Neuromechanical linkage between the head and forearm during running.

OBJECTIVES: The main objective was to test the hypothesis of a neuromechanical link in humans between the head and forearm during running mediated by the biceps brachii and superior trapezius muscles. We hypothesized that this linkage helps stabilize the head and combats rapid forward pitching during running which may interfere with gaze stability. MATERIALS AND METHODS: Thirteen human participants walked and ran on a treadmill while motion capture recorded body segment kinematics and electromyographic sensors recorded muscle activation. To test perturbations to the linkage system we compared participants running normally as well as with added mass to the face and the hand. RESULTS: The results confirm the presence of a neuromechanical linkage between the head and forearm mediated by the biceps and superior trapezius during running but not during walking. In running, the biceps and superior trapezius activations were temporally linked during the stride cycle, and adding mass to either the head or hand increased activation in both muscles, consistent with our hypothesis. During walking the forces acting on the body segments and muscle activation levels were much smaller than during running, indicating no need for a linkage to keep the head and gaze stable. DISCUSSION: The results suggest that the evolution of long distance running in early Homo may have favored selection for reduced rotational inertia of both the head and forearm through synergistic muscle activation, contributing to the transition from australopith head and forelimb morphology to the more human-like form of Homo erectus. Selective pressures from the evolution of bipedal walking were likely much smaller, but may explain in part the intermediate form of the australopith scapula between that of extant apes and humans.

Impact loading and locomotor-respiratory coordination significantly influence breathing dynamics in running humans.

Locomotor-respiratory coupling (LRC), phase-locking between breathing and stepping rhythms, occurs in many vertebrates. When quadrupedal mammals gallop, 1∶1 stride per breath coupling is necessitated by pronounced mechanical interactions between locomotion and ventilation. Humans show more flexibility in breathing patterns during locomotion, using LRC ratios of 2∶1, 2.5∶1, 3∶1, or 4∶1 and sometimes no coupling. Previous studies provide conflicting evidence on the mechanical significance of LRC in running humans. Some studies suggest LRC improves breathing efficiency, but others suggest LRC is mechanically insignificant because 'step-driven flows' (ventilatory flows attributable to step-induced forces) contribute a negligible fraction of tidal volume. Yet, although step-driven flows are brief, they cause large fluctuations in ventilatory flow. Here we test the hypothesis that running humans use LRC to minimize antagonistic effects of step-driven flows on breathing. We measured locomotor-ventilatory dynamics in 14 subjects running at a self-selected speed (2.6±0.1 ms(-1)) and compared breathing dynamics in their naturally 'preferred' and 'avoided' entrainment patterns. Step-driven flows occurred at 1-2X step frequency with peak magnitudes of 0.97±0.45 Ls(-1) (mean ±S.D). Step-driven flows varied depending on ventilatory state (high versus low lung volume), suggesting state-dependent changes in compliance and damping of thoraco-abdominal tissues. Subjects naturally preferred LRC patterns that minimized antagonistic interactions and aligned ventilatory transitions with assistive phases of the step. Ventilatory transitions initiated in 'preferred' phases within the step cycle occurred 2x faster than those in 'avoided' phases. We hypothesize that humans coordinate breathing and locomotion to minimize antagonistic loading of respiratory muscles, reduce work of breathing and minimize rate of fatigue. Future work could address the potential consequences of locomotor-ventilatory interactions for elite endurance athletes and individuals who are overweight or obese, populations in which respiratory muscle fatigue can be limiting.

The evolution of marathon running : capabilities in humans.

Humans have exceptional capabilities to run long distances in hot, arid conditions. These abilities, unique among primates and rare among mammals, derive from a suite of specialised features that permit running humans to store and release energy effectively in the lower limb, help keep the body's center of mass stable and overcome the thermoregulatory challenges of long distance running. Human endurance running performance capabilities compare favourably with those of other mammals and probably emerged sometime around 2 million years ago in order to help meat-eating hominids compete with other carnivores.

The human gluteus maximus and its role in running.

The human gluteus maximus is a distinctive muscle in terms of size, anatomy and function compared to apes and other non-human primates. Here we employ electromyographic and kinematic analyses of human subjects to test the hypothesis that the human gluteus maximus plays a more important role in running than walking. The results indicate that the gluteus maximus is mostly quiescent with low levels of activity during level and uphill walking, but increases substantially in activity and alters its timing with respect to speed during running. The major functions of the gluteus maximus during running are to control flexion of the trunk on the stance-side and to decelerate the swing leg; contractions of the stance-side gluteus maximus may also help to control flexion of the hip and to extend the thigh. Evidence for when the gluteus maximus became enlarged in human evolution is equivocal, but the muscle's minimal functional role during walking supports the hypothesis that enlargement of the gluteus maximus was likely important in the evolution of hominid running capabilities.

Endurance running and the evolution of Homo.

Striding bipedalism is a key derived behaviour of hominids that possibly originated soon after the divergence of the chimpanzee and human lineages. Although bipedal gaits include walking and running, running is generally considered to have played no major role in human evolution because humans, like apes, are poor sprinters compared to most quadrupeds. Here we assess how well humans perform at sustained long-distance running, and review the physiological and anatomical bases of endurance running capabilities in humans and other mammals. Judged by several criteria, humans perform remarkably well at endurance running, thanks to a diverse array of features, many of which leave traces in the skeleton. The fossil evidence of these features suggests that endurance running is a derived capability of the genus Homo, originating about 2 million years ago, and may have been instrumental in the evolution of the human body form.