Center for Neuromotor and Biomechanics Research, University of Houston, Houston, TX 77054, USA.
J Exp Biol. 2012 Dec 1;215(Pt 23):4104-14. doi: 10.1242/jeb.075275. Epub 2012 Sep 12.
This study explored the muscle strain and activation patterns of several key flight muscles of the pigeon (Columba livia) during takeoff and landing flight. Using electromyography (EMG) to measure muscle activation, and sonomicrometry to quantify muscle strain, we evaluated the muscle function patterns of the pectoralis, biceps, humerotriceps and scapulotriceps as pigeons flew between two perches. These recordings were analyzed in the context of three-dimensional wing kinematics. To understand the different requirements of takeoff, midflight and landing, we compared the activity and strain of these muscles among the three flight modes. The pectoralis and biceps exhibited greater fascicle strain rates during takeoff than during midflight or landing. However, the triceps muscles did not exhibit notable differences in strain among flight modes. All observed strain, activation and kinematics were consistent with hypothesized muscle functions. The biceps contracted to stabilize and flex the elbow during the downstroke. The humerotriceps contracted to extend the elbow at the upstroke-downstroke transition, followed by scapulotriceps contraction to maintain elbow extension during the downstroke. The scapulotriceps also appeared to contribute to humeral elevation. Greater muscle activation intensity was observed during takeoff, compared with mid-flight and landing, in all muscles except the scapulotriceps. The timing patterns of muscle activation and length change differed among flight modes, yet demonstrated that pigeons do not change the basic mechanical actions of key flight muscles as they shift from flight activities that demand energy production, such as takeoff and midflight, to maneuvers that require absorption of energy, such as landing. Similarly, joint kinematics were consistent among flight modes. The stereotypy of these neuromuscular and joint kinematic patterns is consistent with previously observed stereotypy of wing kinematics relative to the pigeon's body (in the local body frame) across these flight behaviors. Taken together, these observations suggest that the control of takeoff and landing flight primarily involves modulation of overall body pitch to effect changes in stroke plane angle and resulting wing aerodynamics.
本研究探讨了鸽子(Columba livia)起飞和降落飞行过程中几个关键飞行肌肉的肌肉应变和激活模式。使用肌电图(EMG)测量肌肉激活,超声测量肌肉应变,我们评估了鸽子在两个栖木之间飞行时胸大肌、二头肌、肱三头肌和肩胛胸肌的肌肉功能模式。这些记录在三维翅膀运动学的背景下进行了分析。为了了解起飞、中飞和降落的不同要求,我们比较了这三种飞行模式下这些肌肉的活动和应变。胸大肌和二头肌在起飞时比在中飞或降落时表现出更大的肌束应变率。然而,三头肌在飞行模式之间没有表现出明显的应变差异。所有观察到的应变、激活和运动学都与假设的肌肉功能一致。在下降冲程中,二头肌收缩以稳定和弯曲肘部。在上升-下降转换时,肱三头肌收缩以伸展肘部,然后肩胛胸肌收缩以在下降冲程中保持肘部伸展。肩胛胸肌似乎也有助于肱骨抬高。与中飞和降落相比,除了肩胛胸肌之外,所有肌肉在起飞时的肌肉激活强度都更高。在起飞、中飞和降落时,肌肉激活和长度变化的时间模式不同,但表明鸽子在从需要能量产生的飞行活动(如起飞和中飞)转换为需要吸收能量的机动(如降落)时,不会改变关键飞行肌肉的基本机械动作。同样,关节运动学在飞行模式之间是一致的。这些神经肌肉和关节运动学模式的刻板性与之前观察到的翅膀运动学相对于鸽子身体(在局部身体框架中)在这些飞行行为中的刻板性一致。总之,这些观察结果表明,起飞和降落飞行的控制主要涉及对整体身体俯仰的调节,以改变冲程平面角度和由此产生的机翼空气动力学。
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