burst timing of voluntary step cycle. In many cases the bursts of EMG activity during passive movements exceeded activity in homonymous muscles during voluntary stepping. The foot loading imitation exerted essential influence on distal parts of moving extremity during voluntary as well passive movements, that was expressed in the appearance of movements in the ankle joint and accompanied by emergence and increasing of phasic EMG activity of shank muscles. The excitability of motoneurons during passive movements was greater then during voluntary ones. The changes and modulation of H-reflex throughout the step cycle without restriction of joint mobility and during exclusion of hip joint mobility were similar. The knee joint fixation exerted the greater influence. It is supposed that imposed movements activate the same mechanisms of rhythm generation as a supraspinal commands during voluntary movements. In the conditions of passive movements the presynaptic inhibition depend on afferent influences from moving leg in the most degree then on central commands. It seems that afferent inputs from pressure receptors of foot in the condition of "air-stepping" actively interact with central program of stepping and, irrespective of type of the performing movements (voluntary or passive), form the final pattern activity.     

Motor unit firing patterns of lower leg muscles during isometric plantar flexion with flexed knee joint position

The ankle flexor and extensor muscles are essential for pedal movements associated with car driving. Neuromuscular activation of lower leg muscles is influenced by the posture during a given task, such as the flexed knee joint angle during car driving. This study aimed to investigate the influence of flexion of the knee joint on recruitment threshold-dependent motor unit activity in lower leg muscles during isometric contraction. Twenty healthy participants performed plantar flexor and dorsiflexor isometric ramp contractions at 30 % of the maximal voluntary contraction (MVC) with extended (0°) and flexed (130°) knee joint angles. High-density surface electromyograms were recorded from medial gastrocnemius (MG), soleus (SOL), and tibialis anterior (TA) muscles and decomposed to extract individual motor units. The torque-dependent change (Δpps /Δ%MVC) of the motor unit activity of MG (recruited at 15 %MVC) and SOL (recruited at 5 %MVC) muscles was higher with a flexed compared with an extended knee joint (p < 0.05). The torque-dependent change of TA MU did not different between the knee joint angles. The motor units within certain limited recruitment thresholds recruited to exert plantar flexion torque can be excited to compensate for the loss of MG muscle torque output with a flexed knee joint.     

An analysis of bailer movements responsible for gill ventilation in the crab,cancer novae‐zelandiae

The movements of the bailer during normal ventilation can be resolved into two components, a cycle of pronation and supination being superimposed on a cycle of protraction and retraction. Pronation leads protraction with a phase angle of about 90° in a normal cycle. Pronation is accompanied by flexion of the bailer.

The skeletal anatomy of the bailer is such as to restrict movements of the bailer to those described above. Further the pronated and supinated positions of the limb represent the two stable positions of a skeletal click mechanism, the operation of which may help to resolve the functional duality of the promotor and remotor muscles.

This functional duality arises because the muscles are positioned so as to produce either protraction or supination of the limb. Other muscles in the limb are monofunctional. The bulk of muscle tissue responsible for protraction and supination seems to be greater than that responsible for pronation and retraction.

The sequence of muscular activity during the ventilation cycle follows that expected for a sequence, of protraction, supination, retraction and pronation. Overlap in the periods of activity of the bifunctional muscles and muscles responsible for pronation may also help to resolve the functional duality of the former.

The amplitude of bailer excursion (protraction‐retraction) is not greatly affected by changes in frequency. An advance in the onset of activity in some muscles at higher ventilation ? frequencies suggests that the system is tailored to produce a constant beat amplitude at all frequencies.

Pauses in ventilation occur with the bailer in the retracted position, and it is maintained in this position by tonic activity in the appropriate muscle. During normal ventilation the relative contraction duration of this muscle is positively correlated with cycle period, so that pauses apparently represent a prolongation of the normal retracted phase. The relative contraction durations of some other muscles are negatively correlated with cycle period. The different signs of these correlations may be related to the type of endogenous oscillator present in the central nervous system.     

