The mechanics of head-supported load carriage by Nepalese porters.

In the Everest valley of Nepal, because of the rugged mountain terrain, roads are nothing more than dirt paths and all material must be conveyed on foot. The Nepalese porters routinely carry head-supported loads, which often exceed their body mass, over long distances up and down the steep mountain footpaths. In Africa, women transport their loads economically thanks to an energy-saving gait adaptation. We hypothesized that the Nepalese porters may have developed a corresponding mechanism. To investigate this proposition, we measured the mechanical work done during level walking in Nepalese porters while carrying different loads at several speeds. Our results show that the Nepalese porters do not use an equivalent mechanism as the African women to reduce work. In contrast, the Nepalese porters develop an equal amount of total mechanical work as Western control subjects while carrying loads of 0 to 120% of their body mass at all speeds measured (0.5-1.7 m s-1), making even more impressive their ability to carry loads without any apparent mechanically determined tricks. Nevertheless, our results show that the Nepalese porters have a higher efficiency, at least at slow speeds and high loads.

Biomechanics of locomotion in Asian elephants.

Elephants are the biggest living terrestrial animal, weighing up to five tons and measuring up to three metres at the withers. These exceptional dimensions provide certain advantages (e.g. the mass-specific energetic cost of locomotion is decreased) but also disadvantages (e.g. forces are proportional to body volume while supportive tissue strength depends on their cross-sectional area, which makes elephants relatively more fragile than smaller animals). In order to understand better how body size affects gait mechanics the movement of the centre of mass (COM) of 34 Asian elephants (Elephas maximus) was studied over their entire speed range of 0.4-5.0 m s(-1) with force platforms. The mass-specific mechanical work required to maintain the movements of the COM per unit distance is approximately 0.2 J kg(-1) m(-1) (about 1/3 of the average of other animals ranging in size from a 35 g kangaroo rat to a 70 kg human). At low speeds this work is reduced by a pendulum-like exchange between the kinetic and potential energies of the COM, with a maximum energy exchange of approximately 60% at 1.4 m s(-1). At high speeds, elephants use a bouncing mechanism with little exchange between kinetic and potential energies of the COM, although without an aerial phase. Elephants increase speed while reducing the vertical oscillation of the COM from about 3 cm to 1 cm.

Effect of an increase in gravity on the power output and the rebound of the body in human running.

The effect of an increase in gravity on the mechanics of running has been studied by using a force platform fixed to the floor of an aircraft undergoing flight profiles, resulting in a simulated gravity of 1.3 g. The power spent to maintain the motion of the centre of mass of the body is approximately 1.3 times greater than on Earth, due to a similar increase of both the power spent against gravity and to sustain the forward speed changes. This indicates that the average vertical displacement per unit distance and the average direction of the push are unchanged. The increase in power is mainly due to an increase in step frequency rather than to an increase in the work done at each step. The increase in step frequency in turn is mainly due to a decreased duration of the effective aerial phase (when the vertical force is less than body weight), rather than an increase in the stiffness of the bouncing system. The maximal speed where step frequency can match the resonant frequency of the bouncing system is increased by approximately 5 km h(-1) at 1.3 g. These results suggest a similar running mechanics at higher gravity, maintained at the expense of greater energy expenditure.

Effect of load and speed on the energetic cost of human walking.

It is well established that the energy cost per unit distance traveled is minimal at an intermediate walking speed in humans, defining an energetically optimal walking speed. However, little is known about the optimal walking speed while carrying a load. In this work, we studied the effect of speed and load on the energy expenditure of walking. The O(2) consumption and CO(2) production were measured in ten subjects while standing or walking at different speeds from 0.5 to 1.7 m s(-1) with loads from 0 to 75% of their body mass (M(b)). The loads were carried in typical trekker's backpacks with hip support. Our results show that the mass-specific gross metabolic power increases curvilinearly with speed and is directly proportional to the load at any speed. For all loading conditions, the gross metabolic energy cost (J kg(-1) m(-1)) presents a U-shaped curve with a minimum at around 1.3 m s(-1). At that optimal speed, a load up to 1/4 M(b) seems appropriate for long-distance walks. In addition, the optimal speed for net cost minimization is around 1.06 m s(-1) and is independent of load.

Mechanical work and muscular efficiency in walking children.

