Exercise enhances axonal growth and functional recovery in the regenerating spinal cord

We investigated whether enhancing locomotory activity could accelerate the axonal growth underlying the significant recovery of function after a complete spinal transection in the eel, Anguilla. Eels with low spinal transections (at about 60% body length) were kept in holding tanks, where they were inactive, or made to swim continually against a water current at about one body length/s. Their locomotion was periodically assessed by measuring tail beat frequencies at different swimming speeds. Axonal growth was determined from anterograde labeling with 1,1'-diotadecyl-3,3,3',3'-tetramethylindocarbocyanine perchlorate, inserted postmortem into the spinal cord, just rostral to the transection. Twenty days after surgery, there were significantly more labeled growth cones more than 2 mm caudal from the transection in the exercised fish (74.6+/-2.3%; cf. 34.5+/-1.1%). This difference was still observed at 40 days (57.9+/-1.6% cf. 42.1+/-2% >2 mm), but the regenerated axons were of similar maximum lengths by 120 days (9.8+/-0.3 cf. 7.7+/-2.8 mm). After surgery, each eel undulated its whole body faster at any given swimming speed, thus changing the linear relationship between tail beat frequency and forward speed established before transection. The slope increased by up to 112.5+/-27.4% over the first 8 days post-surgery in inactive animals, while a smaller rise (45.6+/-10.5%) was observed in exercised fish during this period. Thereafter, the slope progressively declined to pre-surgery levels in both groups of animals, but the recovery occurred within 20+/-4 days in exercised eels, as opposed to 40+/-5 days in inactive fish. The locomotory performance of sham-operated fish was unaffected by 10 days of continual locomotion and remained similar to that of naïve eels, pre-transection. These data show that elevated locomotory activity enhances axonal growth and accelerates recovery of locomotory function.