Sagittal spine movements of small therian mammals during asymmetrical gaits

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Abstract
SUMMARY: Mammalian locomotion is characterized by the use of asymmetrical gaits associated with extensive flexions and extensions of the body axis. Although the impact of sagittal spine movements on locomotion is well known, little information is available on the kinematics of spinal motion. Intervertebral joint movements were studied in two metatherian and three eutherian species during the gallop and halfbound using high-speed cineradiography. Fast-Fourier transformation was used to filter out high frequency digitizing errors and keep the lower frequency sinusoid oscillations that characterize the intervertebral angular movements. Independent of their regional classification as thoracic or lumbar vertebrae, 7±1 presacral intervertebral joints were involved in sagittal bending movements. In only one species, no more than five intervertebral joints contributed to the resulting `pelvic movement'. In general, the trunk region involved in sagittal bending during locomotion did not correspond to the traditional subdivisions of the vertebral column (e.g. as thoracic and lumbar or pre- and postdiaphragmatic region). Therefore, these classifications do not predict the regions involved in spinal oscillations during locomotion. Independent of the gait, maximum flexion of the spine was observed in the interval between the last third of the swing phase and touch-down. This results in a retraction of the pelvis and hindlimbs before touch-down and, we hypothesize, enhances the stability of the system. Maximum extension occurred during the first third of the swing phase (i.e. after lift-off) in all species. In general, the observed timing of dorsoventral oscillations of the spine are in accordance with that observed in other mammals and with activity data of respiratory and epaxial back muscles. Although no strict craniocaudal pattern was observable, the more cranial intervertebral joints tend to flex and extend earlier than the more caudal ones. This is in accordance with the organization and the activation of the paravertebral musculature in mammals. The amplitude of intervertebral joint movements increased caudally, reaching its highest values in the presacral joint. The more intense sagittal bending movements in the caudal intervertebral joints are reflected by the muscle fiber type composition of the back muscles involved. Despite the highly similar amplitude of `pelvic motion', touch-down and lift-off positions of the pelvis were clearly different between the species with a long, external tail and those with no external tail.

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