The effect of age and body size on the total mechanical work done during walking is studied in children of 3-12 years of age and in adults. The total mechanical work per stride (W tot) is measured as the sum of the external work, W ext (i.e. the work required to move the centre of mass of the body relative to the surroundings), and the internal work, W int (i.e. the work required to move the limbs relative to the centre of mass of the body, W int,k, and the work done by one leg against the other during the double contact period, W int,dc). Above 0.5 m s(-1), both W ext) and W int,k, normalised to body mass and per unit distance (J kg(-1) m(-1)), are greater in children than in adults; these differences are greater the higher the speed and the younger the subject. Both in children and in adults, the normalised W int,dc shows an inverted U-shape curve as a function of speed, attaining a maximum value independent of age but occurring at higher speeds in older subjects. A higher metabolic energy input (J kg(-1) m(-1)) is also observed in children, although in children younger than 6 years of age, the normalised mechanical work increases relatively less than the normalised energy cost of locomotion. This suggests that young children have a lower efficiency of positive muscular work production than adults during walking. Differences in normalised mechanical work, energy cost and efficiency between children and adults disappear after the age of 10.

The double contact phase in walking children.

During walking, when both feet are on the ground (the double contact phase), the legs push against each other, and both positive and negative work are done simultaneously. The work done by one leg on the other (W(int,dc)) is not counted in the classic measurements of the positive muscular work done during walking. Using force platforms, we studied the effect of speed and age (size) on W(int,dc). In adults and in 3-12-year-old children, W(int,dc) (J kg(-1) m(-1)) as a function of speed shows an inverted U-shaped curve, attaining a maximum value that is independent of size but that occurs at higher speeds in larger subjects. Normalising the speed with the Froude number shows that W(int,dc) is maximal at about 0.3 in both children and adults. Differences due to size disappear for the most part when normalised with the Froude number, indicating that these speed-dependent changes are primarily a result of body size changes. At its maximum, W(int,dc) represents more than 40% of W(ext) (the positive work done to move the centre of mass of the body relative to the surroundings) in both children and adults.

Pendular energy transduction within the step in human walking.

During walking, the centre of mass of the body moves like that of a 'square wheel': with each step cycle, some of its kinetic energy, E(k), is converted into gravitational potential energy, E(p), and then back into kinetic energy. To move the centre of mass, the locomotory muscles must supply only the power required to overcome the losses occurring during this energy transduction. African women carry loads of up to 20% of their body weight on the head without increasing their energy expenditure. This occurs as a result of an unexplained, more effective energy transduction between E(k) and E(p) than that of Europeans. In this study we measured the value of the E(k) to E(p) transduction at each instant in time during the step in African women and European subjects during level walking at 3.5-5.5 km h(-1), both unloaded and carrying loads spanning 20-30% of their body weight. A simulation of the changes in E(k) and E(p) during the step by sinusoidal curves was used for comparison. It was found that loading improves the transduction of E(p) to E(k) during the descent of the centre of mass. The improvement is not significant in European subjects, whereas it is highly significant in African women.

Energy transfer during stress relaxation of contracting frog muscle fibres.

1. A contracting muscle resists stretching with a force greater than the force it can exert at a constant length, T(o). If the muscle is kept active at the stretched length, the excess tension disappears, at first rapidly and then more slowly (stress relaxation). The present study is concerned with the first, fast tension decay. In particular, it is still debated if and to what extent the fast tension decay after a ramp stretch involves a conservation of the elastic energy stored during stretching into cross-bridge states of higher chemical energy. 2. Single muscle fibres of Rana temporaria and Rana esculenta were subjected to a short ramp stretch (approximately 15 nm per half-sarcomere at either 1.4 or 0.04 sarcomere lengths s(-1)) on the plateau of the force-length relation at temperatures of 4 and 14 degrees C. Immediately after the end of the stretch, or after discrete time intervals of fixed-end contraction and stress relaxation at the stretched length (Delta t(isom) = 0.5-300 ms), the fibre was released against a force ~T(o). Fibre and sarcomere stiffness during the elastic recoil to T(o) (phase 1) and the subsequent transient shortening against T(o) (phase 2), which is expression of the work enhancement by stretch, were measured after different Delta t(isom) and compared with the corresponding fast tension decay during Delta t(isom). 3. The amplitude of fast tension decay is large after the fast stretch, and small or nil after the slow stretch. Two exponential terms are necessary to fit the fast tension decay after the fast stretch at 4 degrees C, whereas one is sufficient in the other cases. The rate constant of the dominant exponential term (0.1-0.2 ms(-1) at 4 degrees C) increases with temperature with a temperature coefficient (Q(10)) of approximately 3. 4. After fast stretch, the fast tension decay during Delta t(isom) is accompanied in both species and at both temperatures by a corresponding increase in the amplitude of phase 2 shortening against T(o) after Delta t(isom): a maximum of approximately 5 nm per half-sarcomere is attained when the fast tension decay is almost complete, i.e. 30 ms after the stretch at 4 degrees C and 10 ms after the stretch at 14 degrees C. After slow stretch, when fast tension decay is small or nil, the increase in phase 2 shortening is negligible. 5. The increase in phase 2 work during fast tension decay (Delta W(out)) is a constant fraction of the elastic energy simultaneously set free by the recoil of the undamped elastic elements. 6. Delta W(out) is accompanied by a decrease in stiffness, indicating that it is not due to a greater number of cross-bridges. 7. It is concluded that, during the fast tension decay following a fast ramp stretch, a transfer of energy occurs from the undamped elastic elements to damped elements within the sarcomeres by a temperature-dependent mechanism with a dominant rate constant consistent with the theory proposed by A. F. Huxley and R. M. Simmons in 1971.

