Biology subjects

Cusumano, J. P.

Publications and source records attributed to Cusumano, J. P..

13 recordsLinked to original sources

bioRxiv2026

How Visual Context Influences Lateral Stepping Regulation While Walking on Winding Paths

Goal-directed walking involves regulating foot placements to achieve specific tasks. This requires visuomotor integration. Perceptual, cognitive, and contextual salience guide attention and motor planning for navigation. Here, we quantified how perceptual salience informs lateral foot placement while walking. Participants walked along prescribed virtual paths (straight or winding), thus keeping contextual salience (the task itself) constant. We manipulated perceptual salience by systematically altering environment richness (rich vs. sparse) and path color contrast (high vs. low). We thus tested the extent to which stepping control is determined only by the walking path (i.e. its shape), or is also influenced by task-relevant salience of the paths (only), and/or the salience of the surrounding environment. We quantified head pitch angle to approximate gaze direction. We quantified lateral stepping regulation from a Goal Equivalent Manifold framework. On winding (vs. straight) paths, participants looked down more (lower mean head pitch) and more consistently (less variable head pitch). They took narrower, more variable steps and corrected stepping errors more strongly. This confirmed the predominant influence of task (contextual salience) on stepping control. On low (vs. high) contrast paths, participants looked down more and exhibited greater variability in lateral position on their path with weaker error correction. Higher contrast paths elicited stronger and more consistent stepping control. Walking in sparse (vs. rich) environments yielded somewhat less consistent, but still significant changes in head pitch and stepping regulation. Salience manipulations affected stepping differently on straight versus winding paths. Therefore, perceptual salience influences step-to-step control during naturalistic walking.

neuroscience↗

bioRxiv2026

Visual Context Influences Lateral Balance While Walking on Winding Paths

Effectively navigating daily environments necessitates achieving adaptability while maintaining stability. This requires integrating sensory feedback, primarily from the visual system, to maintain balance and maneuverability as we walk. This study examined the impact of visual information salience on lateral balance and stepping. Twenty-eight healthy adults (16F/12M; 26.16{+/-}4.23 years) participated. Participants walked along each of two virtual paths (Straight vs. Winding), having each of two path color contrasts (High vs. Low), in each of two environments with differing visual richness (Rich vs. Sparse). We quantified stepping errors as the percentage of steps landing outside designated path boundaries. We computed means () and standard deviations ({sigma}) of the minimum mediolateral margins of stability (MoSL), and we computed lateral Probability of Instability (PoIL) to assess participants risk of taking unstable (MoSL < 0) steps. On Straight paths, participants made more stepping errors on Low (vs. High) contrast paths for both environments, and exhibited decreased{sigma} (MoSL) in Sparse (vs. Rich) environments on paths of both visual contrasts. On Winding paths, participants made the most stepping errors on Low Contrast paths in Sparse environments. They walked with smaller (MoSL) and exhibited higher PoIL on Low Contrast paths in both environments, and they exhibited higher PoIL in Sparse environments on paths of both visual contrasts. The effects of diminished visual information were far more pronounced on Winding paths (vs. Straight), hindering performance and balance maintenance, as these conditions challenged both mechanical and sensory mechanisms underlying balance control.

neuroscience↗

bioRxiv2024

Probability of Lateral Instability While Walking on Winding Paths

People with balance impairments often struggle performing turns or lateral maneuvers, which can increase risk of falls and injuries. Here we asked how peoples mediolateral balance is impacted when walking on non-straight winding paths. Twenty-four healthy adults (12F/12M; 25.8{+/-}3.5 yrs) participated. Each walked on each of six paths projected onto a treadmill, comprised of three pseudo-random path oscillation frequency combinations (straight, slowly-winding, quickly-winding), each presented at either wide or narrow width. We quantified stepping errors as the percent of steps taken off each path. We quantified minimum mediolateral Margin of Stability (MoSL) at each step and calculated means () and standard deviations ({sigma}) for each trial. We calculated lateral Probability of Instability (PoIL) as participants statistical risk of taking unstable (MoSL < 0) steps. On narrower paths, participants made more stepping errors and walked with smaller (MoSL) on all paths (p < 0.001), and exhibited increased PoIL on the straight and slowly-winding paths (p < 0.001). On winding paths, participants made progressively more stepping errors and walked with smaller (MoSL) for increasingly sinuous narrow paths (all p < 0.001) and wide quickly-winding paths (all p < 0.001). They also consistently walked with larger{sigma} (MoSL), and increased PoIL on higher sinuosity paths of both widths (all p < 0.001). Though many took numerous unstable steps, no participant fell. Our results demonstrate healthy adults ability both to trade off increased risk of lateral instability for greater maneuverability, and to employ highly-versatile stepping strategies to maintain balance while walking.

