Asymmetric bipedal locomotion – an adaptive response to incomplete spinal injury in the chick

Abstract The purpose of this study was to compare the asymmetric gait induced by unilateral spinal cord injury in chicks with asymmetric gaits of other bipeds and quadrupeds. After lateral hemisection of the left thoracic spinal cord, kinetic (ground reaction forces) and kinematic (distance and timing) data were recorded as chicks moved overground unrestrained. Ground reaction forces were analyzed to obtain the mechanical energy changes throughout the stride. Kinematic measurements were obtained over a range of speeds to determine the velocity-dependent characteristics of the gait. Hemisected chicks adopted an asymmetric hopping gait in which the animals hopped from the right leg (contralateral to the lesion) onto the left (ipsilateral) leg but then fell forward onto the right leg. Mechanical energy fluctuations throughout a single stride (i.e., two steps) approximated the oscillations that occur during a single walking step of control animals. When examined over a range of velocities, asymmetries in limb timing remained constant, but distance measurements such as step length became more symmetric as speed increased.The results show that, after spinal hemisection, adaptations of the remaining neural circuitry permitted the production of a locomotor pattern that, in addition to providing effective support and propulsion, incorporated some of the energy-conserving mechanisms of the normal walk. Adjustment of this novel locomotor pattern for different velocities further demonstrates the flexibility of locomotor circuitry. Comparisons with other studies shows that this gait shares some temporal and energetic features with asymmetric gaits of several bipedal species, including humans. In particular, hemisected chicks and some hemiplegic humans adopt an asymmetric gait in which maximum energy recovery occurs during the stance of the affected limb; these similarities probably relate to common mechanical constraints imposed on bipedal forms of terrestrial locomotion.

[1]  P A Costigan,et al.  Mechanical energy patterns in gait of cerebral palsied children with hemiplegia. , 1987, Physical therapy.

[2]  J. Steeves,et al.  Phasic cutaneous input facilitates locomotor recovery after incomplete spinal injury in the chick. , 1995, Journal of neurophysiology.

[3]  G. Cavagna,et al.  Mechanical work in terrestrial locomotion: two basic mechanisms for minimizing energy expenditure. , 1977, The American journal of physiology.

[4]  S Carlsöö,et al.  Kinetic analysis of the gait in patients with hemiparesis and in patients with intermittent claudication. , 1974, Scandinavian journal of rehabilitation medicine.

[5]  M. P. Murray Gait as a total pattern of movement. , 1967, American journal of physical medicine.

[6]  A. Bekoff Neuroethological approaches to the study of motor development in chicks: achievements and challenges. , 1992, Journal of neurobiology.

[7]  P A Costigan,et al.  Mechanical energy of walking of stroke patients. , 1986, Archives of physical medicine and rehabilitation.

[8]  J. Steeves,et al.  Ontogeny of bipedal locomotion: walking and running in the chick. , 1996, The Journal of physiology.

[9]  Murray Mp,et al.  Gait as a total pattern of movement. , 1967 .

[10]  S. Gatesy,et al.  Bipedal locomotion: effects of speed, size and limb posture in birds and humans , 1991 .

[11]  J Whitall,et al.  A developmental study of the interlimb coordination in running and galloping. , 1989, Journal of motor behavior.

[12]  G. Cavagna,et al.  Energetics and mechanics of terrestrial locomotion. III. Energy changes of the centre of mass as a function of speed and body size in birds and mammals. , 1982, The Journal of experimental biology.

[13]  A. Bekoff,et al.  Neural control of limb coordination. I. Comparison of hatching and walking motor output patterns in normal and deafferented chicks , 1987, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[14]  S. Olney,et al.  Temporal, kinematic, and kinetic variables related to gait speed in subjects with hemiplegia: a regression approach. , 1994, Physical therapy.

[15]  K. Pearson,et al.  Reversal of the influence of group Ib afferents from plantaris on activity in medial gastrocnemius muscle during locomotor activity. , 1993, Journal of neurophysiology.

[16]  L Finch Hemiplegic gait : new treatment strategies , 1986 .

[17]  T J Roberts,et al.  Muscular Force in Running Turkeys: The Economy of Minimizing Work , 1997, Science.

[18]  J. Kauer,et al.  Neural control of hatching: fate of the pattern generator for the leg movements of hatching in post-hatching chicks , 1984, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[19]  A. Bekoff,et al.  Constrained and flexible features of rhythmical hindlimb movements in chicks: kinematic profiles of walking, swimming and airstepping. , 1992, The Journal of experimental biology.

[20]  N. Heglund,et al.  Speed, stride frequency and energy cost per stride: how do they change with body size and gait? , 1988, The Journal of experimental biology.

[21]  D. F. Hoyt,et al.  Gait and the energetics of locomotion in horses , 1981, Nature.

[22]  L. Tesio,et al.  Motion of the center of gravity of the body in clinical evaluation of gait. , 1985, American journal of physical medicine.