Key points• Neuroscientists often suggest that we adapt our movements to minimize energy use; however, recent studies have provided conflicting evidence in this regard.• In the present study, we show that motor learning robustly increases the economy of locomotion during split-belt treadmill adaptation.• We also demonstrate that reductions in metabolic power scale with the magnitude of adaptation and are also associated with a reduction in muscle activity throughout the lower limbs.• Our results provide strong evidence that increasing economy may be a key criterion driving the systematic changes in co-ordination during locomotor adaptation.• These findings may also facilitate the design of novel interventions to improve locomotor learning in stroke survivors.Abstract Many theories of motor control suggest that we select our movements to reduce energy use. However, it is unclear whether this process underlies short-term motor adaptation to novel environments. Here we asked whether adaptation to walking on a split-belt treadmill leads to a more economical walking pattern. We hypothesized that adaptation would be accompanied by a reduction in metabolic power and muscle activity and that these reductions would be temporally correlated. Eleven individuals performed a split-belt adaptation task where the belt speeds were set at 0.5 and 1.5 m s −1 . Adaptation was characterized by step length symmetry, which is the normalized difference in step length between the legs. Metabolic power was calculated based on expired gas analysis, and surface EMG was used to record the activity of four bilateral leg muscles (tibialis anterior, lateral gastrocnemius, vastus lateralis and biceps femoris). All participants initially walked with unequal step lengths when the belts moved at different speeds, but gradually adapted to take steps of equal length. Additionally, net metabolic power was reduced from early adaptation to late adaptation (early, 3.78 ± 1.05 W kg −1 ; and late, 3.05 ± 0.79 W kg −1 ; P < 0.001). This reduction in power was also accompanied by a bilateral reduction in EMG throughout the gait cycle. Furthermore, the reductions in metabolic power occurred over the same time scale as the improvements in step length symmetry, and the magnitude of these improvements predicted the size of the reduction in metabolic power. Our results suggest that increasing economy may be a key criterion driving locomotor adaptation.

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We reasoned that with an optimal aiding horizontal force, the reduction in metabolic rate would reflect the cost of generating propulsive forces during normal walking. Furthermore, the reductions in ankle extensor electromyographic (EMG) activity would indicate the propulsive muscle actions. We applied horizontal forces at the waist, ranging from 15% body weight aiding to 15% body weight impeding, while subjects walked at 1.25 m/s. With an aiding horizontal force of 10% body weight, 1) the net metabolic cost of walking decreased to a minimum of 53% of normal walking, 2) the mean EMG of the medial gastrocnemius (MG) during the propulsive phase decreased to 59% of the normal walking magnitude, and yet 3) the mean EMG of the soleus (Sol) did not decrease significantly. Our data indicate that generating horizontal propulsive forces constitutes nearly half of the metabolic cost of normal walking. Additionally, it appears that the MG plays an important role in forward propulsion, whereas the Sol does not.

To investigate the metabolic cost and muscular actions required for the initiation and propagation of leg swing, we applied a novel combination of external forces to subjects walking on a treadmill. We applied a forward pulling force at each foot to assist leg swing, a constant forward pulling force at the waist to provide center of mass propulsion, and a combination of these foot and waist forces to evaluate leg swing. When the metabolic cost and muscle actions were at a minimum, the condition was considered optimal. We reasoned that the difference in energy consumption between the optimal combined waist and foot force trial and the optimal waist force-only trial would reflect the metabolic cost of initiating and propagating leg swing during normal walking. We also reasoned that a lower muscle activity with these assisting forces would indicate which muscles are normally responsible for initiating and propagating leg swing. With a propulsive force at the waist of 10% body weight (BW), the net metabolic cost of walking decreased to 58% of normal walking. With the optimal combination, a propulsive force at the waist of 10% BW plus a pulling force at the feet of 3% BW the net metabolic cost of walking further decreased to 48% of normal walking. With the same combination, the muscle activity of the iliopsoas and rectus femoris muscles during the swing phase was 27 and 60% lower, respectively, but the activity of the medial gastrocnemius and soleus before swing did not change. Thus our data indicate that approximately 10% of the net metabolic cost of walking is required to initiate and propagate leg swing. Additionally, the hip flexor muscles contribute to the initiation and propagation leg swing.

The sensory and neural mechanisms underlying postural control have received much attention in recent decades but remain poorly understood. Our objectives were 1) to establish the decerebrate cat as an appropriate model for further research into the sensory mechanisms of postural control and 2) to observe what elements of the postural response can be generated by the brain stem and spinal cord. Ten animals were decerebrated using a modified premammillary technique, which consists of a premammillary decerebration that is modified with a vertical transection near the subthalamic nucleus to eliminate spontaneous locomotion. Horizontal support surface perturbations were applied to all four limbs and electromyographic recordings were collected from 14 muscles of the right hindlimb. Muscle activation was quantified with tuning curves, which compared increases and decreases in muscle activity to background and graphed the difference against perturbation direction. Parallels were drawn between these tuning curves, which were further quantified with a principal direction and breadth (range of directions of muscle activation), and data collected by other researchers from the intact animal. We found a strong similarity in the direction and breadth of the tuning curves generated in the decerebrate and intact cat. These results support our hypothesis that directionally specific tuning of muscles in response to support surface perturbations does not require the cortex, further indicating a strong role for the brain stem and spinal cord circuits in mediating directionally appropriate muscle activation patterns.

Teasing preparations of cat extraocular muscles (EOM) were used to study the arrangement of muscle fibers and the distribution of the different cholinesterase-positive sites, i.e. (1) large motor endplates, (2) small motor endings of the 'en grappe' type, (3) myotendinous junctions and (4) myomyous junctions. The distribution of these cholinesterase-positive structures gives clear evidence of a complex muscle architecture of cat EOM. In the global layer of cat EOM, only multiply innervated muscle fibers run the whole length of the muscle. The focally innervated muscle fibers are generally shorter; they are usually arranged in series of two to three fibers being interconnected by myomyous junctions. Moreover, muscle fiber splitting is frequently present resulting in a netlike arrangement of muscle fibers. Most of the myomyous junctions occur between focally innervated muscle fibers, but also end-to-side connections of focally to multiply innervated muscle fibers are observed; multiply innervated muscle fi0ers have not been found connected to each other. In this layer, large motor endplates are distributed in several bands between origin and insertion. In the orbital layer all muscle fibers run from tendon to tendon, focally as well as multiply innervated ones. Here, large motor endplates are confined to a band in the middle of the muscle, and myomyous junctions are generally absent. Some functional implications of this complex architecture of cat EOM are discussed.

extent of symmetry. As expected, during level walking, the GPE and KE curves were out of phase, of similar magnitude, and nearly mirror images so that the fluctuations in combined (GPE+KE) energy were attenuated. During downhill walking, the fluctuations in the combined energy of the center of mass were smaller than those on the level, i.e. mechanical energy exchange was more effective. During uphill walking, the fluctuations in the combined energy of the center of mass were larger than those on the level, i.e. mechanical energy exchange was less effective. Mechanical energy exchange occurred during downhill, level and uphill walking, but it was most effective during downhill walking.

mechanics during walking do not necessarily correspond to high mechanical work and may not result in a high metabolic cost.