bioengineering↗

bioRxiv2024

Adapting Lateral Stepping Control to Walk on Winding Paths

Most often, gait biomechanics is studied during straight-ahead walking. However, real-life walking imposes various lateral maneuvers people must navigate. Such maneuvers challenge peoples lateral balance and can induce falls. Determining how people regulate their stepping movements during such complex walking tasks is therefore essential. Here, 24 adults (12F/12M; Age 25.8{+/-}3.5yrs) walked on wide or narrow virtual paths that were either straight, slowly-winding, or quickly-winding. From each trial, we computed time series of participants step widths and their lateral body positions relative to their path. We applied our Goal Equivalent Manifold framework - an analysis of how task-level redundancy impacts motor regulation - to quantify how participants adjusted their step width and lateral position from step to step as they walked on these paths. On the narrower paths, participants walked with narrower steps and less lateral position and step width variability. They did so by correcting step-to-step deviations in lateral position more, while correcting step-to-step deviations in step width less. On the winding paths, participants took both narrower and more variable steps. Interestingly, on slowly-winding paths, participants corrected step-to-step deviations in step width more by correcting step-to-step deviations in lateral position less: i.e., they prioritized maintaining step width over position. Conversely, on quickly-winding paths, participants strongly corrected step-to-step deviations in both step width and lateral position: i.e., they prioritized maintaining both approximately equally, consistent with trying to maximize their maneuverability. These findings have important implications for persons with gait impairments who may have elevated fall risk.

bioengineering↗

bioRxiv2024

A model of task-level human stepping regulation yields semistable walking

A simple lateral dynamic walker, with swing leg dynamics and three adjustable input parameters, is used to study how motor regulation affects frontal plane stepping. Motivated by experimental observations and phenomenological models, we imposed task-level multiobjective regulation targeting the walkers optimal lateral foot placement at each step. The regulator prioritizes achieving step width and lateral body position goals to varying degrees by choosing a mixture parameter. Our model thus integrates a lateral mechanical template, which captures fundamental mechanics of frontal-plane walking, with a lateral motor regulation template, an empirically verified model of how humans manipulate lateral foot placements in a goal-directed manner. The model captures experimentally observed stepping fluctuation statistics and demonstrates how linear empirical models of stepping dynamics can emerge from first-principles nonlinear mechanics. We find that task-level regulation gives rise to a goal equivalent manifold in the systems extended state space of mechanical states and inputs, a subset of which contains a continuum of period-1 gaits forming a semistable set: perturbations off of any of its gaits result in transients that return to the set, though typically to different gaits.

