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First published online October 27, 2003
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Motor control of locomotor hindlimb posture in the American alligator (Alligator mississippiensis)

Stephen M. Reilly1 and Richard W. Blob2,*

1 Department of Biological Sciences, Ohio University, Athens, OH 45701, USA
2 Department of Biological Sciences, Clemson University, Clemson, SC 29634, USA



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Fig. 1. Diagrammatic illustrations of alternative models to explain increases in femoral loading during upright locomotion in alligators (adapted from Blob, 1998Go, 2001Go; Blob and Biewener, 1999Go, 2001Go). For visual clarity of the forces and moments bearing on the model, the views are presented from an oblique posterolateral perspective (this causes the femur to appear not to project perpendicular to the vertebral axis as it does at mid-stance). For each arrow depicting a force or moment, a difference in thickness (not length) between the left and right sides of a panel indicates a difference in force or moment magnitude between the postures illustrated in those panels (with thicker arrows indicating larger forces or moments). Note that panels A and B are drawn to different scales (A is magnified for clarity), and that comparisons of force and moment magnitudes are not intended between A and B. Because the ground reaction force (GRF) does not differ in magnitude between sprawling and upright steps (Blob and Biewener, 2001Go), both models are based on changes in the action of hindlimb muscles between these postures. (A) Bending induced by the ground reaction force (red arrow) places the dorsal femur in compression (-), and the ventral femur in tension (+). In sprawling posture (left), the GRF might have a longer moment arm about the hip than in upright posture (right), resulting in a larger abductor moment that would tend to rotate the femur dorsally. To keep the hip joint in equilibrium, the hip adductors might exert a larger force in sprawling posture (left) and a smaller force in upright posture (right). Because the hip adductors bend the femur in the opposite direction from the GRF, larger adductor forces during sprawling steps could more effectively mitigate strains induced by the GRF, resulting in lower dorsal and ventral stresses and strains during sprawling steps. (B) As limb posture becomes more upright, the center of pressure of the GRF shifts away from the ankle, increasing the moment arm of the GRF at the ankle (RGRF). Consequently, ankle extensors (e.g. gastrocnemius) must exert higher forces during upright steps in order to counter the larger ankle flexor moment and maintain joint equilibrium. Because gastrocnemius also spans the knee, it makes a greater contribution to the flexor moment at the knee during more upright steps, and knee extensors (femorotibialis and iliotibialis, on the dorsal aspect of the femur) must exert greater force to counter this moment and maintain equilibrium at the knee. Increases in knee extensor forces could then raise dorsal and ventral femoral strains and stresses as alligators use more upright posture. Data from previous force platform studies (Blob and Biewener, 2001Go) are consistent with the model proposed in B, but changes in muscle activity patterns have not been tested prior to this study. rankext, moment arm of ankle extensor muscles at ankle (no change between sprawling and upright stance).

 


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Fig. 2. Hindlimb muscles from which electromyographic (EMG) data were collected in alligators walking with a range of postures. Lines of actions are indicated for muscles primarily active in either stance phase (red), swing phase (blue) or with activity in both phases (black). Muscle abbreviations are indicated in the text (under Myology).

 


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Fig. 3. Modulation of the timing and intensity of hindlimb muscle activity between sprawling and high-walk locomotion in alligators. Significant changes in motor patterns (Table 2) as limb posture becomes more erect are illustrated by comparing mean electromyographic (EMG) patterns for more sprawling postures (about 30° of femoral adduction; gray bars) to those for more erect postures (about 50° of femoral adduction; black bars). Open bars indicate muscle patterns for which there was no significant change in motor patterns (timing or amplitude) as femoral adduction angle increased. Differences in bar thickness between sprawling and upright stance indicate significant differences in mean amplitude (in all cases where such differences occur, black bars for upright posture are thicker than gray bars for sprawling posture). The number of individuals from which EMG data were collected is listed in parentheses after each muscle name. In the top panel, mean three-dimensional limb segmental kinematics for the low (gray line) and high (black lines) walking postures are shown, illustrating differences between these postures (from Reilly and Elias, 1998Go). Note that more erect postures are produced by consistently greater femoral adduction, knee extension and ankle extension during the stance phase. Abbreviations: FTE, flexor tibialis externus; ADDFEM1, adductor femoris, head 1; PIFE3, puboischiofemoralis externus, head 3; PIT (a), puboischiotibialis (stance burst); PIT (b), puboischiotibialis (swing burst); FEMTIB, femorotibialis internus; ILTIB1, iliotibialis, head 1; ILTIB2, iliotibialis, head 2; GAST, gastrocnemius; PIFI2, puboischiofemoralis internus, head 2; ILFEM, iliofemoralis; AMB 1, ambiens, head 1; TA, tibialis anterior.

 


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Fig. 4. Representative regressions of electromyographic (EMG) variables on femoral adduction angle for hindlimb extensor muscles of individual experimental alligators. Larger angles indicate more upright posture, as illustrated by the sketches at the bottom of the figure. N=30 for both regressions. (A) Mean burst amplitude for the femorotibialis of Alligator 2 (=FEMTIB AREA DUR of Table 2). (B) Mean burst amplitude for the gastrocnemius of Alligator 4 (=GAST AREA DUR of Table 2). Burst amplitudes increase significantly with the use of upright posture for both muscles.

 





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