Mechanical energy expenditure on human movement and the anthropomorphic mechanism]

Mechanical energy expenditures of the man and anthropomorphic locomotion machine during movement are compared theoretically. Sources of the mechanical energy affecting movement of human's lower extremity are modelled by 8 muscles, 3 of which are the two-joint muscles. The model of the lower extremity of anthropomorphic locomotion machine is moved by joint moments. It was shown that in the same movement the model of the human lower extremity can spend less mechanical energy than that of the model of the anthropomorphic locomotion machine. It is caused by the presence of two-joint muscles in the first model. Such an economy of mechanical energy expenditures realized by the two-joint muscle is possible at simultaneous execution of three conditions: 1) signs of the muscle powers, which are produced by that muscle at both joints, are opposite; 2) moments produced by that muscle at each of both joints have the same direction with the joint moments at these joints; 3) one-joint antagonistic muscles are not active. An expression which makes it possible to estimate the mechanical energy savings by the two-joint muscles during humans' movement was developed.     

Theoretical considerations on cocontraction of sets of agonistic and antagonistic muscles

It is well known that static, non-linear minimization of the sum of the stress in muscles to a certain power cannot predict cocontraction of pairs of one-joint antagonistic muscles. In this report, we prove that for a single joint either all agonistic muscles cocontract or all are silent. For two-joint muscles, we show that lengthening and shortening of muscles corresponds closely to zero force and non-zero force states, respectively. This gives a new physiological interpretation of situations in which cocontraction of pairs of two-joint antagonistic muscles is predicted by these static non-linear optimization approaches.     

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).     

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.     

Muscle mechanical work requirements during normal walking: the energetic cost of raising the body's center-of-mass is significant

Inverted pendulum models of walking predict that little muscle work is required for the exchange of body potential and kinetic energy in single-limb support. External power during walking (product of the measured ground reaction force and body center-of-mass (COM) velocity) is often analyzed to deduce net work output or mechanical energetic cost by muscles. Based on external power analyses and inverted pendulum theory, it has been suggested that a primary mechanical energetic cost may be associated with the mechanical work required to redirect the COM motion at the step-to-step transition. However, these models do not capture the multi-muscle, multi-segmental properties of walking, co-excitation of muscles to coordinate segmental energetic flow, and simultaneous production of positive and negative muscle work. In this study, a muscle-actuated forward dynamic simulation of walking was used to assess whether: (1). potential and kinetic energy of the body are exchanged with little muscle work; (2). external mechanical power can estimate the mechanical energetic cost for muscles; and (3.) the net work output and the mechanical energetic cost for muscles occurs mostly in double support. We found that the net work output by muscles cannot be estimated from external power and was the highest when the COM moved upward in early single-limb support even though kinetic and potential energy were exchanged, and muscle mechanical (and most likely metabolic) energetic cost is dominated not only by the need to redirect the COM in double support but also by the need to raise the COM in single support.     

Functional differentiation of fore- and hindlimb muscles inMacaca fuscata determined on the basis of enzymatic activities

When the functional differentiation of 83 kinds of limb and trunk muscles ofMacaca fuscata was investigated on the basis of the activities of two glycolytic enzymes [lactate dehydrogenase (LDH) and aldolase] and one oxidative enzyme [succinate dehydrogenase (SDH)], the forelimb rather than the hindlimb muscles proved have higher oxidative activities. These results indicated that, inMacaca fuscata, the forelimb muscles have a higher resistance to fatigue, and that the hindlimb muscles have a higher tetanic tension on the basis of the relationships between enzymatic activities and functional properties of the muscle fiber types. These findings were interpreted in relation to the fact thatMacaca fuscata is a quadrupedal primate with arboreal habits, as compared with nonprimate terrestrial quadrupeds. The two-joint muscles and the superficial muscles contract more rapidly than do the other muscles in the hindlimb, thereby suggesting that both types of muscles readily adapt to quick movement.     