Individual muscle contributions to knee joint impedance following a sudden perturbation: An in vivo inverted pendulum model

Previous research has suggested that muscle forces, generated by reflexes, contribute to joint stability prior to the more coordinated voluntary muscle forces. The purpose of the current study was to quantify the behaviour of the leg muscles, through the calculation of individual muscle contributions to joint rotational impedance (MJRI), with a specific interest in the neuromuscular contribution in the period following shortly after a sudden knee extension perturbation. The knee was selected as an in vivo system to represent an inverted pendulum model. Kinematic and sEMG data were collected while subjects were in a prone position and exposed to sudden knee extension perturbations. A biomechanical model was used to estimate muscle forces and moments about the knee and these data were then used to calculate instantaneous MJRI. Data indicated that pre-voluntary muscle forces do contribute significantly to MJRI following a sudden knee extension perturbation as there was a 40% increase in total MJRI in the flexion/extension and valgus/varus axes immediately following the perturbation, suggesting their importance in stabilizing the joint immediately after a disturbance. Additionally, knowledge of perturbation timing was shown to increase anticipatory MJRI levels, pre-perturbation (p < 0.05), indicating that it is advantageous for the neuromuscular system to prepare for a sudden disturbance. In conclusion, the data show that the neuromuscular feedback system significantly contributes to MJRI and it is believed that this behaviour enhances joint impedance following a sudden knee extension perturbation.     

Electromyographic analysis of rotator cuff muscles in patients with rotator cuff tendinopathy: A systematic review

The shoulder is inherently an unstable joint which heavily relies on the neuromuscular activation of the rotator cuff (RC) complex for stability during movement. Currently, there is no consensus regarding how the activity of RC muscles is affected among individuals with a RC tendinopathy (RCTe). This study reviewed the evidence of studies comparing the electromyographic (EMG) activity of any RC muscle of shoulders with a symptomatic RCTe to asymptomatic shoulders. Eight databases were searched. Data from 343 participants (201 symptomatic and 209 asymptomatic shoulders) were analyzed from 10 out of 402 included studies. Strong evidence for the infraspinatus and supraspinatus during isometric contractions and limited evidence for the supraspinatus and infraspinatus during isokinetic contractions suggest that the muscular activity is not altered among individuals with a RCTe during these types of contraction. Very limited evidence indicates reduced muscle activity for the infraspinatus and subscapularis in the presence of a RCTe during isotonic contractions, and no alterations for the supraspinatus or teres minor were identified. Lastly, conflicting to moderate evidence suggests alterations in RC muscle activity during unrestrained movements and swimming. These findings indicate that EMG deficits associated with a RCTe can best be appreciated during unrestrained movements.     

System identification of muscle–joint interactions of the cat hind limb during locomotion

Neurophysiological experiments in walking cats have shown that a number of neural control mechanisms are involved in regulating the movements of the hind legs during locomotion. It is experimentally hard to isolate individual mechanisms without disrupting the natural walking pattern and we therefore introduce a different approach where we use a model to identify what control is necessary to maintain stability in the musculo-skeletal system. We developed a computer simulation model of the cat hind legs in which the movements of each leg are produced by eight limb muscles whose activations follow a centrally generated pattern with no proprioceptive feedback. All linear transfer functions, from each muscle activation to each joint angle, were identified using the response of the joint angle to an impulse in the muscle activation at 65 postures of the leg covering the entire step cycle. We analyzed the sensitivity and stability of each muscle action on the joint angles by studying the gain and pole plots of these transfer functions. We found that the actions of most of the hindlimb muscles display inherent stability during stepping, even without the involvement of any proprioceptive feedback mechanisms, and that those musculo-skeletal systems are acting in a critically damped manner, enabling them to react quickly without unnecessary oscillations. We also found that during the late swing, the activity of the posterior biceps/semitendinosus (PB/ST) muscles causes the joints to be unstable. In addition, vastus lateralis (VL), tibialis anterior (TA) and sartorius (SAT) muscle-joint systems were found to be unstable during the late stance phase, and we conclude that those muscles require neuronal feedback to maintain stable stepping, especially during late swing and late stance phases. Moreover, we could see a clear distinction in the pole distribution (along the step cycle) for the systems related to the ankle joint from that of the other two joints, hip or knee. A similar pattern, i.e., a pattern in which the poles were scattered over the s-plane with no clear clustering according to the phase of the leg position, could be seen in the systems related to soleus (SOL) and TA muscles which would indicate that these muscles depend on neural control mechanisms, which may involve supraspinal structures, over the whole step cycle.     

