Blob R W, Biewener A A
Department of Zoology, Division of Fishes, Field Museum of Natural History, Chicago, IL 60605, USA.
J Exp Biol. 2001 Mar;204(Pt 6):1099-122. doi: 10.1242/jeb.204.6.1099.
In vivo measurements of strain in the femur and tibia of Iguana iguana (Linnaeus) and Alligator mississippiensis (Daudin) have indicated three ways in which limb bone loading in these species differs from patterns observed in most birds and mammals: (i) the limb bones of I. iguana and A. mississippiensis experience substantial torsion, (ii) the limb bones of I. iguana and A. mississippiensis have higher safety factors than those of birds or mammals, and (iii) load magnitudes in the limb bones of A. mississippiensis do not decrease uniformly with the use of a more upright posture. To verify these patterns, and to evaluate the ground and muscle forces that produce them, we collected three-dimensional kinematic and ground reaction force data from subadult I. iguana and A. mississippiensis using a force platform and high-speed video. The results of these force/kinematic studies generally confirm the loading regimes inferred from in vivo strain measurements. The ground reaction force applies a torsional moment to the femur and tibia in both species; for the femur, this moment augments the moment applied by the caudofemoralis muscle, suggesting large torsional stresses. In most cases, safety factors in bending calculated from force/video data are lower than those determined from strain data, but are as high or higher than the safety factors of bird and mammal limb bones in bending. Finally, correlations between limb posture and calculated stress magnitudes in the femur of I. iguana confirm patterns observed during direct bone strain recordings from A. mississippiensis: in more upright steps, tensile stresses on the anterior cortex decrease, but peak compressive stresses on the dorsal cortex increase. Equilibrium analyses indicate that bone stress increases as posture becomes more upright in saurians because the ankle and knee extensor muscles exert greater forces during upright locomotion. If this pattern of increased bone stress with the use of a more upright posture is typical of taxa using non-parasagittal kinematics, then similar increases in load magnitudes were probably experienced by lineages that underwent evolutionary shifts to a non-sprawling posture. High limb bone safety factors and small body size in these lineages could have helped to accommodate such increases in limb bone stress.
对绿鬣蜥(Iguana iguana,林奈)和密西西比鳄(Alligator mississippiensis,达丁)股骨和胫骨应变的体内测量表明,这些物种的肢体骨骼加载方式与大多数鸟类和哺乳动物中观察到的模式存在三方面差异:(i)绿鬣蜥和密西西比鳄的肢体骨骼承受大量扭转;(ii)绿鬣蜥和密西西比鳄的肢体骨骼比鸟类或哺乳动物的肢体骨骼具有更高的安全系数;(iii)密西西比鳄肢体骨骼中的负荷大小不会随着更直立姿势的使用而均匀降低。为了验证这些模式,并评估产生这些模式的地面力和肌肉力,我们使用测力平台和高速视频,从亚成体绿鬣蜥和密西西比鳄收集了三维运动学和地面反作用力数据。这些力/运动学研究的结果总体上证实了从体内应变测量推断出的加载方式。地面反作用力在两个物种中都对股骨和胫骨施加扭力矩;对于股骨,这个力矩增强了股尾肌施加的力矩,表明存在较大的扭转应力。在大多数情况下,根据力/视频数据计算出的弯曲安全系数低于根据应变数据确定的安全系数,但与鸟类和哺乳动物肢体骨骼弯曲时的安全系数一样高或更高。最后,绿鬣蜥股骨中肢体姿势与计算出的应力大小之间的相关性证实了在密西西比鳄直接骨骼应变记录期间观察到的模式:在更直立的步态中,前皮质上的拉应力降低,但背皮质上的峰值压应力增加。平衡分析表明,在蜥蜴中,随着姿势变得更直立,骨骼应力会增加,因为在直立运动过程中,踝关节和膝关节伸肌会施加更大的力。如果这种随着使用更直立姿势而骨骼应力增加的模式对于使用非矢状面运动学的类群是典型的,那么经历了向非 sprawl 姿势进化转变 的谱系可能也经历了类似的负荷大小增加。这些谱系中高的肢体骨骼安全系数和小体型可能有助于适应肢体骨骼应力的这种增加。