Structural attributes contributing to locomotor performance in the ostrich

As the fastest long-endurance runner, the bipedal ostrich (Struthio camelus) was selected as a prime model organism to investigate the physical attributes underlying this advanced locomotor performance. A specific integrative approach combining morphological, morphometric, kinematic and pedobarographic methods was developed. The comparative morphometric analysis of the hind limbs of all ratite species revealed that leg segment ratios in the ostrich are the most specialised for efficient locomotion, especially when taking into consideration its unique supra-jointed toe posture. In addition, the crural muscle mass is more concentrated towards the hip joint in the ostrich than in its ratite relatives. According to the Law of the Pendulum, this concentration of mass towards the pivot point – in concert with the relatively longest and lightest distal leg elements – represents a mechanical optimisation of limb swinging capacities. While musculature clearly drives limb movement, the passive guidance and constraint of motion range by ligamentous structures combined with joint surface contours allows a high level of energy output efficiency during all stages of locomotion and ensures articular stability during slow locomotion as well as high-speed performance. So far, the influence of these passive effects in locomotion has been largely ignored. In order to quantify the guiding effect of these anatomical structures, kinematic data of adult ostriches during walking and running were collected. Subsequently, these data were compared with results from manual manipulation experiments performed with the limbs of anatomical specimens – both fully intact and with muscles removed – leaving only the ligament system intact. This investigation revealed that the range of motion among leg segments was nearly identical in all sample groups, especially in regard to maximum extension values. This indicates that ostrich hind limb dynamics are managed to a significant degree by passive elements that ensure a controlled swing-plane with minimal deviation from an optimal attitude. Further dissections allowed some of these features to be described in detail, with an emphasis on functional-morphological examination of the intertarsal joint. The intertarsal joint contains a significant locking mechanism, briefly mentioned in historical documents, but described and functionally analysed herein for the first time. The functional examination qualified the interplay of three collateral ligaments, the tendinous M. fibularis brevis and specific joint surface protrusions as the basis for this effect which remains absent in smaller ground-dwelling bird species. A proximate quantification, based on comparative morphological and kinematic data, revealed function of Struthio's passively locked intertarsal joint as a potent stabiliser in the supporting limb during the ground-contact phase of locomotion. During stance phase, it is crucial that the supporting limb is stabilised internally and in relation to the substrate. As yet, no study exists concerning use and loading of the actual ground contact elements. The toes must absorb body mass, guarantee stable grip and provide energetic push off. Obvious specialisations of the ostrich's phalangeal complex include toe reduction (leaving only 3rd and 4th toe), claw reduction (only at 3rd toe) and a permanently elevated metatarsophalangeal joint. Using a relatively new methodology to examine in vivo toe function, pedobarography was employed on specifically trained ostriches to allow extensive collection of Centre of Pressure (CoP) and load distribution (LD) data. In contrast to a relatively predictable CoP trajectory at all speeds, conspicuous LD differences were observed between slow and fast trials. Load was distributed rather inconsistently during walking, while a typical tripod-like toe-print occurred in all running trials to presumably deliver additional stability during the comparatively short stance phase. Significant grip is provided by the highly directed impact of the 3rd toe claw-tip, suggesting its important function as a positional anchor during running. Pedobarographic analysis further showed the importance of the 4th toe as an outrigger to maintain balance, rendering a future reduction highly unlikely. In conclusion, the application of interdisciplinary methodologies allowed comprehensive data collection and integration of the model organism within its ecological context. The data gained from this thesis increases the current knowledge about ostrich locomotion by identifying distinct structural attributes as essential elements for extreme cursorial performance. The present data may alter existing models for calculation of the metabolic cost of terrestrial locomotion and aid in the reconstruction of theropod locomotion, as these branch sciences often overlook the important role of ligaments and passively-coupled motion cycles in reducing the cost of locomotion

The intertarsal joint of the ostrich (Struthio camelus): Anatomical examination and function of passive structures in locomotion

The ostrich (Struthio camelus) is the largest extant biped. Being flightless, it exhibits advanced cursorial abilities primarily evident in its characteristic speed and endurance. In addition to the active musculoskeletal complex, its powerful pelvic limbs incorporate passive structures wherein ligaments interact with joint surfaces, cartilage and other connective tissue in their course of motion. This arrangement may enable energy conservation by providing joint stabilisation, optimised limb segment orientation and automated positioning of ground contact elements independently of direct muscle control