Mechanical energy costs of human movement: an approach to evaluating the transfer possibilities of two-joint muscles

Different methods of calculating the mechanical energy cost of a movement presented in the literature can give results differing by an order of magnitude. The assumptions made concerning the transfer of energy between different parts of the body are part of the problem. This investigation assesses the role of transfer in energy saving and specifically, the possibility of two-joint muscles reducing the mechanical energy cost of a movement compared to a system having one-joint muscles only. An algorithm was developed which recruited one-joint or both one- and two-joint muscles to supply the net joint moments. The work performed under these two conditions was then compared. It was found that activation of both one- and two-joint musculature reduced the mechanical work cost during walking by between 7 and 29% over that required by single-joint musculature alone. This investigation supports suggestions in the literature that one of the functions of two-joint musculature is to reduce the mechanical energy cost and probably the metabolic cost of movement.     

The effects of sloped surfaces on locomotion: backward walking as a perturbation

The purpose of this study was to examine lower extremity kinetics and muscle activity during backward slope walking to clarify the relationship between joint moments and powers and muscle activity patterns observed in forward slope walking. Nine healthy volunteers walked backward on an instrumented ramp at three grades (-39% (-21 degrees ), 0% (level), +39% (+21 degrees )). EMG activity was recorded from major lower extremity muscles. Joint kinetics were obtained from kinematic and force platform data. The knee joint moment and power generation increased significantly during upslope walking; hip joint moment and power absorption increased significantly during downslope walking. When compared to data from forward slope walking, these backward walking data suggest that power requirements of a task dictate the muscle activity pattern needed to accomplish that movement. During downslope walking tasks, power absorption increased and changes in muscle activity patterns were directly related to the changes in the joint moment patterns. In contrast, during upslope walking tasks, power generation increased and changes in the muscle activity were related to the changes in the joint moments only at the 'primary' joint; at adjacent joints the changes in muscle activity were unrelated to the joint moment pattern. The 'paradoxical' changes in the muscle activity at the adjacent joints are possibly related to the activation of biarticular muscles required by the increased power generation at the primary joint. In total, these data suggest that changing power requirements at a joint impact the control of muscle activity at that and adjacent joints.     

Dimensions and moment arms of the hind- and forelimb muscles of common chimpanzees (Pan troglodytes).

This paper supplies quantitative data on the hind- and forelimb musculature of common chimpanzees (Pan troglodytes) and calculates maximum joint moments of force as a contribution to a better understanding of the differences between chimpanzee and human locomotion. We dissected three chimpanzees, and recorded muscle mass, fascicle length, and physiological cross-sectional area (PCSA). We also obtained flexion/extension moment arms of the major muscles about the limb joints. We find that in the hindlimb, chimpanzees possess longer fascicles in most muscles but smaller PCSAs than are predicted for humans of equal body mass, suggesting that the adaptive emphasis in chimpanzees is on joint mobility at the expense of tension production. In common chimpanzee bipedalism, both hips and knees are significantly more flexed than in humans, necessitating muscles capable of exerting larger moments at the joints for the same ground force. However, we find that when subject to the same stresses, chimpanzee hindlimb muscles provide far smaller moments at the joints than humans, particularly the quadriceps and plantar flexors. In contrast, all forelimb muscle masses, fascicle lengths, and PCSAs are smaller in humans than in chimpanzees, reflecting the use of the forelimbs in chimpanzee, but not human, locomotion. When subject to the same stresses, chimpanzee forelimb muscles provide larger moments at the joints than humans, presumably because of the demands on the forelimbs during locomotion. These differences in muscle architecture and function help to explain why chimpanzees are restricted in their ability to walk, and particularly to run bipedally.     

Segment interactions within the swing leg during unloaded and loaded running

In this study, designed to determine the effect of lower extremity inertia manipulation on joint kinetics and segment energetics during the swing phase, 15 male distance runners were filmed as they performed treadmill running (3.35 m s-1) under five load conditions: no added load and loads of 0.25 kg and 0.50 kg added to each thigh or each foot. Results of this study demonstrated that the energetics of the lower extremity movements during the swing phase of the running cycle were dominated by mechanical energy transfers between adjacent segments attributed to the joint reaction forces, which acted to redistribute mechanical energy within the system. These contributions were considerably greater than those of the net joint moments, which primarily reflected muscular generation and dissipation of mechanical energy. Lower extremity loading caused little change in the movement pattern of the swing leg. However, increases in the joint reaction forces and net moments and in the amount of work done and the energy transfer attributed to the reaction forces and moments were observed, but were limited to the joints proximal to the location of the added load. These results were consistent with the increased aerobic demand associated with increases in lower extremity inertia that have been reported elsewhere and also have implications for the manner in which the neuromuscular system controls the motion of the legs during running.     

