Unpowered exoskeletons with springs in parallel to human plantar flexor muscle-tendons can reduce the metabolic cost of walking. We used ultrasound imaging to look 'under the skin' and measure how exoskeleton stiffness alters soleus muscle contractile dynamics and shapes the user's metabolic rate during walking. Eleven participants (4F, 7M; age: 27.7 ± 3.3 years) walked on a treadmill at 1.25 m s −1 and 0% grade with elastic ankle exoskeletons (rotational stiffness: 0-250 Nm rad −1) in one training and two testing days. Metabolic savings were maximized (4.2%) at a stiffness of 50 Nm rad −1. As exoskeleton stiffness increased, the soleus muscle operated at longer lengths and improved economy (force/activation) during early stance, but this benefit was offset by faster shortening velocity and poorer economy in late stance. Changes in soleus activation rate correlated with changes in users' metabolic rate (p = 0.038, R 2 = 0.44), highlighting a crucial link between muscle neuromechanics and exoskeleton performance; perhaps informing future 'muscle-in-the loop' exoskeleton controllers designed to steer contractile dynamics toward more economical force production. The plantar flexors serve an essential role in human walking by providing more than 50% of the leg's total positive mechanical energy 1 and up to 60% of the mechanical power output for redirecting the body's center of mass during push-off 2. The importance of the ankle plantar flexors in legged locomotion is linked to their morphology. Muscle-tendons (MTs) with short pennate muscle fibers and long compliant in-series tendons are well suited for economical locomotion because the elastic tendons store and return mechanical energy over each step 3-5. During steady-state walking, the interaction of the plantar flexors and Achilles tendon is tuned. Throughout early stance, 0% (heel strike) to 40% stride, plantar flexor muscle fascicles generate force isometrically while the Achilles tendon stretches and stores mechanical energy 6. In late stance, 40% to 60% (toe-off) stride, the plantar flexors shorten and the tendon rapidly recoils, providing a burst of positive mechanical power 1,4,7. This coordinated MT interaction permits the plantar flexor muscle fascicles to operate over a narrow region of their force-length (F-L) curve and remain at slow shortening velocities which are favorable contractile conditions for economical force production 7-12. Any disruption to this 'catapult' like mechanism may decrease the ankle's efficient mechanical power production and worsen walking efficiency. Exoskeletons are a class of wearable devices that often act in parallel with human MTs to restore or augment human movement. An increasing number of studies 13 are establishing that both tethered 14-18 and portable 19-21 lower-limb exoskeletons can deliver mechanical power to the body to reduce metabolic demand during walking in young healthy individuals 14-22 , individuals post-stroke 23 , and older adults 24,25. Recently, our group has demonstrated that exoskeletons need not deli...