Factors underlying the perturbation resistance of the trunk in the first part of a lifting movement

In the first part of lifting movements, the trunk movement is surprisingly resistant to perturbations. This study examined which factors contribute to this perturbation resistance of the trunk during lifting. Three possible mechanisms were studied: force-length-velocity characteristics of muscles, the momentum of the trunk as well as the effect of passive extending of the elbows. A forward dynamics modelling and simulation approach was adopted with two different input signals: (1) stimulation of Hill-type muscles versus (2) net joint moments. Experimental data collected during an unperturbed lifting movement were used as a reference, which a simulated lifting movement had to resemble. Subsequently, the simulated lifting movement was perturbed by applying 10 kg extra mass at the wrist (both before and after lift-off and with/without a fixed elbow), without modifying the input signals. The momentum of the trunk appeared to be insufficient to explain the perturbation resistance of trunk movements as found experimentally. In addition to the momentum of the trunk, the force-length-velocity characteristics of the muscles are necessary to account for the observed perturbation resistance. Initial extension of the elbow due to the mass perturbation delayed the propagation of the load to the shoulder. However, this delay is reduced due to the impedance at the elbow provided by the characteristics of muscles spanning the elbow. So, the force-length-velocity characteristics of the muscles spanning the elbow joint increase the perturbation at the trunk.     

Is the trunk movement more perturbed after an asymmetric than after a symmetric perturbation during lifting?

Low back injury is associated with sudden movements and loading. Trunk motion after sudden loading depends on the stability of the spine prior to loading and on the trunk muscle activity in response to the loading. Both factors are not axis-symmetric. Therefore, it was hypothesized that the effects on trunk dynamics would be larger after an asymmetric than after a symmetric perturbation. Ten subjects lifted a crate in which, prior to lifting, a mass was displaced to the front or to the side without the subjects being aware of this. Crate and subject movements, crate reaction forces and muscle activity were recorded. From this, the stability prior to the perturbation was estimated, and the trunk angular kinematics and moments at the lumbo-sacral joint were calculated. Both perturbations only minimally affected the trunk kinematics, although the stability of the spine prior to the lifting movement was higher in the sagittal plane than in the frontal plane. In both conditions the stability appeared to be sufficient to absorb the applied perturbation.     

Analysis of a muscular control system in human movements

In this work, we have studied a muscular control system under experimental conditions for analyzing the dynamic behavior of individual muscles and theoretical considerations for elucidating its control strategy. Movement of human limbs is achieved by joint torques and each torque is specified as the sum of torques generated by muscle forces. The behavior of individual muscles is controlled by the neural input which is estimated by means of an electromyogram (EMG). In this study, the EMGs for a flexor and an extensor are measured in elbow joint movements and the dynamic behavior of individual muscles is analyzed. As a result, it is verified that both a flexor and an extensor are activated throughout the entire movement and that the activation of muscles is controlled above a specific limit independent of the hand-held load. Subsequently, a system model for simulating elbow joint movements is developed which includes the muscle dynamic relationship between the neural input and the isometric force. The minimum limit of muscle activation that has been confirmed in experiments is provided as a constraint of the neural input and the criterion is defined by a derivative of the isometric force of individual muscles. The optimal trajectories formulated under these conditions are quantitatively compared with the experimentally observed trajectories, and the control strategy of a muscular control system is studied. Finally, a muscular control system in multi-joint arm movements is discussed with regard to the comparative analysis between observed and optimal trajectories. Received: 7 April 1999 / Accepted in revised form: 27 July 1999     

Plane of vertebral movement eliciting muscle lengthening history in the low back influences the decrease in muscle spindle responsiveness of the cat

Proprioceptive feedback is thought to play a significant role in controlling both lumbopelvic and intervertebral orientations. In the lumbar spine, a vertebra's positional history along the dorsal-ventral axis has been shown to alter the position, movement, and velocity sensitivity of muscle spindles in the multifidus and longissimus muscles. These effects appear due to muscle history. Because spinal motion segments have up to 6 degrees of freedom for movement, we were interested in whether the axis along which the history is applied differentially affects paraspinal muscle spindles. We tested the null hypothesis that the loading axis, which creates a vertebra's positional history, has no effect on a lumbar muscle spindle's subsequent response to vertebral position or movement. Identical displacements were applied along three orthogonal axes directly at the L(6) spinous process using a feedback motor system under displacement control. Single-unit nerve activity was recorded from 60 muscle spindle afferents in teased filaments from L(6) dorsal rootlets innervating intact longissimus or multifidus muscles of deeply anesthetized cats. Muscle lengthening histories along the caudal-cranial and dorsal-ventral axis, compared with the left-right axis, produced significantly greater reductions in spindle responses to vertebral position and movement. The spinal anatomy suggested that the effect of a lengthening history is greatest when that history had occurred along an axis lying within the anatomical plane of the facet joint. Speculation is made that the interaction between normal spinal mechanics and the inherent thixotropic property of muscle spindles poses a challenge for feedback and feedforward motor control of the lumbar spine.     

