Coordination strategies for limb forces during weight-bearing locomotion in normal rats, and in rats spinalized as neonates.

Some rats spinally transected as neonates (ST rats) achieve weight-supporting independent locomotion. The mechanisms of coordinated hind-limb weight support in such rats are not well understood. To examine these we compared ST rats (with better than 60% of weight supported steps) and normal rats that were trained to cross an instrumented runway. Ground reaction forces, coordination of hind-limb and forelimb forces and the motions of the center of pressure (CoP) were assessed. Normal rats crossed the runway with a diagonal trot. On average hind-limbs bore about 80% of the vertical load carried by forelimbs (45% body weight on hind-limbs 55% on forelimbs), although this varied. Forelimbs and hind-limbs acted synergistically to generate decelerative and propulsive rostrocaudal forces, which averaged 15% of body weight with maximums of 50%. Lateral forces were very small (<8% of body weight). Center of pressure progressed in jumps along a straight line with mean lateral deviations <1 cm. ST rats hind-limbs bore about 60% of the vertical load of forelimbs (37% body weight on hind-limbs, 63% on forelimbs), significantly less compared to intact rats (P < 0.05). ST rats showed similar mean rostrocaudal forces, but with significantly larger maximum fluctuations of up to 80% of body weight (P < 0.05). Joint force-plate recordings showed forelimbs and hind-limb rostrocaudal forces in ST rats were opposing and significantly different from intact rats (P < 0.05). Lateral forces were approximately 20% of body weight and significantly larger than in normal rats (P < 0.05). Center of pressure zig-zagged, with mean lateral deviations of approximately 2 cm and a significantly larger range (P < 0.05). The haunches were also observed to roll more than normal rats. The locomotor strategy of injured rats using limbs in opposition was presumably less efficient but their complex gait was statically stable. Because forelimbs and hind-limbs acted in opposition, the trunk was held compressed. Force coordination was likely managed largely by the voluntary control in forelimbs and trunk.

Trunk sensorimotor cortex is essential for autonomous weight-supported locomotion in adult rats spinalized as P1/P2 neonates.

Unlike adult spinalized rats, approximately 20% of rats spinalized as postnatal day 1 or 2 (P1/P2) neonates achieve autonomous hindlimb weight support. Cortical representations of mid/low trunk occur only in such rats with high weight support. However, the importance of hindlimb/trunk motor cortex in function of spinalized rats remains unclear. We tested the importance of trunk sensorimotor cortex in their locomotion using lesions guided by cortical microstimulation in P1/P2 weight-supporting neonatal spinalized rats and controls. In four intact control rats, lesions of hindlimb/trunk cortex caused no treadmill deficits. All spinalized rats lesioned in trunk cortex (n = 16: 4 transplant, 6 transect, 6 transect + fibrin glue) lost an average of about 40% of their weight support. Intact trunk cortex was essential to their level of function. Lesion of trunk cortex substantially increased roll of the hindquarters, which correlated to diminished weight support, but other kinematic stepping parameters showed little change. Embryonic day 14 (E14) transplants support development of the trunk motor representations in their normal location. We tested the role of novel relay circuits arising from the grafts in such cortical representations in E14 transplants using the rats that received (noncellular) fibrin glue grafting at P1/P2 (8 allografts and 32 xenografts). Fibrin-repaired rats with autonomous weight support also had trunk cortical representations similar to those of E14 transplant rats. Thus acellular repair and intrinsic plasticity were sufficient to support the observed features. Our data show that effective cortical mechanisms for trunk control are essential for autonomous weight support in P1/P2 spinalized rats and these can be achieved by intrinsic plasticity.

Motor strategies used by rats spinalized at birth to maintain stance in response to imposed perturbations.

Some rats spinalized P1/P2 achieve autonomous weight-supported locomotion and quiet stance as adults. We used force platforms and robot-applied perturbations to test such spinalized rats (n = 6) that exhibited both weight-supporting locomotion and stance, and also normal rats (n = 8). Ground reaction forces in individual limbs and the animals' center of pressure were examined. In normal rats, both forelimbs and hindlimbs participated actively to control horizontal components of ground reaction forces. Rostral perturbations increased forelimb ground reaction forces and caudal perturbations increased hindlimb ground reaction forces. Operate rats carried 60% body weight on the forelimbs and had a more rostral center of pressure placement. The pattern in normal rats was to carry significantly more weight on the hindlimbs in quiet stance (roughly 60%). The strategy of operate rats to compensate for perturbations was entirely in forelimbs; as a result, the hindlimbs were largely isolated from the perturbation. Stiffness magnitude of the whole body was measured: its magnitude was hourglass shaped, with the principal axis oriented rostrocaudally. Operate rats were significantly less stiff--only 60-75% of normal rats' stiffness. The injured rats adopt a stance strategy that isolates the hindlimbs from perturbation and may thus prevent hindlimb loadings. Such loadings could initiate reflex stepping, which we observed. This might activate lumbar pattern generators used in their locomotion. Adult spinalized rats never achieve independent hindlimb weight-supported stance. The stance strategy of the P1 spinalized rats differed strongly from the behavior of intact rats and may be difficult for rats spinalized as adults to master.