Mechanical power and efficiency in running children.

The effect of age and body size on the total mechanical power output (Wtot) during running was studied in children of 3-12 years of age and in adults. Wtot was measured as the sum of the power required to move the body's centre-of-mass relative to the surroundings (the "external power", Wext) plus the power required to move the limbs relative to the body's centre-of-mass (the "internal power", Wint). At low and intermediate speeds (less than about 13 km h-1) the higher step frequency used by young children resulted in a decrease of up to 40-50% in the mass-specific external power and an equal increase in the mass-specific internal power relative to adults. Due to this crossed effect, the mass-specific Wtot is nearly independent of age. At high speeds the mass-specific Wtot is 20-30% larger in young children than in adults, due to a greater forward deceleration of the centre-of-mass at each step. The efficiency of positive work production, calculated as the positive mechanical power divided by the net energy consumption rate, appears to be similar in children and adults (i.e. 0.40-0.55).

The energy cost of walking in children.

Size, morphology and motor skills change dramatically during growth and this probably has an effect on the cost of locomotion. In this study, the effects of age and speed on the energy expended while walking were determined during growth. The rate of oxygen consumption and carbon dioxide production were measured in 3- to 12-year-old children and in adults while standing and walking at different speeds from 0.5 m x s(-1) to near their maximum aerobic walking speed. Standing energy expenditure rate decreases with age from 3.42 +/- 0.48 W x kg(-1) (mean +/- SD, n = 6) in the 3- to 4-year-olds to 1.95 +/- 0.22 W x kg(-1) (n = 6) in young adults. At all ages the gross cost of transport has a minimum which decreases from 5.9 J x kg(-1) x m(-1) in 3- to 4-year-olds to 3.6 J x kg(-1) x m(-1) after 10 years of age. The speed at which this minimum occurs increases from 1.2 m x s(-1) to 1.5 m x s(-1) over the same age range. At low and intermediate walking speeds the net cost of transport is similar in children and adults (about 2 J x kg(-1) x m(-1)). In young children walking at their highest speeds the net cost of transport is 70% (3- to 4-year-olds) to 40% (5- to 6-year-olds) greater than in adults.

The role of gravity in human walking: pendular energy exchange, external work and optimal speed.

During walking on Earth, at 1.0 g of gravity, the work done by the muscles to maintain the motion of the centre of mass of the body (W(ext)) is reduced by a pendulum-like exchange between gravitational potential energy and kinetic energy. The weight-specific W(ext) per unit distance attains a minimum of 0.3 J x kg(-1) x m(-1) at about 4.5 km x h(-1) in adults. The effect of a gravity change has been studied during walking on a force platform fixed to the floor of an aircraft undergoing flight profiles which resulted in a simulated gravity of 0.4 and 1.5 times that on Earth. At 0.4 g, such as on Mars, the minimum W(ext) was 0.15 J x kg(-1) x m(-1), half that on Earth and occurred at a slower speed, about 2.5 km x h(-1). The range of walking speeds is about half that on Earth. At 1.5 g, the lowest value of W(ext) was 0.60 J x kg(-1) x m(-1), twice that on Earth; it was nearly constant up to about 4.3 km x h(-1) and then increased with speed. The range of walking speeds is probably greater than that on Earth. A model is presented in which the speed for an optimum exchange between potential and kinetic energy, the 'optimal speed', is predicted by the balance between the forward deceleration due to the lift of the body against gravity and the forward deceleration due to the impact against the ground. In conclusion, over the range studied, gravity increases the work required to walk, but it also increases the range of walking speeds.