An EMG-driven model applied for predicting metabolic energy consumption during movement

The relationship between mechanical work and metabolic energy cost during movement is not yet clear. Many studies demonstrated the utility of forward-dynamic musculoskeletal models combined with experimental data to address such question. The aim of this study was to evaluate the applicability of a muscle energy expenditure model at whole body level, using an EMG-driven approach.Four participants performed a 5-min squat exercise on unilateral leg press at two different frequencies and two load levels. Data collected were kinematics, EMG, forces and moments under the foot and gas-exchange data. This same task was simulated using a musculoskeletal model, which took EMG and kinematics as inputs and gave muscle forces and muscle energetics as outputs. Model parameters were taken from literature, but maximal isometric muscle force was optimized in order to match predicted joint moments with measured ones. Energy rates predicted by the model were compared with energy consumption measured by the gas-exchange data.Model results on metabolic energy consumption were close to the values obtained through indirect calorimetry. At the higher frequency level, the model underestimated measured energy consumption. This underestimation can be explained with an increase in energy consumption of the non-muscular mass with movement velocity.In conclusion, results obtained in comparing model predictions with experimental data were promising. More research is needed to evaluate this way of computing mechanical and metabolic work.     

Elastic energy savings and active energy cost in a simple model of running

The energetic economy of running benefits from tendon and other tissues that store and return elastic energy, thus saving muscles from costly mechanical work. The classic “Spring-mass” computational model successfully explains the forces, displacements and mechanical power of running, as the outcome of dynamical interactions between the body center of mass and a purely elastic spring for the leg. However, the Spring-mass model does not include active muscles and cannot explain the metabolic energy cost of running, whether on level ground or on a slope. Here we add explicit actuation and dissipation to the Spring-mass model, and show how they explain substantial active (and thus costly) work during human running, and much of the associated energetic cost. Dissipation is modeled as modest energy losses (5% of total mechanical energy for running at 3 m s^-1) from hysteresis and foot-ground collisions, that must be restored by active work each step. Even with substantial elastic energy return (59% of positive work, comparable to empirical observations), the active work could account for most of the metabolic cost of human running (about 68%, assuming human-like muscle efficiency). We also introduce a previously unappreciated energetic cost for rapid production of force, that helps explain the relatively smooth ground reaction forces of running, and why muscles might also actively perform negative work. With both work and rapid force costs, the model reproduces the energetics of human running at a range of speeds on level ground and on slopes. Although elastic return is key to energy savings, there are still losses that require restorative muscle work, which can cost substantial energy during running.     

Muscle energy transfer during a given wave-like movement of human body segments

The paper deals with a movement of two voluntary segments fixed in a joint and connected by a muscle in a multi-segment biomechanical system of human body. The muscle model is a four-element mechanical system. The mechanical movement energy brought into the "segments-muscle" system from the segments preceding the next ones is studied. The movement in which the total multi-segment system of the human body participates is described by the wave equation. Conditions concerning applying active muscle efforts and correlating velocities of muscle ends movement which provide the maximal value of transferred energy have been found. It is shown that the use of "artificial muscles" type devices promotes activization of energy transfer processes between segments.     

Resolving Shifting Patterns of Muscle Energy Use in Swimming Fish

Muscle metabolism dominates the energy costs of locomotion. Although in vivo measures of muscle strain, activity and force can indicate mechanical function, similar muscle-level measures of energy use are challenging to obtain. Without this information locomotor systems are essentially a black box in terms of the distribution of metabolic energy. Although in situ measurements of muscle metabolism are not practical in multiple muscles, the rate of blood flow to skeletal muscle tissue can be used as a proxy for aerobic metabolism, allowing the cost of particular muscle functions to be estimated. Axial, undulatory swimming is one of the most common modes of vertebrate locomotion. In fish, segmented myotomal muscles are the primary power source, driving undulations of the body axis that transfer momentum to the water. Multiple fins and the associated fin muscles also contribute to thrust production, and stabilization and control of the swimming trajectory. We have used blood flow tracers in swimming rainbow trout (Oncorhynchus mykiss) to estimate the regional distribution of energy use across the myotomal and fin muscle groups to reveal the functional distribution of metabolic energy use within a swimming animal for the first time. Energy use by the myotomal muscle increased with speed to meet thrust requirements, particularly in posterior myotomes where muscle power outputs are greatest. At low speeds, there was high fin muscle energy use, consistent with active stability control. As speed increased, and fins were adducted, overall fin muscle energy use declined, except in the caudal fin muscles where active fin stiffening is required to maintain power transfer to the wake. The present data were obtained under steady-state conditions which rarely apply in natural, physical environments. This approach also has potential to reveal the mechanical factors that underlie changes in locomotor cost associated with movement through unsteady flow regimes.     

