Kram R, Wong B, Full R J
Department of Integrative Biology, University of California at Berkeley, 94720-3140, USA.
J Exp Biol. 1997 Jul;200(Pt 13):1919-29. doi: 10.1242/jeb.200.13.1919.
We tested the hypothesis that fast-running hexapeds must generate high levels of kinetic energy to cycle their limbs rapidly compared with bipeds and quadrupeds. We used high-speed video analysis to determine the three-dimensional movements of the limbs and bodies of cockroaches (Blaberus discoidalis) running on a motorized treadmill at 21 cm s-1 using an alternating tripod gait. We combined these kinematic data with morphological data to calculate the mechanical energy produced to move the limbs relative to the overall center of mass and the mechanical energy generated to rotate the body (head + thorax + abdomen) about the overall center of mass. The kinetic energy involved in moving the limbs was 8 microJ stride-1 (a power output of 21 mW kg-1, which was only approximately 13% of the external mechanical energy generated to lift and accelerate the overall center of mass at this speed. Pitch, yaw and roll rotational movements of the body were modest (less than +/- 7 degrees), and the mechanical energy required for these rotations was surprisingly small (1.7 microJ stride-1 for pitch, 0.5 microJ stride-1 for yaw and 0.4 microJ stride-1 for roll) as was the power (4.2, 1.2 and 1.1 mW kg-1, respectively). Compared at the same absolute forward speed, the mass-specific kinetic energy generated by the trotting hexaped to swing its limbs was approximately half of that predicted from data on much larger two- and four-legged animals. Compared at an equivalent speed (mid-trotting speed), limb kinetic energy was a smaller fraction of total mechanical energy for cockroaches than for large bipedal runners and hoppers and for quadrupedal trotters. Cockroaches operate at relatively high stride frequencies, but distribute ground reaction forces over a greater number of relatively small legs. The relatively small leg mass and inertia of hexapeds may allow relatively high leg cycling frequencies without exceptionally high internal mechanical energy generation.
与两足动物和四足动物相比,快速奔跑的六足动物必须产生高水平的动能才能快速循环其肢体。我们使用高速视频分析来确定蟑螂(Discoidalis蜚蠊)在电动跑步机上以21厘米/秒的速度使用交替三脚架步态奔跑时肢体和身体的三维运动。我们将这些运动学数据与形态学数据相结合,以计算相对于整体质心移动肢体所产生的机械能以及使身体(头部 + 胸部 + 腹部)围绕整体质心旋转所产生的机械能。移动肢体所涉及的动能为8微焦耳/步(功率输出为21毫瓦/千克,这仅约为此速度下提升和加速整体质心所产生的外部机械能的13%)。身体的俯仰、偏航和滚动旋转运动幅度较小(小于±7度),并且这些旋转所需的机械能出奇地小(俯仰为1.7微焦耳/步,偏航为0.5微焦耳/步,滚动为0.4微焦耳/步),功率也很小(分别为4.2、1.2和1.1毫瓦/千克)。在相同的绝对前进速度下进行比较,小跑的六足动物摆动其肢体所产生的质量比动能约为根据更大体型的两足和四足动物数据预测值的一半。在等效速度(小跑中间速度)下进行比较,蟑螂的肢体动能在总机械能中所占比例比大型两足奔跑者和跳跃者以及四足小跑者要小。蟑螂以相对较高的步频运行,但将地面反作用力分布在更多相对较小的腿上。六足动物相对较小的腿部质量和惯性可能允许相对较高的腿部循环频率,而无需产生异常高的内部机械能。