Reilly Stephen M, Blob Richard W
Department of Biological Sciences, Ohio University, Athens, OH 45701, USA.
J Exp Biol. 2003 Dec;206(Pt 23):4327-40. doi: 10.1242/jeb.00688.
Crocodilians are unusual among quadrupedal tetrapods in their frequent use of a wide variety of hindlimb postures, ranging from sprawling to a more erect high walk. In this study, we use synchronized kinematic videos and electromyographic recordings to test how the activity patterns of hindlimb muscles in American alligators (Alligator mississippiensis Daudin) differ between sprawling and more upright postures. Previous force platform analyses suggested that upright posture in alligators would require greater activation by hindlimb extensors to counter increases in the flexor moments exerted about joints by the ground reaction force during upright stance. Consistent with these predictions, ankle extensors (gastrocnemius) and knee extensors (femorotibialis internus and iliotibialis 2) exhibit increases in signal intensity during the use of more upright stance. Bone loading data also predicted that activation patterns for hip adductors spanning the length of the femur would not differ between sprawling and more upright posture. Correspondingly, motor patterns of the adductor femoris were not altered as posture became more upright. However, the adductor puboischiofemoralis externus 3, which inserts far proximally on the femur, displays significant increases in burst intensity that could contribute to the greater femoral adduction that is integral to upright posture. In contrast to patterns in alligators, in mammals EMG burst intensity typically decreases during the use of upright posture. This difference in the motor control of limb posture between these taxa may be related to differences in the relative sizes of their feet. Alligator feet are large relative to the hindlimb and, as a result, the ground reaction force shifts farther from the limb joints during upright steps than in mammals, increasing flexor moments at joints and requiring alligator extensor muscles to exert greater forces to keep the limb in equilibrium. However, several alligator hindlimb muscles show no differences in motor pattern between sprawling and upright posture. The wide range of motor pattern modulations between different postures in alligators suggests considerable independence of neural control among the muscles of the alligator hindlimb.
在四足四足动物中,鳄鱼不同寻常之处在于它们频繁使用各种不同的后肢姿势,从 sprawled(四肢展开的姿势)到更为直立的高步行走。在本研究中,我们使用同步运动学视频和肌电图记录来测试美国短吻鳄(密西西比鳄,Alligator mississippiensis Daudin)后肢肌肉在 sprawled 姿势和更直立姿势之间的活动模式有何不同。先前的力平台分析表明,短吻鳄的直立姿势需要后肢伸肌更大程度的激活,以抵消直立站立时地面反作用力在关节处施加的屈肌力矩增加。与这些预测一致,在使用更直立姿势时,踝关节伸肌(腓肠肌)和膝关节伸肌(股胫内侧肌和髂胫肌 2)的信号强度增加。骨骼负荷数据还预测,横跨股骨长度的髋内收肌的激活模式在 sprawled 姿势和更直立姿势之间不会有差异。相应地,随着姿势变得更加直立,股内收肌的运动模式没有改变。然而,远在股骨近端附着的耻骨坐骨股外肌 3 的爆发强度显著增加,这可能有助于直立姿势所必需的更大程度的股骨内收。与短吻鳄的模式不同,在哺乳动物中,肌电图爆发强度在使用直立姿势时通常会降低。这些类群在肢体姿势运动控制上的这种差异可能与它们足部相对大小的差异有关。短吻鳄的脚相对于后肢较大,因此,在直立行走时,地面反作用力比在哺乳动物中更远离肢体关节,增加了关节处的屈肌力矩,需要短吻鳄的伸肌施加更大的力来保持肢体平衡。然而,几种短吻鳄后肢肌肉在 sprawled 姿势和直立姿势之间的运动模式没有差异。短吻鳄不同姿势之间广泛的运动模式调制表明,短吻鳄后肢肌肉的神经控制具有相当大的独立性。