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First published online August 22, 2008
Journal of Experimental Biology 211, 2735-2751 (2008)
Published by The Company of Biologists 2008
doi: 10.1242/jeb.018820
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The movements of limb segments and joints during locomotion in African and Asian elephants

Lei Ren1, Melanie Butler1, Charlotte Miller1, Heather Paxton1, Delf Schwerda2, Martin S. Fischer2 and John R. Hutchinson1,*

1 Structure and Motion Laboratory, Department of Veterinary Basic Sciences, The Royal Veterinary College, University of London, Hatfield, Hertfordshire AL9 7TA, UK
2 Institut fuer Spezielle Zoologie und Evolutionsbiologie, mit Phyletischem Museum, Jena 07743, Germany


Figure 1
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  		Fig. 1. Marker placements on a representative African elephant. (A) Subject G (Table 1) in oblique right lateral view showing all skin markers (back markers shown are not used in this study). (B) Relationship of markers with underlying skeleton. (C) Definitions of segment and joint angles [picture modified from Shoshani (Shoshani, 1992Go)]. Palpated anatomical positions of markers (Smuts and Bezuidenhout, 1993Go; Smuts and Bezuidenhout, 1994Go): lateral side of greater tubercle of humerus, lateral epicondyle of humerus, styloid process of ulna, toenail of manus digit 3, caudal side of accessory carpal, greater trochanter of femur (just caudal to tuber coxae of ilium), lateral epicondyle of femur (just caudal and proximal to patella), lateral malleolus of fibula, middle of toenail of pes digit 3, caudal side of calcaneal tuber. We only used the calcaneus and carpal markers to identify touch-down/lift-off events (see Materials and Methods). The segmental angles were calculated relative to a vertical line through the proximal marker of each segment; only shown precisely for the upper arm and thigh segments.

 

Figure 2
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  		Fig. 2. Representative limb segment angular trajectories during a stride for an elephant (Subject H, Table 1) moving at normal walking speed; 1.37±0.28 m s–1 (Fr=0.11; N=5). Stance phase is shown in blue; swing phase in red. Note that the stance phase for the forelimb (right side) is offset ~15% of a stride relative to the hindlimb, as the data were collected synchronously; this relative limb phase offset is typical for a lateral sequence walk in elephants (Hutchinson et al., 2003Go; Hutchinson et al., 2006Go).

 

Figure 3
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  		Fig. 3. Representative limb joint angular trajectories during a stride, shown for the same elephant as in Fig. 2. Stance phase is shown in blue, swing phase in red.

 

Figure 4
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  		Fig. 4. Representative limb segment (thigh and upper arm; A–B) and joint (C–F) angular trajectories during a stride for an elephant (Subject N, Table 1) moving at a fast walk/slow run (thin lines; 2.15±0.24 m s–1, Fr=0.31; N=2) and a fast run (thick lines; 4.25±0.19 m s–1, Fr=1.22; N=3) to show the effect of a doubling of speed on limb kinematics. Stance phase is shown in blue, swing phase in red.

 

Figure 5
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  		Fig. 5. Representative limb segment (thigh and upper arm; A–B) and joint (C–F) angular velocities during a stride, shown for the same elephant as in Fig. 4. Stance phase is shown in blue, swing phase in red.

 

Figure 6
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  		Fig. 6. Forelimb segment/joint angles (mean values ±1 s.e.m.) at particular events (ROM = range of motion) in one stride compared across our speed range; A–G=speed categories (see supplementary material Tables S1 and S2). Dimensionless speed (û) ranges are: A, very slow, <0.25 m s–1; B, slow, 0.25–0.30 m s–1; C, normal, 0.30–0.35 m s–1; D, medium fast, 0.35–0.50 m s–1; E, fast, 0.50–0.75 m s–1; F, very fast, 0.75–1.0 m s–1; G, Fr>1 run, û>1.0.

 

Figure 7
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  		Fig. 7. Hindlimb segment/joint angles (mean values ±1 s.e.m.) at particular events (ROM = range of motion) in one stride compared across our speed range; A–G=speed categories (see supplementary material Tables S1 and S2); dimensionless speed (û) ranges are: A, very slow, <0.25 m s–1; B, slow, 0.25–0.30 m s–1; C, normal, 0.30-0.35 m s–1; D, medium fast, 0.35–0.50 m s–1; E, fast, 0.50–0.75 m s–1; F, very fast, 0.75–1.0 m s–1; G, Fr>1 run, û>1.0.

 

Figure 8
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  		Fig. 8. Forelimb segment/joint angular velocities (mean values ±1 s.e.m.) at particular events (RAV = range of angular velocity) in one stride compared across our speed range; A G=speed categories (see supplementary material Tables S1 and S2); dimensionless speed (û) ranges are: A, very slow, <0.25 m s–1; B, slow, 0.25–0.30 m s–1; C, normal, 0.30–0.35 m s–1; D, medium fast, 0.35–0.50 m s–1; E, fast, 0.50–0.75 m s–1; F, very fast, 0.75–1.0 m s–1; G, Fr>1 run, û>1.0.

 

Figure 9
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  		Fig. 9. Hindlimb segment/joint angular velocities (mean values ±1 s.e.m.) at particular events (RAV = range of angular velocity) in one stride compared across our speed range; A–G=speed categories (see supplementary material Tables S1 and S2); dimensionless speed (û) ranges are: A, very slow, <0.25 m s–1; B, slow, 0.25–0.30 m s–1; C, normal, 0.30–0.35 m s–1; D, medium fast, 0.35–0.50 m s–1; E, fast, 0.50–0.75 m s–1; F, very fast, 0.75–1.0 m s–1; G, Fr>1 run, û>1.0.

 

Figure 10
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  		Fig. 10. Stick figure portrayals of elephant limb motions for subject O (Table 1) in both stance and swing phases for the fore- and hindlimbs at normal walking (1.26 m s–1; Fr=0.11; shown at 20 Hz), and fast running (4.81 m s–1; Fr=1.56; shown at 40 Hz). Because of space constraints the hip and shoulder positions are kept constant during the swing phase. Also represented as online supplementary material (Movies 1 and 2).

 

Figure 11
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  		Fig. 11. Comparison of hindlimb segment/joint kinematics between humans and elephants. Elephant hindlimb data shown are for subject N (Table 1) fast walking at 1.93±0.07 m s–1 (Fr=0.25; N=2) and human limb data shown are from Ren et al. [(Ren et al., in review) age 28 years, mass 69 kg]; walking at 1.45±0.08 m s–1 (Fr=0.25).

 

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© The Company of Biologists Ltd 2008