bioengineering↗

bioRxiv2024

How Older Adults Maintain Lateral Balance While Walking on Narrowing Paths

BackgroundOlder adults have difficulty maintaining side-to-side balance while navigating daily environments. Losing balance in such circumstances can lead to falls. We need to better understand how older adults adapt lateral balance to navigate environment-imposed task constraints. Research QuestionHow do older adults adjust mediolateral balance while walking along continually-narrowing paths, and what are the stability implications of these adjustments? MethodsEighteen older (71.6{+/-}6.0 years) and twenty younger (21.7{+/-}2.6 years) healthy adults traversed 25m-long paths that gradually narrowed from 45cm to 5cm. Participants switched onto an adjacent path when they chose. We quantified participants lateral center-of-mass dynamics and lateral Margins of Stability (MoSL) as paths narrowed. We quantified lateral Probability of Instability (PoIL) as the cumulative probability that participants would take a laterally unstable (MoSL<0) step as they walked. We also extracted these outcomes where participants switched paths. ResultsAs paths narrowed, all participants exhibited progressively smaller average MoSL and increasingly larger PoIL. However, their MoSL variability was largest at both the narrowest and widest path sections. Older adults exhibited consistently both larger average and more variable MoSL across path widths. Taken into account together, these resulted in either comparable or somewhat larger PoIL as paths narrowed. Older adults left the narrowing paths sooner, on average, than younger. As they did so, older adults exhibited significantly larger average and more variable MoSL, but somewhat smaller PoIL than younger. SignificanceOur results directly challenge the predominant interpretation that larger average MoSL as indicating "greater stability", which we argue is inconsistent with the principles underlying its derivation. In contrast, analyzing step-to-step gait dynamics, together with estimating PoIL allows one to properly quantify instability risk. Furthermore, the adaptive strategies uncovered using these methods suggest potential targets for future interventions to reduce falls in older adults.

bioengineering↗

bioRxiv2023

Generalizing Stepping Concepts To Non-Straight Walking

People rarely walk in straight lines. Instead, we make frequent turns or other maneuvers. Spatiotemporal parameters fundamentally characterize gait. For straight walking, these parameters are well-defined for that task of walking on a straight path. Generalizing these concepts to non-straight walking, however, is not straightforward. People also follow non-straight paths imposed by their environment (store aisle, sidewalk, etc.) or choose readily-predictable, stereotypical paths of their own. People actively maintain lateral position to stay on their path and readily adapt their stepping when their path changes. We therefore propose a conceptually coherent convention that defines step lengths and widths relative to known walking paths. Our convention simply re-aligns lab-based coordinates to be tangent to a walkers path at the mid-point between the two footsteps that define each step. We hypothesized this would yield results both more correct and more consistent with notions from straight walking. We defined several common non-straight walking tasks: single turns, lateral lane changes, walking on circular paths, and walking on arbitrary curvilinear paths. For each, we simulated idealized step sequences denoting "perfect" performance with known constant step lengths and widths. We compared results to path-independent alternatives. For each, we directly quantified accuracy relative to known true values. Results strongly confirmed our hypothesis. Our convention returned vastly smaller errors and introduced no artificial stepping asymmetries across all tasks. All results for our convention rationally generalized concepts from straight walking. Taking walking paths explicitly into account as important task goals themselves thus resolves conceptual ambiguities of prior approaches.

neuroscience↗

bioRxiv2023

How Older Adults Regulate Lateral Stepping on Narrowing Walking Paths

Walking humans often navigate complex, varying walking paths. To reduce falls, we must first determine how older adults purposefully vary their steps in contexts that challenge balance. Here, 20 young (21.7{+/-}2.6 yrs) and 18 older (71.6{+/-}6.0 yrs) healthy adults walked on virtual paths that slowly narrowed (from 45 cm to as narrow as 5 cm). Participants could switch onto an "easier" path whenever they chose. We applied our Goal Equivalent Manifold framework to quantify how participants adjusted their lateral stepping variability and step-to-step corrections of step width and lateral position as these paths narrowed. We also extracted these characteristics where participants switched paths. As paths narrowed, all participants reduced their lateral stepping variability, but older adults less so. To stay on the narrowing paths, young adults increasingly corrected step-to-step deviations in lateral position more, by correcting step-to-step deviations in step width less. Conversely, as older adults also increasingly corrected lateral position deviations, they did so without sacrificing correcting step-to-step deviations in step width, presumably to preserve balance. While older adults left the narrowing paths sooner, several of their lateral stepping characteristics remained similar to those of younger adults. While older adults largely maintained overall walking performance per se, they did so by changing how they balanced the competing stepping regulation requirements intrinsic to the task: maintaining position vs. step width. Thus, balancing how to achieve multiple concurrent stepping goals while walking provides older adults the flexibility they need to appropriately adapt their stepping on continuously narrowing walking paths.