Signaling of Ankle Joint Position by Receptors in Different Muscles

Plots were made of multiunit activity versus ankle joint position for receptors in each of the 12 muscles crossing the cat ankle joint, except peroneus tertius, by recording from populations of afferent fibers in muscle nerves. The discharge was measured 15 or 30 sec after terminating the movements that altered the position of the joint. These recordings were dominated by large-spike activity that would be expected to originate mainly from primary spindle endings. Seven of the 12 muscles also cross other joints. Their responses at a given ankle joint position were so altered by changes in the position of the knee or toe joints that they could not reliably signal the position of the ankle joint. As judged from multiunit recording, receptors in each of the five muscles specific to the ankle joint were influenced by more than one axis of ankle joint displacement.

Single-unit recording from dorsal root filaments was used to determine whether primary or secondary spindle receptors in soleus and tibialis anterior could selectively signal one axis of ankle joint rotation. Individual soleus receptors were tested both on the flexion extension axis and with a combined adduction–eversion movement.

For 38 of the 70 soleus receptors examined (54%), firm adduction–eversion produced a level of activity greater than that caused by 10° of flexion, and for 77% the level of activity was greater than that caused by 5° of flexion. For 168 of the 184 tibialis anterior receptors studied (91%), firm abduction inversion produced a level of activity greater than that caused by 10° of extension. Thus few receptors were found that responded exclusively to one axis of rotation.

One way in which the position of the ankle joint could be specified in the face of multiaxial receptor activity is by examining the receptor discharge from more than one muscle. A suggestion for how the nervous system might do this is given in the discussion.     

Evolution of locomotion in arachnida: The hydraulic pressure pump of the giant whipscorpion,Mastigoproctus Giganteus (Uropygi)

Many arachnids lack extensor muscles at the femoropatellar (knee) joint of their legs and extend this joint with hydraulic pressure during locomotion. Pressure is generated through compression of the prosoma, but there is disagreement about which muscles are involved in this process. Many arachhnologists consider contraction of the musculi laterales, a group of modified extrinsic leg muscles, as the cause of high prosomal pressure and regard hydraulic extension as a derived feature. However, integration of results from phylogenetic and comparative anatomical studies supports the view that hydraulic extension is primitive in Arachnida and that fluid pressure is generated by contraction of endosternal suspensor muscles. The functional predictions of the musculi laterales and endosternite hypotheses were tested by measuring muscle activity and prosomal pressure during unrestrained locomotion in a primitively “extensorless” arachnid, the giant whipscorpion. The results corroborate the endosternite model and refute the musculi laterales model. Changes in the prosomal pressure baseline were correlated with changes in endosternal muscle activity, while the musculi laterales fired in a step-coupled pattern of discrete bursts that appeared to be incapable of generating the pressure observed during locomotion. Step-coupled fluctuations in prosomal pressure were observed but were apparently caused by rapid flexing of the femoropatellar joints of the fourth leg pair rather than contraction of the musculi laterales.     

Muscle contraction history: modified Hill versus an exponential decay model

In recent years, it has been recognised that improvements to classic models of muscle mechanical behaviour are often necessary for properly modelling co-ordinated multi-joint actions. In this respect, the purpose of the present study was to improve on modelling stretch-induced force enhancement and shortening-induced force depression of muscle contraction. For this purpose, two models were used: a modified Hill model and a model based loosely on mechano-chemistry of the cross-bridge cycle (exponential decay model). The models were compared with a classic Hill model and experimental data. Parameter values were based, as much as possible, on experimental findings in the literature, and tested with new experiments on the gastrocnemius of the rat. Both models describe many features of slow-ramp movements well during short contractions (300–500 ms), but long-duration behaviour is described only partly. The exponential decay model does not incorporate a force–velocity curve. Therefore, its good performance indicates that the status of the classic force–velocity characteristic may have to be reconsidered. Like movement-induced force depression and enhancement, it seems a particular manifestation of time-dependent force behaviour of muscle, rather than a fundamental property of muscle (like the length–tension curve). It is argued that a combination of the exponential decay model (or other models based on the mechano-chemistry of contraction) and structurally based models may be fruitful in explaining this time-dependent contraction behaviour. Furthermore, not in the least because of its relative simplicity, the exponential decay model may prove more suitable for modelling multi-joint movements than the Hill model. Received: 19 March 1999 / Accepted in revised form: 9 June 2000