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

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

Peculiarities of Activation of Muscles of the Shoulder Belt in Voluntary Two-Joint Movements of the Upper Limb

We studied coordination of central motor commands (СMCs) coming to muscles of the shoulder and shoulder belt in the course of single-joint and two-joint movements including flexion and extension of the elbow and shoulder joints. Characteristics of rectified and averaged EMGs recorded from a few muscles of the upper limb were considered correlates of the CMC parameters. Special attention was paid to coordination of CMCs coming to two-joint muscles that are able to function as common flexors (m. biceps brachii, caput breve, BBcb) and common extensors (m. triceps brachii, caput longum, TBcl) of the elbow and shoulder joints. Upper limb movements used in the tests included planar shifts of the arm from one spatial point to another resulting from either simultaneous changes in the angles of the shoulder and elbow joints or isolated sequential (two-stage) changes in these joint angles. As was found, shoulder muscles providing movements of the elbow with changes in the angle of the elbow joint, i.e., BBcb and TBcl, were also intensely involved in the performance of single-joint movements in the shoulder joint. The CMCs coming to two-joint muscles in the course of two-joint movements appeared, in the first approximation, as sums of the commands received by these muscles in the course of corresponding single-joint movements in the elbow and shoulder joints. Therefore, if we interpret the isolated forearm movement performed due to a change in the angle of the elbow joint as the main motor event, while the shoulder movement is considered the accessory one, we can conclude that realization of a two-joint movement of the upper-limb distal part is based on superposition of CMCs related to basic movements (main and accessory). Neirofiziologiya/Neurophysiology, Vol. 41, No. 1, pp. 48–56, January–February, 2009.     

A quantitative assessment of the "tendon" action of two-joint muscles]

Two-joint muscles are able to transmit mechanical energy between the links of the body having no common joint ("tendon action" of the muscles). It is proposed to calculate difference between control moment power in a joint and the sum of powers developed by all muscles serving this joint in order to determine the direction and rate of mechanical energy transfer through the two-joint muscles. It was shown that in the shock-absorbing phase of support in running two-joint muscles the energy transfers from distal to proximal links (from foot to thigh, and from shank to pelvis), in take-off phase-from proximal links to distal ones (from pelvis to shank, and from thigh to foot).     

Peculiarities of Activation of the Upper Limb Muscles in Realization of Two-Joint Movements in Humans

In tests on humans, we recorded EMG activity from the muscles flexing and extending the forearm and shoulder in the course of realization of sequential single-joint and simultaneous two-joint movements of the upper limb. As was shown, the shoulder muscles m. biceps brachii and m. triceps brachii are involved in flexion/extension of both elbow and shoulder joints. Central commands sent to the above muscles in the course of a two-joint movement could be considered a superposition of the central commands coming to the same muscles in realization of the corresponding sequential single-joint movements with the same changes in the angles of the elbow and shoulder joints. External loadings applied in the direction of extension of the elbow and shoulder joints induced, in general, similar changes in coordination of the activity of muscles moving the forearm and shoulder under conditions of both single-joint and two-joint movements. These facts allow us to suppose that coordination of the muscle activity in two-joint movements depends to a greater extent on the forces influencing limb links than on the mode of realization of the movements (two sequential single-joint movements vs a two-joint movement corresponding to the above motor events).     

