18 19 20 21 22 23 24 provide further insight into how the healthy and impaired nervous system regulates dynamic 49 balance during walking. Introduction 51 Bipedal locomotion is inherently unstable due to the small base of support, long single-52 limb support times, and sensorimotor transmission delays [1]. As a result, we must frequently 53 generate corrective responses to maintain balance in response to both internal and external 54 perturbations [2,3]. For example, to recover from unexpected perturbations such as slips or trips 55 while walking, the nervous system generates reactive control strategies involving simultaneous, 56 coordinated responses of both the upper and lower limbs [4,5]. These reactive, interlimb 57 responses to perturbations can restore stability by generating changes in angular momentum that 58 counteract the body's rotation toward the ground. 59 One conventional method to capture whole-body rotational dynamics during perturbation 60 responses is to compute whole-body angular momentum (WBAM). WBAM reflects the net 61 influence of all the body segments' rotation relative to a specified axis, which is commonly taken 62 to project through the body's center of mass [6-8]. WBAM is highly regulated as its value 63 remains close to zero during normal, unperturbed walking [9,10]. During perturbed walking, 64 angular momentum dramatically deviates from that measured during unperturbed walking [6,7], 65 and this deviation captures the features of body rotation that, if not arrested, would lead to a fall.66 To regain balance when encountering unexpected perturbations, the central nervous system 67 activates muscles to accelerate body segments and restore angular momentum across multiple 68 recovery steps [11,12].
69Angular momentum can also capture balance impairments in populations with gait 70 asymmetries and sensorimotor deficits such as amputees and stroke survivors. These individuals 71 often have a higher peak-to-peak range of angular momentum than healthy controls [13][14][15][16], and 72 the presence of gait asymmetries may contribute to balance impairments in these populations. 73 For example, the magnitude of step length asymmetry in people-post stroke is negatively 74 correlated with scores on the Berg Balance Scale, indicating that step length asymmetry is 75 associated with increased fall risk [17]. 76 An important question for clinical researchers is whether there is a causal relationship 77 between gait asymmetry and the ability to maintain balance in response to perturbations during 78 walking. Previous work demonstrated that whole-body dynamics, as measured by WBAM, do 79 not change in response to imposed gait asymmetries in healthy individuals [7]. However, the 80 strategy that the central nervous system uses to stabilize whole-body dynamics remains to be 81 determined. There are two distinct hypotheses capable of explaining the negligible influence of 82 asymmetry on whole-body angular momentum. First, the central nervous system may generate 83 stereotypical, invariant intersegmenta...