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.     

Avian Jaw Function: Adaptation of the Seven–Muscle System and a Review

Avian jaw function is the most interesting part of the feeding apparatus, and essential in the life of birds. The usual seven jaw muscles in birds are highly adapted for diverse food-getting devices through muscular modifications as well as changes in kinesis of the skeletal components of the skull. In the first part I have described from an introspection of my earlier works, the functional morphology of the seven jaw muscles in different birds in four functional groups such as, adductors of the lower jaw, depressor of the lower jaw, protractors of the upper jaw and retractors-cum-adductors of the upper and lower jaws. Emphasis has been laid on the differential force production by these muscles, depending on the nature of their connective tissue attachments on the skeletal parts and changes in the kinesis of the skeletal parts. The contraction of the muscles and movements of the skeletal parts are rhythmically synchronized in such a way that their concerted action performs adaptively in different feeding adaptations. The differential force production by the one-joint and two-joint muscles in terms of ‘torque’ analysis is important in jaw kinesis. The second part of the text is a historical review of some notable works centred around the avian jaw muscles, jaw kinesis, tongue muscles, synchronization with the movements of the tongue apparatus and adaptational as well as evolutionary significance of the feeding apparatus in different feeding strategies.     

Minimizing center of mass vertical movement increases metabolic cost in walking.

A human walker vaults up and over each stance limb like an inverted pendulum. This similarity suggests that the vertical motion of a walker's center of mass reduces metabolic cost by providing a mechanism for pendulum-like mechanical energy exchange. Alternatively, some researchers have hypothesized that minimizing vertical movements of the center of mass during walking minimizes the metabolic cost, and this view remains prevalent in clinical gait analysis. We examined the relationship between vertical movement and metabolic cost by having human subjects walk normally and with minimal center of mass vertical movement ("flat-trajectory walking"). In flat-trajectory walking, subjects reduced center of mass vertical displacement by an average of 69% (P = 0.0001) but consumed approximately twice as much metabolic energy over a range of speeds (0.7-1.8 m/s) (P = 0.0001). In flat-trajectory walking, passive pendulum-like mechanical energy exchange provided only a small portion of the energy required to accelerate the center of mass because gravitational potential energy fluctuated minimally. Thus, despite the smaller vertical movements in flat-trajectory walking, the net external mechanical work needed to move the center of mass was similar in both types of walking (P = 0.73). Subjects walked with more flexed stance limbs in flat-trajectory walking (P < 0.001), and the resultant increase in stance limb force generation likely helped cause the doubling in metabolic cost compared with normal walking. Regardless of the cause, these findings clearly demonstrate that human walkers consume substantially more metabolic energy when they minimize vertical motion.     