The evolution of hindlimb tendons and muscles on the line to crown-group birds

The anatomy and functions of muscle-tendon complexes and their bony attachments in birds and their outgroups show how the major pelvic limb muscle groups evolved. Fossils reveal that most changes evolved after the divergence of archosaurs in the Triassic, particularly in the dinosaurian precursors to birds. Three-dimensional limb control became concentrated at the hip joint; more distal joints and muscles were restricted to flexion or extension early in dinosaur evolution. Hip extensors expanded even though the primary femoral retractor M. caudofemoralis longus was reduced. Hip flexors and two-joint "hamstring" muscles were simplified to a few large heads. Knee extensors increased their sizes and moment arms early in bipedal dinosaurs, but the patella and cranial cnemial crest evolved later in birds. Lower limb muscles expanded as ossifications such as the hypotarsus increased their moment arms. The ossification of lower limb tendons, particularly in extensors, is a recent novelty of birds. Muscles and tendons that develop large forces, stresses, and moments to stabilize or move the limbs became increasingly prominent on the line to birds. Locomotion evolved in a stepwise pattern that only recently produced the derived limb control mechanisms of crown-group birds, such as the strongly flexed hip and knee joints.     

Neuromusculoskeletal computer modeling and simulation of upright, straight-legged, bipedal locomotion of Australopithecus afarensis (A.L. 288-1)

The skeleton of Australopithecus afarensis (A.L. 288-1, better known as "Lucy") is by far the most complete record of locomotor morphology of early hominids currently available. Even though researchers agree that the postcranial skeleton of Lucy shows morphological features indicative of bipedality, only a few studies have investigated Lucy's bipedal locomotion itself. Lucy's energy expenditure during locomotion has been the topic of much speculation, but has not been investigated, except for several estimates derived from experimental data collected on other animals. To gain further insights into how Lucy may have walked, we generated a full three-dimensional (3D) reconstruction and forward-dynamic simulation of upright bipedal locomotion of this ancient human ancestor. Laser-scanned 3D bone geometries were combined with state-of-the-art neuromusculoskeletal modeling and simulation techniques from computational biomechanics. A detailed full 3D neuromusculoskeletal model was developed that encompassed all major bones, joints (10), and muscles (52) of the lower extremity. A model of muscle force and heat production was used to actuate the musculoskeletal system, and to estimate total energy expenditure during locomotion. Neural activation profiles for each of the 52 muscles that produced a single step of locomotion, while at the same time minimizing the energy consumed per meter traveled, were searched through numerical optimization. The numerical optimization resulted in smooth locomotor kinematics, and the predicted energy expenditure was appropriate for upright bipedal walking in an individual of Lucy's body size.     

Experimentally reduced hip abductor function during walking: Implications for knee joint loads

Hip and knee functions are intimately connected and reduced hip abductor function might play a role in development of knee osteoarthritis (OA) by increasing the external knee adduction moment during walking. The purpose of this study was to test the hypothesis that reduced function of the gluteus medius (GM) muscle would lead to increased external knee adduction moment during level walking in healthy subjects. Reduced GM muscle function was induced experimentally, by means of intramuscular injections of hypertonic saline that produced an intense short-term muscle pain and reduced muscle function. Isotonic saline injections were used as non-painful control. Fifteen healthy subjects performed walking trials at their self-selected walking speed before and immediately after injections, and again after 20 min of rest, to ensure pain recovery. Standard gait analyses were used to calculate three-dimensional trunk and lower extremity joint kinematics and kinetics. Surface electromyography (EMG) of the glutei, quadriceps, and hamstring muscles were also measured. The peak GM EMG activity had temporal concurrence with peaks in frontal plane moments at both hip and knee joints. The EMG activity in the GM muscle was significantly reduced by pain (?39.6%). All other muscles were unaffected. Peaks in the frontal plane hip and knee joint moments were significantly reduced during pain (?6.4% and ?4.2%, respectively). Lateral trunk lean angles and midstance hip joint adduction and knee joint extension angles were reduced by ?1°. Thus, the gait changes were primarily caused by reduced GM function. Walking with impaired GM muscle function due to pain significantly reduced the external knee adduction moment. This study challenge the notion that reduced GM function due to pain would lead to increased loads at the knee joint during level walking.     