Effect of stretching on undamped elasticity in muscle fibres from Rana temporaria.

Muscle stiffness was measured from the undamped elastic recoil taking place when the force attained during ramp stretches of muscle fibres, tetanized on the plateau of the tension-length relation, was suddenly reduced to the isometric value developed before the stretch, T0. Sarcomere elastic recoil was measured on a tendon-free segment of the fibre by means of a striation follower. After small ramp stretches, stiffness increases to a value 1.33x greater than that measured during release from a state of isometric contraction to 0.9 T0. While the relative increase in stiffness is equal to that reported for fibres of Rana esculenta (Piazzesi et al., 1992), the absolute value of stiffness measured during release from isometric contraction is just over half. As stretch amplitude is increased, on the plateau of the force-length relation, stiffness decreases toward the isometric value. This finding shows that the decrease in stiffness with large stretches cannot be due to a decrease in myofilament overlap (as may be the case when stretching occurs on the descending limb of the tension-length relation, Sugi & Tsuchiya, 1988), but must be due to an effect of the ramp stretch per se. For a given stretch amplitude, the after-stretch transient shortening against T0 taking place after the elastic recoil (which is expression of the work enhancement induced by stretching, Cavagna et al., 1986, 1994) is similar in fibres with very different stiffness of their undamped elastic elements. This suggests that this work enhancement is not due to the recoil of damped elastic structures recruited during stretching because of sarcomere length inhomogenity, a condition which would result in a decrease in stiffness (Morgan et al., 1996).

Mechanics and energetics of human locomotion on sand.

Moving about in nature often involves walking or running on a soft yielding substratum such as sand, which has a profound effect on the mechanics and energetics of locomotion. Force platform and cinematographic analyses were used to determine the mechanical work performed by human subjects during walking and running on sand and on a hard surface. Oxygen consumption was used to determine the energetic cost of walking and running under the same conditions. Walking on sand requires 1.6-2.5 times more mechanical work than does walking on a hard surface at the same speed. In contrast, running on sand requires only 1.15 times more mechanical work than does running on a hard surface at the same speed. Walking on sand requires 2.1-2.7 times more energy expenditure than does walking on a hard surface at the same speed; while running on sand requires 1.6 times more energy expenditure than does running on a hard surface. The increase in energy cost is due primarily to two effects: the mechanical work done on the sand, and a decrease in the efficiency of positive work done by the muscles and tendons.

Muscle work enhancement by stretch. Passive visco-elasticity or cross-bridges?

Tetanized frog muscle fibres subjected to ramp stretches on the plateau of the tension-length relation, followed by an isotonic release against a load equal to the maximum isometric tension (T0), exhibit a well defined transient shortening against T0 which was attributed to the release of mechanical energy stored during stretching within the damped element of the cross-bridges. However, this interpretation has recently been challenged, and 'transient shortening against T0' has instead been attributed to elastic elements strained because of non-uniform distribution of lengthening within the fibre volume. The 'excess length change', resulting from the recoil of these elastic elements, was found i) to increase continuously with stretch amplitude up to 50 nm per h.s. with a 100 nm per h.s. strain, ii) to decrease steadily with the decrease in force during stress relaxation after the ramp stretch, and iii) to increase on the descending limb of the tension-length relation where sarcomere inhomogeneity is greater. In contrast, the transient shortening against T0: i) reaches a plateau at 8 nm per half sarcomere after about 50 nm per half sarcomere strain, ii) remains constant during the temperature dependent, fast phase of stress relaxation, when the excess in force above isometric reduces to about one half, iii) also occurs on the ascending limb of the tension-length relation where sarcomere inhomogeneity is drastically reduced. As a consequence of these differences we conclude that transient shortening and 'excess length change' do not "reflect the same underlying process".

Energy-saving gait mechanics with head-supported loads.

In many areas of the world that lack a transportation infrastructure, people routinely carry extraordinary loads supported by their heads, for example the Sherpa of the Himalayas and the women of East Africa. It has previously been shown that African women from the Kikuyu and Luo tribes can carry loads substantially more cheaply than army recruits; however, the mechanism for their economy has remained unknown. Here we investigate, using a force platform, the mechanics of carrying head-supported loads by Kikuyu and Luo women. The weight-specific mechanical work, required to maintain the motion of the common centre of mass of the body and load, decreases with load in the African women, whereas it increases in control subjects. The decrease in work by the African women is a result of a greater conservation of mechanical energy resulting from an improved pendulum-like transfer of energy during each step, back and forth between gravitational potential energy and kinetic energy of the centre of mass.