A comparison of muscular mechanical energy expenditure and internal work in cycling

The hypothesis that the sum of the absolute changes in mechanical energy (internal work) is correlated with the muscular mechanical energy expenditure (MMEE) was tested using two elliptical chainrings, one that reduced and one that increased the internal work (compared to circular). Upper and lower bounds were put on the extra MMEE (work done by net joint torques in excess of the external work) with respect to the effect of intercompensation between joint torques due to biarticular muscles. This was done by having two measures of MMEE, one that allowed no intercompensation and one that allowed complete intercompensation between joints spanned by biarticular muscles. Energy analysis showed no correlation between internal work and the two measures of MMEE. When compared to circular, the chainring that reduced internal work increased MMEE, and phases of increased crank velocity associated with the elliptical shape resulted in increased power absorbed by the upstroke leg as it was accelerated against gravity. The resulting negative work necessitated additional positive work. Thus, the hypothesis that the internal work is correlated with MMEE was found to be invalid, and the total mechanical work done cannot be estimated by summing the internal and external work. Changes in the dynamics of cycling caused by a non-circular chainring may affect performance and must be considered during the non-circular chainring design process.     

Myoanatomy of the marine tardigrade Halobiotus crispae (Eutardigrada: Hypsibiidae)

The muscular architecture of Halobiotus crispae (Eutardigrada: Hypsibiidae) was examined by means of fluorescent‐coupled phalloidin in combination with confocal laser scanning microscopy and computer‐aided three‐dimensional reconstruction, in addition to light microscopy (Nomarski), scanning electron microscopy, and transmission electron microscopy (TEM). The somatic musculature of H. crispae is composed of structurally independent muscle fibers, which can be divided into a dorsal, ventral, dorsoventral, and a lateral musculature. Moreover, a distinct leg musculature is found. The number and arrangement of muscles differ in each leg. Noticeably, the fourth leg contains much fewer muscles when compared with the other legs. Buccopharyngeal musculature (myoepithelial muscles), intestinal musculature, and cloacal musculature comprise the animal's visceral musculature. TEM of stylet and leg musculature revealed ultrastructural similarities between these two muscle groups. Furthermore, microtubules are found in the epidermal cells of both leg and stylet muscle attachments. This would indicate that the stylet and stylet glands are homologues to the claw and claw glands, respectively. When comparing with previously published data on both heterotardigrade and eutardigrade species, it becomes obvious that eutardigrades possess very similar numbers and arrangement of muscles, yet differ in a number of significant details of their myoanatomy. This study establishes a morphological framework for the use of muscular architecture in elucidating tardigrade phylogeny. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Mechanical energy and power flow analysis of wheelchair use with different camber settings

It has been suggested that minimisation of energy cost is one of the primary determinants of wheelchair designs. Wheel camber is one important parameter related to wheelchair design and its angle may affect usability during manual propulsion. However, there is little available literature addressing the effect of wheel camber on the mechanical energy or power flow involved in manual wheelchair propulsion. Twelve normal subjects (mean age, 22.3 years; SD, 1.6 years) participated in this study. A video-tracking system and an instrumented wheel were used to collect 3D kinematic and kinetic data. Wheel camber of 0° and 15° was chosen to examine the difference between mechanical power and power flow of the upper extremity during manual wheelchair propulsion. The work calculated from power flow and the discrepancy between the mechanical work and power flow work of upper extremity had significantly greater values with increased camber. The upper arm had a larger active muscle power compared with that in the forearm and hand segments. While propelling the increased camber, the magnitude of both the proximal and distal joint power and proximal muscle power was increased in all three segments. While the propelling wheel with camber not only needs a greater energy cost but also there is greater energy loss.     

The minimization of muscular energy expenditure during inspiration in linear models of the respiratory system

The consequences of requiring a general linear model of respiratory mechanics to inspire a fixed volume in a fixed time in a way that minimizes various measures of muscular energy expenditure are examined. For such a model no volume profile minimizes the time-integral of the applied pressure developed by the respiratory muscles, although this integral is not independent of the profile. Minimizing the mechanical work done by the respiratory muscles, on the other hand, requires that the inspiratory flow be constant. These results support the hypothesis that neither the pressure integral nor the mechanical work are individually minimized over an inspiration by an animal or man at rest. Minimization of a weighted sum of the pressure integral and the work done may be a more physiologically reasonable criterion by which the respiratory muscles direct inspiration. This combined cost function predicts a non-constant optimum velocity profile.