Ankle flexor muscles in the cat: Length-active tension and muscle unit properties as related to locomotion

The mechanical properties of the whole muscle and fast-twitch muscle units of the cat hindlimb pretibial flexors have been explored and related to normal locomotion. Tibialis anterior (TA) is parallel-fibered and functionally crosses a single joint, the ankle, whereas extensor digitorum longus (EDL) is pinnate and spans the ankle, knee, metatarsophalangeal and interphalangeal joints. The active tetanic tension of TA remains near its peak value over a range of muscle lengths associated with normal ankle movement. In contrast, the length-tension curve of EDL is sharply peaked. However, normal corollary action of the knee, ankle and metatarsophalangeal joints during stepping minimizes EDL's excursion and maintains it at or near a length optimal for peak tension development. EDL is capable of producing synchronous but sterotyped digit and ankle movements while TA provides for independent ankle flexion at all relevant joint angles. The mechanical properties of 84 TA and 98 EDL fast-twitch muscle units were studied by measuring twitch contraction time (≤45 msec), peak tetanic tension, response to repetitive stimulation, and contractile fatigue resistance during electrical stimulation of single alpha axons, functionally isolated from ventral root filaments. These mechanical properties were essentially similar for both muscles with the exception of mean peak tetanic tension which was 30% lower for TA units (14 gm-wt) than for EDL units (20 gm-wt). A high proportion of units in both muscles demonstrated fatigue resistance which is reflective of the repetitive, phasic demand upon these muscles during locomotion.     

Joit torque and energy patterns in normal gait

Experiments were conducted on normal level gait to determine the synergistic patterns present in the forces causing joint moments and those associated with the generation, absorption and transfer of mechanical energy. The following generalizations can be made about the patterns: (i) During swing phase three forces (gravitational, muscle and knee joint acceleration) are responsible for shank rotation, and are shown to act together during both acceleration and deceleration.—(ii) The patterns of generation, absorption and transfer of mechanical energy at the joints are detailed. These patterns demonstrate inter-segment transfers of energy through the joint centres, and through the muscles, as well as the more recognized generation and absorption by the muscles themselves.—As a result of the complexity shown in these patterns it is cautioned that fundamental relationships that may have been derived from controlled biomechanical experiments (such as horizontal flexion and extension of the forearm) are not likely to apply to more normal movements such as gait.     

Lower extremity control and dynamics during backward angular impulse generation in forward translating tasks

Observation of complex whole body movements suggests that the nervous system coordinates multiple operational subsystems using some type of hierarchical control. When comparing two forward translating tasks performed with and without backward angular impulse, we have learned that both trunk-leg coordination and reaction force-time characteristics are significantly different between tasks. This led us to hypothesize that differences in trunk-leg coordination and reaction force generation would induce between-task differences in the control of the lower extremity joints during impulse generation phase of the tasks. Eight highly skilled performers executed a series of forward jumps with and without backward rotation (reverse somersault and reverse timer, respectively). Sagittal plane kinematics, reaction forces, and electromyograms of lower extremity muscles were acquired during the take-off phase of both tasks. Lower extremity joint kinetics were calculated using inverse dynamics. The results demonstrated between-task differences in the relative angles between the lower extremity segments and the net joint forces/reaction force and the joint angular velocity profiles. Significantly less knee extensor net joint moments and net joint moment work and greater hip extensor net joint moments and net joint moment work were observed during the push interval of the reverse somersault as compared to the reverse timer. Between-task differences in lower extremity joint kinetics were regulated by selectively activating the bi-articular muscles crossing the knee and hip. These results indicate that between-task differences in the control of the center of mass relative to the reaction force alters control and dynamics of the multijoint lower extremity subsystem.     

Identification of passive elastic joint moments in the lower extremities.

Musculotendon actuators produce active and passive moments at the joints they span. Due to the existence of bi-articular muscles, the passive elastic joint moments are influenced by the angular positions of adjacent joints. To obtain quantitative information about this passive elastic coupling between lower limb joints, we examined the passive elastic joint properties of the hip, knee, and ankle joint of ten healthy subjects. Passive elastic joint moments were found to considerably depend on the adjacent joint angles. We present a simple mathematical model that describes these properties on the basis of a double-exponential expression. The model can be implemented in biomechanical models of the lower extremities, which are generally used for the simulation of multi-joint movements such as standing-up, walking, running, or jumping.     