bioengineering↗

bioRxiv2023

How Healthy Older Adults Enact Lateral Maneuvers While Walking

BackgroundWalking requires frequent maneuvers to navigate changing environments with shifting goals. Humans accomplish maneuvers and simultaneously maintain balance primarily by modulating their foot placement, but a direct trade-off between these two objectives has been proposed. As older adults rely more on foot placement to maintain lateral balance, they may be less able to adequately adapt stepping to perform lateral maneuvers. Research QuestionHow do older adults adapt stepping to enact lateral lane-change maneuvers, and how do physical and perceived ability influence their task performance? MethodsTwenty young (21.7 {+/-} 2.6 yrs) and 18 older (71.6 {+/-} 6.0 yrs) adults walked on a motorized treadmill in a virtual environment. Following an audible and visual cue, participants switched between two parallel paths, centered 0.6m apart, to continue walking on their new path. We quantified when participants initiated the maneuver following the cue, as well as their step width, lateral position, and stepping variability ellipses at each maneuver step. ResultsYoung and older adults did not differ in when they initiated the maneuver, but participants with lower perceived ability took longer to do so. Young and older adults also did not exhibit differences in step width or lateral positions at any maneuver step, but participants with greater physical ability reached their new path faster. While only older adults exhibited stepping adaptations prior to initiating the maneuver, both groups traded-off stability for maneuverability to enact the lateral maneuver. SignificancePhysical and perceived balance ability, rather than age per se, differentially influenced maneuver task performance. Humans must make decisions related to the task of walking itself and do so based on both physical and perceived factors. Understanding and targeting these interactions may help improve walking performance among older adults.

neuroscience↗

bioRxiv2022

Humans Adapt Multi-Objective Control of Stepping To Perform Lateral Maneuvers While Walking

To successfully traverse their environment, humans often perform maneuvers to achieve desired task goals while simultaneously maintaining balance. Humans accomplish these tasks primarily by modulating their foot placements. As humans are more unstable laterally, we must better understand how humans modulate lateral foot placement. We previously developed a theoretical framework and corresponding computational models to describe how humans regulate lateral stepping during straight-ahead continuous walking. We identified goal functions for step width and lateral body position that define the walking task and determine the set of all possible task solutions as Goal Equivalent Manifolds (GEMs). Here, we used this framework to determine if humans can regulate lateral stepping during non-steady-state lateral maneuvers by minimizing errors consistent with these goal functions. Twenty young healthy adults each performed four lateral lane-change maneuvers in a virtual reality environment. Extending our general lateral stepping regulation framework, we first re-examined the requirements of such transient walking tasks. Doing so yielded new theoretical predictions regarding how steps during any such maneuver should be regulated to minimize error costs, consistent with the goals required at each step and with how these costs are adapted at each step during the maneuver. Humans performed the experimental lateral maneuvers in a manner consistent with our theoretical predictions. Furthermore, their stepping behavior was well modeled by allowing the parameters of our previous lateral stepping models to adapt from step to step. To our knowledge, our results are the first to demonstrate humans might use evolving cost landscapes in real time to perform such an adaptive motor task and, furthermore, that such adaptation can occur quickly - over only one step. Thus, the predictive capabilities of our general stepping regulation framework extend to a much greater range of walking tasks beyond just normal, straight-ahead walking. AUTHOR SUMMARYWhen we walk in the real world, we rarely walk continuously in a straight line. Indeed, we regularly have to perform other tasks like stepping aside to avoid an obstacle in our path (either fixed or moving, like another person coming towards us). While we have to be highly maneuverable to accomplish such tasks, we must also maintain balance to avoid falling while doing so. This is challenging because walking humans are inherently more unstable side-to-side. Sideways falls are particularly dangerous for older adults as they can lead to hip fractures. Here, we establish a theoretical basis for how people might accomplish such maneuvers. We show that humans execute a simple lateral lane-change maneuver consistent with our theoretical predictions. Importantly, our simulations show they can do so by adapting at each step the same step-to-step regulation strategies they use to walk straight ahead. Moreover, these same control processes also explain how humans trade-off side-to-side stability to gain the maneuverability they need to perform such lateral maneuvers.

