Human locomotor adaptation is necessary to maintain flexibility of walking. Several lines of research suggest that the cerebellum plays a critical role in motor adaptation. In this study we investigated the effects of noninvasive stimulation of the cerebellum to enhance locomotor adaptation. We found that anodal cerebellar transcranial direct current stimulation (tDCS) applied during adaptation expedited the adaptive process while cathodal cerebellar tDCS slowed it down, without affecting the rate of de-adaptation of the new locomotor pattern. Interestingly, cerebellar tDCS affected the adaptation rate of spatial but not temporal elements of walking. It may be that spatial and temporal control mechanisms are accessible through different neural circuits. Our results suggest that tDCS could be used as a tool to modulate locomotor training in neurological patients with gait impairments.

New walking patterns can be learned over short timescales (i.e., adapted in minutes) using a split-belt treadmill that controls the speed of each leg independently. This leads to storage of a modified spatial and temporal motor pattern that is expressed as an aftereffect in regular walking conditions. Because split-belt walking is a novel task for adults and children alike, we used it to investigate how motor adaptation matures during human development. We also asked whether the immature pattern resembles that of people with cerebellar dysfunction, because we know that this adaptation depends on cerebellar integrity. Healthy children (3-18 years old) and adults, and individuals with cerebellar damage were adapted while walking on split belts (1:2 speed ratio). Adaptation and de-adaptation rates were quantified separately for temporal and spatial parameters. All healthy children and adults tested could learn the new timing at the same rate and showed significant aftereffects. However, children younger than 6 years old were unable to learn the new spatial coordination. Furthermore, children as old as age 11 years old showed slower rates of adaptation and de-adaptation of spatial parameters of walking. Young children showed patterns similar to cerebellar patients, with greater deficits in spatial versus temporal adaptation. Thus, although walking is a well-practiced, refined motor skill by late childhood (i.e., 11 years of age), the processes underlying learning new spatial relationships between the legs are still developing. The maturation of locomotor adaptation follows at least two time courses, which we propose is determined by the developmental state of the cerebellum.

Adaptation is an error-driven motor learning process that can account for predictable changes in the environment (e.g. walking on ice) or in ourselves (e.g. injury). Our ability to recall and build upon adapted motor patterns across days is essential to this learning process. We investigated how different training paradigms affect the day-to-day memory of an adapted walking pattern. Healthy human adults walked on a split-belt treadmill, and returned the following day to assess recall, re-learning rate, and performance. In the first experiment, one group adapted and de-adapted (i.e. washed-out the learning) several times on day one to practice the initial stage of learning where errors are large; another group adapted only one time and then practiced in the adapted (“learned”) state where errors were small. On day two, they performed washout trials before re-adapting. The group that repeatedly practiced the initial portion of adaptation where errors are large showed the fastest re-learning on the second day. In fact, the memory was nearly as strong as that of a third group that was left overnight in the adapted state and was not washed-out prior to re-exposure on the second day. This demonstrates that alternating exposures to early adaptation and washout can enhance re-adaptation. In the second experiment, we tested whether the opposite split-belt pattern interferes with day two re-learning. Surprisingly, it did not, and instead was similar to practicing in the adapted state. These results show that the structure of the initial phase of learning influences the ease of motor re-learning.

New walking patterns can be learned over short time scales (i.e., adapted in minutes) using a split-belt treadmill that controls the speed of each leg independently. This leads to storage of a modified motor pattern that is expressed as an aftereffect in regular walking conditions and must be de-adapted to return to normal. Here we asked whether the nervous system adapts a general walking pattern that is used across many speeds or a specific pattern affecting only the two speeds experienced during split-belt training. In experiment 1, we tested three groups of healthy adult subjects walking at different split-belt speed combinations and then assessed aftereffects at a range of speeds. We found that aftereffects were largest at the slower speed that was used in split-belt training in all three groups, and it decayed gradually for all other speeds. Thus adaptation appeared to be more strongly linked to the slow walking speed. This result suggests a separation in the functional networks used for fast and slow walking. We tested this in experiment 2 by adapting walking to split belts and then determining how much fast regular walking washed out the slow aftereffect and vice versa. We found that 23-38% of the aftereffect remained regardless of which speed was washed out first. This demonstrates that there is only partial overlap in the functional networks coordinating different walking speeds. Taken together, our results suggest that there are some neural networks for controlling locomotion that are recruited specifically for fast versus slow walking in humans, similar to recent findings in other vertebrates.