Moving cheaply: energetics of walking in the African elephant.

Large animals have a much better fuel economy than small ones, both when they rest and when they run. At rest, each gram of tissue of the largest land animal, the African elephant, consumes metabolic energy at 1/20 the rate of a mouse; using existing allometric relationships, we calculate that it should be able to carry 1 g of its tissue (or a load) for 1 km at 1/40 the cost for a mouse. These relationships between energetics and size are so consistent that they have been characterized as biological laws. The elephant has massive legs and lumbers along awkwardly, suggesting that it might expend more energy to move about than other animals. We find, however, that its energetic cost of locomotion is predicted remarkably well by the allometric relationships and is the lowest recorded for any living land animal.

External, internal and total work in human locomotion.

The muscle-tendon work performed during locomotion can, in principle, be measured from the mechanical energy of the centre of mass of the whole body and the kinetic energy due to the movements of the body segments relative to the centre of mass of the body. Problems arise when calculating the muscle-tendon work from increases in mechanical energy, largely in correctly attributing these increases either to energy transfer or to muscle-tendon work. In this study, the kinetic and gravitational potential energy of the centre of mass of the whole human body was measured (using a force platform) simultaneously with calculation of the kinetic and potential energy of the body segments due to their movements relative to the body centre of mass (using cinematography) at different speeds of walking and running. Upper and lower boundaries to the total work were determined by including or excluding possible energy transfers between the segments of each limb, between the limbs and between the centre of mass of the body and the limbs. It appears that the muscle-tendon work of locomotion is most accurately measured when energy transfers are only included between segments of the same limb, but not among the limbs or between the limbs and the centre of mass of the whole body.

Storage and release of mechanical energy by contracting frog muscle fibres.

1. Stretching a contracting muscle leads to greater mechanical work being done during subsequent shortening by its contractile component; the mechanism of this enhancement is not known. 2. This mechanism has been investigated here by subjecting tetanized frog muscle fibres to ramp stretches followed by an isotonic release against a load equal to the maximum isometric tension, T(o). Shortening against T(o) was taken as direct evidence of an absolute increase in the ability to do work as a consequence of the previous stretch. 3. Ramp stretches (0.5-8.6% sarcomere strain, confined to the plateau of the isometric tension-length relationship) were given at different velocities of lengthening (0.03-1.8 sarcomere lengths s-1). Isotonic release to T(o) took place immediately after the end of the ramp, or 5-800 ms after the end of the largest ramp stretches. The length changes taking place after release were measured both at the fibre end and on a tendon-free segment of the fibre. The experiments were carried out at 4 and 14 degrees C. 4. After the elastic recoil of the undamped elastic elements, taking place during the fall in tension at the instant of the isotonic release, a well-defined shortening took place against T(o) (transient shortening against T(o)). 5. The amplitude and time course of transient shortening against T(o) were similar at the fibre end and in the segment, indicating that it is due to a properly of the sarcomeres and not due to stress relaxation of the tendons. 6. Transient shortening against T(o) increased with sarcomere stretch amplitude up to about 8 nm per half-sarcomere independent of stretch velocity. 7. When a short delay (5-20 ms) was introduced between the end of the stretch and the isotonic release, the transient shortening against T(o) did not change; after longer time delays, the transient shortening against T(o) decreased in amplitude. 8. The velocity of transient shortening against T(o) increased with temperature with a temperature coefficient, Q10, of approximately 2.5. 9. It is suggested that transient shortening against T(o) results from the release of mechanical energy stored within the damped element of the cross-bridges. The cross-bridges are brought into a state of greater potential energy not only during the ramp stretch, but also immediately afterwards, during the first phase of stress relaxation.

Variable gearing during locomotion in the human musculoskeletal system.

Human feet and toes provide a mechanism for changing the gear ratio of the ankle extensor muscles during a running step. A variable gear ratio could enhance muscle performance during constant-speed running by applying a more effective prestretch during landing, while maintaining the muscles near the high-efficiency or high-power portion of the force-velocity curve during takeoff. Furthermore, during acceleration, variable gearing may allow muscle contractile properties to remain optimized despite rapid changes in running speed. Forceplate and kinematic analyses of running steps show low gear ratios at touchdown that increase throughout the contact phase.