neuroscience↗

bioRxiv2022

Humans trade-off energetic cost with fatigue avoidance while walking

Metabolic cost minimization is thought to underscore the neural control of locomotion. Yet, avoiding high muscle activation, a cause of fatigue, often outperforms energy minimization in computational predictions of human gait. Discerning the relative importance of these criteria in human walking has proved elusive, in part, because they have not been empirically decoupled. Here we explicitly decouple whole-body metabolic cost and fatigue-like muscle activation costs (estimated from electromyography) by pitting them against one another using two distinct gait tasks. When experiencing these competing costs, participants (N=10) chose the task that avoided overburdening muscles (fatigue avoidance) at the expense of higher metabolic power (p<0.05). Muscle volume-normalized activation more closely models energy use and was also minimized by the participants decision (p<0.05), demonstrating that muscle activation was, at best, an inaccurate signal for metabolic energy. Energy minimization was only observed when there was no adverse effect on muscle activation costs. By decoupling whole-body metabolic and muscle activation costs, we provide among the first empirical evidence of humans embracing non-energetic optimality in favor of a clearly defined neuromuscular objective. This finding indicates that local muscle fatigue and effort may well be key factors dictating human walking behavior and its evolution.

physiology↗

bioRxiv2022

Viability, task switching, and fall avoidance of the simplest dynamic walker

Humans display great versatility when performing goal-directed tasks while walking. However, the extent to which such versatility helps with fall avoidance remains unclear. We recently demonstrated a functional connection between the motor regulation needed to achieve task goals (e.g. maintaining walking speed) and a simple walkers ability to reject large disturbances. Here, for the same model, we identify the viability kernel--the state space region in which the walker can step forever via at least one sequence of push-off inputs per state. We further find that only a few basins of attraction of the speed-regulated walkers steady-state gaits can fully cover the viability kernel. This highlights a potentially important role of task-level motor regulation in fall avoidance. Therefore, we posit an adaptive hierarchical control/regulation strategy that switches between different task-level regulators to avoid falls. Our hierarchical task switching controller only requires a target value of the regulated observable--a task switch--at each walking step, each chosen from a small, predetermined collection. Because humans have typically already learned to perform such tasks during nominal walking conditions, this suggests that the information cost of biologically implementing such controllers for the nervous system, including cognitive demands in humans, could be relatively low.

bioengineering↗

bioRxiv2021

Rethinking Margin of Stability: Incorporating Step-To-Step Regulation to Resolve the Paradox

Derived from inverted pendulum dynamics, mediolateral Margin of Stability (MoSML) is a mechanically-grounded measure of instantaneous stability. However, average MoSML measures yield paradoxical results. Gait pathologies or perturbations often induce larger (supposedly "more stable") average MoSML, despite clearly destabilizing factors. However, people do not walk "on average" - they walk (and sometimes lose balance) one step at a time. We assert the paradox arises because averaging discards step-to-step dynamics. We present a framework unifying the inverted pendulum with Goal-Equivalent Manifold (GEM) analyses. We identify in the pendulums center-of-mass dynamics constant-MoSML manifolds, including one candidate "stability GEM" signifying the goal to maintain some constant [Formula]. We used this framework to assess step-to-step MoSML dynamics of humans walking in destabilizing environments. While goal-relevant deviations were readily corrected, humans did not exploit equifinality by allowing deviations to persist along this GEM. Thus, maintaining a constant [Formula] is inconsistent with observed step-to-step fluctuations in center-of-mass states. Conversely, the extent to which participants regulated fluctuations in foot placements strongly predicted regulation of center-of-mass fluctuations. Thus, center-of-mass dynamics may arise indirectly as a consequence of regulating mediolateral foot placements. To resolve the paradox caused by averaging MoSML, we present a new statistic, Probability of Instability (PoIL), to predict instability likelihood. Participants exhibited increased PoIL when destabilized (p = 9.45x10-34), despite exhibiting larger ("more stable") average MoSML (p = 1.70x10-15). Thus, PoIL correctly captured peoples increased risk of losing lateral balance, whereas average MoSML did not. PoIL also explains why peoples average MoSML increased in destabilizing contexts.

bioengineering↗