Skip to main content
PLOS One logoLink to PLOS One
. 2022 Oct 5;17(10):e0268090. doi: 10.1371/journal.pone.0268090

Locomotion control during curb descent: Bilateral ground reaction variables covary consistently during the double support phase regardless of future foot placement constraints

Chuyi Cui 1,#, Ashwini Kulkarni 1,#, Shirley Rietdyk 1,#, Satyajit Ambike 1,*
Editor: Hugh Cowley2
PMCID: PMC9534401  PMID: 36197891

Abstract

During community ambulation, anticipatory adaptations in gait are key for navigating built, populated and natural environments. It has been argued that some instability in gait can be functionally beneficial in situations demanding high maneuverability, and while the mechanisms utilized to maintain locomotor balance are well understood, relatively less is known about how the control of gait stability changes to facilitate upcoming maneuvers in challenging environments. The double support phase may be important in this regard; since both feet can push off the ground simultaneously, there is greater control authority over the body’s movement during this phase. Our goal was to identify how this control authority is exploited to prepare for upcoming maneuvers in challenging environments. We used synergy indices to quantify the degree of coordination between the ground reaction forces and moments under the two feet for stabilizing the resultant force and moment on the body during the double support phase of curb descent. In contrast to our expectations, we observed that the kinetic synergy indices during curb descent were minimally influenced by expected foot targeting maneuvers for the subsequent step. Only the resultant moment in the frontal plane showed reduced stability when targeting was required, but the synergy index was still high, indicating that the resultant moment was stable. Furthermore, the synergy indices indicated that the main function of the ground reaction variables is to maintain stability of whole-body rotations during double support, and this prerogative was minimally influenced by the subsequent foot targeting tasks, likely because the cost of losing balance while descending a curb would be higher than the cost of mis-stepping on a visual target. Our work demonstrates the salience of stabilizing body rotations during curb negotiation and improves our understanding of locomotor control in challenging environments.

1. Introduction

Community ambulation requires gait maneuvers to accommodate changes in the environment, such as altering heading to avoid pedestrians or stationary hazards. In such environments, anticipatory control is key for generating locomotor adaptations, and anticipatory gait changes for safe obstacle navigation have been documented [13]. However, few have examined anticipatory adjustments in unpredictable, complex walking environments [46]. It is generally thought that greater stability during human gait is preferred, and strategies that ensure stability have been described [7]. However, in animal locomotion, some instability is functionally desirable in natural habitats when high maneuverability is important [8, 9]. In humans, anticipatory reduction of stability has been observed in prehension and in standing balance. In prehension behavior, the stability of the force produced by the fingers was reduced when a quick change of the force was expected [10]. In standing balance behavior, the center of pressure was less stable when a postural perturbation was expected [11]. The active reduction of stability of the current motor state is a general feedforward mechanism to facilitate a quick motor change to meet future task demands. It is likely that a similar mechanism, i.e., anticipatory reduction of stability, is used in human gait in challenging environments that require quick maneuvers.

Here, we use the theory of synergies [12, 13] to identify anticipatory changes in the stability of key kinetic variables that determine the body’s motion during gait. We quantify stability using synergy indices obtained from uncontrolled manifold analyses of the ground reaction forces and moments [14]. Our premise is that the resultant force and moment acting at the center of mass (CoM), reflecting the inertial effects arising from the current motion of the body segments and the current forces exerted by the muscles, determine future motion via the equations of motion (assuming minimal interference from other external forces). Therefore, stable body movements require consistent resultant forces and moments. Since any change in the motion must be accomplished by changing the underlying kinetics, anticipatory changes should be evident in the kinetics, and particularly in our synergy indices.

In human gait, the whole-body vertical motion is stabilized by a kinetic synergy during the double support phase of unperturbed, steady-state, level walking [15]. That is, the variations in the vertical ground reaction force (GRF) under one foot are compensated by variations in the vertical GRF under the other foot so that the total force is stabilized. We previously extended this result and observed that the resultant force in the medio-lateral (ML) direction and the resultant moment about all three coordinate axes were stabilized by kinetic synergies during the double support phase of curb negotiation [14]. However, it is unknown whether these synergies will be weakened when an unpredictable environment requires quick maneuvers, as having a strong synergy stabilizing the current motion may be counterproductive. The double support phase may be important in this regard: with both feet in contact with the ground, more kinetic degrees of freedom are available, indicating a greater ability to alter the body’s movements.

The purpose of the present study was to quantify how synergies during double support are modulated in preparation for meeting environmental demands. We used a foot-targeting task to simulate a challenging environment where the person may have to quickly alter their foot placement for the step immediately after stepping down from a curb. We examined the kinetic synergies during curb descent when the expectation of the foot placement for the subsequent step was altered across tasks. Specifically, there was no target for the Baseline task, a stationary target at the preferred foot placement for the Fixed task, and a target that may shift forward or laterally in the Anterior- and Lateral-shift tasks, respectively. We hypothesized that kinetic synergies while stepping down from the curb will be weaker (lower stability) for tasks that required targeting, compared to a task without a target (H1), similar to what has been observed in manual and postural tasks [10, 11]. We also hypothesized that kinetic synergies will be weakened more when expecting a target shift versus no expectation of target shift, because the increase in uncertainty of the task imposes higher maneuverability demands, and lower stability will help in achieving greater maneuverability (H2). We hypothesized that changes in the kinetic synergies will depend on the direction of the expected target shift (H3). For example, the synergies stabilizing motion in the frontal plane (ML translation and rotation about the anterior-posterior, or AP axis) would be preferentially diminished if the individual expected to alter movement in that plane (e.g., take a wider step). In contrast, synergies stabilizing motion in the sagittal plane (AP translation and rotation about the ML axis) would be preferentially diminished if the individual expected to alter movement in that plane (e.g., take a longer step).

