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First published online May 2, 2008
Journal of Experimental Biology 211, 1571-1578 (2008)
Published by The Company of Biologists 2008
doi: 10.1242/jeb.013805
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The landing–take-off asymmetry of human running is enhanced in old age

G. A. Cavagna1,*, M. A. Legramandi1 and L. A. Peyré-Tartaruga2

1 Istituto di Fisiologia Umana, Università degli Studi di Milano, 20133 Milan, Italy
2 Exercise Research Laboratory, Federal University of Rio Grande do Sul, 90690-200 Porto Alegre, Brazil


Figure 1
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Fig. 1. Mechanical energy of the centre of mass of the body during running steps at the indicated speeds, in an old subject (A,C,E: 80 years, 73.5 kg mass, 1.71 m height, trained runner), and a young subject (B,D,F: 21 years, 58.5 kg mass, 1.79 m height, trained runner). In each panel the curves show the gravitational potential energy (Ep, blue), the kinetic energy of vertical motion (Ekv, red), the kinetic energy of forward motion (Ekf, brown), the kinetic energy of motion in the sagittal plane (Ek=Ekv+Ekf, green), and the total translational energy of the centre of mass of the body in the sagittal plane (Ecm=Ep+Ek, black). The zero line corresponds to the minimum attained by the Ep curve. The horizontal bars indicate the time of contact during the step (from minimum to minimum of Ep). The time during which positive external work is done (increment of Ecm) is indicated in red, whereas the time of negative external work (decrement of Ecm) is indicated in blue. The gap between red and blue bars indicates the duration of the aerial time. The records were obtained from the signals of a force platform. Note the lower vertical oscillation of the centre of mass in the old subject, indicated by a lower amplitude of the Ep and Ekv curves with a shorter aerial time, lower step duration and an higher step frequency.

 

Figure 2
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Fig. 2. Vertical displacement during contact. (A) Old subjects, (B) young subjects, with data from A (gray) superposed for comparison. The upward and downward vertical displacements of the centre of mass taking place when the foot is in contact with the ground (Sc) are divided by the total upward and downward vertical displacements (Sv) to obtain the fraction of the vertical displacement during contact (Sc/Sv). The fractions so obtained, are plotted separately during the lift (Sc,up/Sv,up; open circles) and during the descent (Sc,down/Sv,down; filled circles). The vertical bars indicate the standard deviation (s.d.) of the mean; numbers near the symbols indicate the number of items in the mean. Lines represent the weighted mean of all the data (Kaleidagraph 4.03). Their only purpose is to be a guide for the eye: they do not describe the underlying physical mechanism. Note that at very low speeds Sc/Sv{approx}1 because the aerial phase is often nil (e.g. Fig. 1A,B). With increasing speed an aerial phase of progressively greater extent takes place during the step and the fraction of vertical displacement during contact decreases. The decrement is less during the lift than during the descent (i.e. during the aerial phase the lift is smaller than the fall). This is particularly true in the old subjects who maintain contact with the ground for almost the whole of the lift (see also light-blue dotted lines in Fig. 3). It must be pointed out here that heel-strike and toe-off, i.e. start and end of foot contact, do not properly describe landing and take-off of the bouncing system. As described in the text, landing and take-off, in a physical sense, coincide with the instants where vertical force becomes greater than body weight (after heel strike) and falls below body weight (before toe-off). The correct landing–take-off asymmetry of the bouncing system is described in Fig. 3.

 

Figure 3
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Fig. 3. The four phases of the bounce of the body. Same subjects and speeds as in Fig. 1. In (A–F) the trend of the EpEk transduction, Rint(t) (black), is illustrated with the simultaneous changes in gravitational potential energy, Ep, and kinetic energy, Ek=Ekv+Ekf (green), normalized to oscillate between zero and one. The colors in the Ep curve distinguish the fractions of the step where the vertical force exerted on the ground is greater than body weight (red), and lower than body weight (blue). The continuous Ep line indicates the contact phase whereas the dotted Ep line (light-blue) indicates the aerial phase (not present in A and B). The four phases correspond to the vertical displacement during the upward acceleration Sce,up (red) and deceleration Sae,up (blue), and the downward acceleration Sae,down (blue) and deceleration Sce,down (red). The vertical dotted lines are drawn through the two peaks of Ek and encompass the fraction of the step where the EpEk transduction occurs, as indicated by the increment of the Rint(t) curve. Note that, particularly in the old subject, the transduction of Ek into Ep during the lift (increment Rint,up of Rint(t), below crossing of the interrupted lines) is smaller than the transduction of Ep into Ek during the downward displacement (increment Rint,down of Rint(t), above crossing of the interrupted lines). This asymmetry is accompanied by a ballistic lift (almost nil in the old subject) smaller than the ballistic fall. In the horizontal tracts of the Rint(t) curve no transduction occurs between Ep and Ek and muscle-tendon units absorb Ep and Ek simultaneously (phase β), and increase Ep and Ek simultaneously (phase {alpha}). Whereas most of β is confined within Sce,down, {alpha} usually extends beyond Sce,up within a large fraction of Sae,up due to a continuing increase of Ek. Also this asymmetry is larger in the old subject. In summary, the following features of the landing–take-off asymmetry of the bounce of the body are shown: (i) a peak of Ek greater during the fall than during the lift, (ii) a transduction of Ep into Ek during the fall greater than the transduction of Ek into Ep during the lift (i.e. Rint,down>Rint,up), and (iii) a simultaneous decrement of Ek and Ep after landing shorter than the simultaneous increment of Ek and Ep before and after take-off (i.e. β<{alpha}). All these deviations from an elastic bounce are larger in the old subject.

 

Figure 4
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Fig. 4. Positive and negative external work durations. (A) Old subjects, (B) young subjects, with the data from A (gray) superposed for comparison. The time during which positive external work is done at each step during the push (open circles and red horizontal bars in Fig. 1), and negative external work is done during the brake (filled circles and blue horizontal bars in Fig. 1) are plotted as a function of the running speed. It can be seen that the duration of positive external work is greater than the duration of negative external work up to a speed of ~13 km h–1 (Table 1) indicating an average force during the brake greater than during the push, which is qualitatively consistent with the force–velocity relation of muscle. This suggests that work done by muscle within the muscle-tendon units at low speeds is progressively substituted, with increasing speed, with elastic energy storage and recovery by tendons (see text). On average, the ratio tpush/tbrake is greater in the old subjects indicating a less elastic behavior (Table 2). Asterisks denote statistically significant difference (P<0.05). Other indications as in Fig. 2.

 

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© The Company of Biologists Ltd 2008