Joint moments and contact forces in the foot during walking

The net force and moment of a joint have been widely used to understand joint disease in the foot. Meanwhile, it does not reflect the physiological forces on muscles and contact surfaces. The objective of the study is to estimate active moments by muscles, passive moments by connective tissues and joint contact forces in the foot joints during walking. Joint kinematics and external forces of ten healthy subjects (all males, 24.7 ± 1.2 years) were acquired during walking. The data were entered into the five-segment musculoskeletal foot model to calculate muscle forces and joint contact forces of the foot joints using an inverse dynamics-based optimization. Joint reaction forces and active, passive and net moments of each joint were calculated from muscle and ligament forces. The maximum joint reaction forces were 8.72, 4.31, 2.65, and 3.41 body weight (BW) for the ankle, Chopart’s, Lisfranc and metatarsophalangeal joints, respectively. Active and passive moments along with net moments were also obtained. The maximum net moments were 8.6, 8.4, 5.4 and 0.8%BW∙HT, respectively. While the trend of net moment was very similar between the four joints, the magnitudes and directions of the active and passive moments varied between joints. The active and passive moments during walking could reveal the roles of muscles and ligaments in each of the foot joints, which was not obvious in the net moment. This method may help narrow down the source of joint problems if applied to clinical studies.     

The integrated function of muscles and tendons during locomotion

The mechanical roles of tendon and muscle contractile elements during locomotion are often considered independently, but functionally they are tightly integrated. Tendons can enhance muscle performance for a wide range of locomotor activities because muscle-tendon units shorten and lengthen at velocities that would be mechanically unfavorable for muscle fibers functioning alone. During activities that require little net mechanical power output, such as steady-speed running, tendons reduce muscular work by storing and recovering cyclic changes in the mechanical energy of the body. Tendon stretch and recoil not only reduces muscular work, but also allows muscle fibers to operate nearly isometrically, where, due to the force-velocity relation, skeletal muscle fibers develop high forces. Elastic energy storage and recovery in tendons may also provide a key mechanism to enable individual muscles to alter their mechanical function, from isometric force-producers during steady speed running to actively shortening power-producers during high-power activities like acceleration or uphill running. Evidence from studies of muscle contraction and limb dynamics in turkeys suggests that during running accelerations work is transferred directly from muscle to tendon as tendon stretch early in the step is powered by muscle shortening. The energy stored in the tendon is later released to help power the increase in energy of the body. These tendon length changes redistribute muscle power, enabling contractile elements to shorten at relatively constant velocities and power outputs, independent of the pattern of flexion/extension at a joint. Tendon elastic energy storage and recovery extends the functional range of muscles by uncoupling the pattern of muscle fiber shortening from the pattern of movement of the body.     

Activation of the Shoulder-Belt and Shoulder Muscles in Two-Joint Arm Movements Performed in Humans with the Action of Opposite Loadings

In tests on four volunteers, we examined coordination of central motor commands (CMCs) controlling slow two-joint movements of the arm within the horizontal plane. Current amplitudes of EMGs recorded from six muscles of the shoulder belt and shoulder and subjected to full-wave rectifying and low-frequency filtration were considered correlates of these commands. In particular, we studied the dependence of coordination of CMCs on the direction of an external force applied to the distal forearm part. As was found, coordination of CMCs significantly depends on the direction of the force flexing the elbow joint. According to our observations, EMGs of definite muscles in the case of performance of a two-joint movement can, in a first approximation, be presented as linear combinations of the EMGs recorded in the course of separate sequential single-joint movements under conditions of shifting the reference point of the hand toward the same point of the operational space as that in the two-joint movement. These data can be interpreted as confirmation of the principle of superposition of elementary CMCs in the performance of complex movements of the extremity.     

Adaptations in the gait pattern with experimental hamstring pain

Pain changes movement but most studies have focused on basic physiological adaptations during non-functional movement tasks. The existing studies on how pain affects lower extremity gross movement biomechanics have primarily involved movements in which the quadriceps is the primary muscle and little attention has been given to how pain in other muscles affects functional movement. The purpose of this study was to investigate the changes in the gait patterns of healthy subjects that occur during experimental muscle pain in the biceps femoris.In a cross-over study design, 14 healthy volunteers underwent EMG assisted 3D gait analyses before, during and after experimental biceps femoris pain induced by intramuscular injections of hypertonic saline. Isotonic saline injections were administered as a non-painful control.The experimental biceps femoris pain led to reductions in hip extensor moments, knee flexor and lateral rotator moments. No changes in lower extremity kinematics and EMG activity in any of the recorded muscles were observed.It is concluded that experimental muscle pain in the biceps femoris leads to changes in the gait pattern in agreement with unloading of the painful muscle. The changes are specific to the painful muscle. The present study provides support to the theory that musculoskeletal pain is a protective signal leading to changes in movement patterns that serve to unload the painful tissue.