2. Methods

2.1 Participants

Twenty-seven young healthy adults participated in this study. Data from three participants were excluded due to bad tracking and one participant was excluded because they consistently skipped down the curb, which led to very brief or no double support phase. Data from 23 young participants were used for subsequent analyses (9 males, age 24.0 ± 3.9 years, weight 74.3 ± 19.0 kg, height 1.69 ± 0.10 m, 1 left leg dominant; assessed using the Waterloo Footedness Questionnaire-Revised [16]). Vision was normal or corrected-to-normal. Exclusion criteria were diagnosed neurological diseases or disorders, musculoskeletal injuries, and need for walking aid. All participants provided written informed consent according to the protocols approved by the Institutional Review Board of Purdue University (Protocol number: IRB-2021-52).

2.2 Protocol

We instructed participants to wear comfortable walking/athletic shoes that they typically wear when walking outside. We decided to study shod walking because young adults likely spend more time walking with shoes than without, especially over uneven terrain. During the experiment, participants walked in their own shoes on an 8 m walkway with an elevated wooden platform or curb (15 cm high, 1 m wide, and 4 m long). We embedded two force plates in the walkway (AMTI Accugait, MA, USA), one in the curb, and one in the ground (Fig 1A). The plates sampled the ground reaction forces and moments at 1000 Hz. A ten-camera motion capture system (Vicon, Oxford, UK) tracked the body’s kinematics at 100 Hz. We placed marker clusters on the lower back and bilateral thigh, shank, and foot segments. We digitized lower-limb joint centers and the toe, heel, and fifth metatarsals of both feet. The force plate and camera data were synchronized using the MotionMonitor software (Innovative Sports Training Inc, IL, USA).

Fig 1.

Fig 1

(A) Experimental Setup. Two force plates were embedded in the walkway, one in the elevated curb, and one in the ground. A projector was used to present visual stepping target on the walkway. (B) Baseline condition. No target was presented. (C) Fixed target condition. The target was presented at the preferred foot landing position and remained stationary. (D, E) Target shift conditions. A target shift in the anterior (D) or lateral (E) direction was triggered with 50% probability when the lead foot contacted the ground force plate (LFCGround). Kinetic synergies were quantified at the double support phase while stepping down using the trials where the target did not shift, as illustrated by the footprints in the initial targets in (D) and (E).

Participants walked at a self-selected speed, stepped down the curb, and continued walking until the end of the walkway. The starting position was identified for each participant during familiarization trials to ensure that participants naturally stepped down the curb with their right foot first. There were four tasks: Baseline, Fixed target, Anterior-shift target, and Lateral-shift target (Fig 1B–1E). These tasks were performed in blocks of 20 or 40 trials; the Baseline task was collected first and the three targeting tasks were block-randomized for each participant. In the Baseline task, participants walked and stepped down the curb with no targets for 20 trials. Average foot placement and step length obtained from these trials determined the target locations for the remaining tasks. The Fixed target task required no anticipation for movement adjustment. In this task, the target was located at the preferred landing position (Fig 1C); it was visible and stationary throughout the trial. Participants were instructed to walk down the walkway as before and step on the target as accurately as possible. This task was performed 20 times. The Anterior- and Lateral-shift tasks required anticipating a possible anterior and lateral maneuver, respectively. For both these tasks, the target was visible at the start of each trial and located at the preferred landing position (Fig 1D and 1E). In 50% of the trials, the target shifted to a new position when the lead foot contacted the ground after crossing the curb (LFCGround, Fig 1A). LFCground was the instant when the vertical GRF exceeded 15 N on the ground force plate. At the beginning of the target shift tasks, participants were told the direction in which the target may shift and were instructed to step on the target as accurately as possible. The forward shift distance was 40% of the average step length [17, 18]. The lateral-shift distance was 20 cm for all participants [17]. The target shift task blocks consisted of 40 trials, with the target shifting for a random set of 20 trials. Participants were given a seated rest break every 20 trials to mitigate fatigue.

The timing of the target shift (at LFCGround, Fig 1A) was set so that the double support phase would include only proactive responses and exclude reactive responses. The longest double support phase in our data across all participants was 193 ms, which is less than the 273 ms reaction time for visual stimuli during walking in young adults [19].

2.3 Data analysis

Pelvis center of mass position, toe and heel positions, GRFs, free moments and center of pressure under both feet were obtained from the MotionMonitor software. Kinematic and kinetic data were filtered with a zero-lag, low pass, 4th-order Butterworth filter with a cut-off frequency of 8 Hz and 20 Hz, respectively.

Spatial-temporal gait measures

We quantified the double support duration, gait speed, step length, step width and foot placement locations. Double support phase was the time interval between LFCGround and the trail foot toe-off from the curb (first instant when vertical GRF on the curb force plate decreased below 15 N). We estimated the whole-body CoM position using the pelvis CoM position. We calculated the instantaneous AP CoM velocity by differentiating the CoM position in the AP direction. Gait speed was the average AP CoM velocity during the double support phase. We used the location of the foot center to quantify foot placement locations for the two steps around the curb and the following target step. The foot center locations were determined as the midpoint of the digitized heel and toe positions when the foot was flat. The foot flat instant was 300 ms after the corresponding heel contacted the ground. Heel contact was identified using the AP heel position [20]. We visually inspected each trial and ensured that the foot was flat at these time instants.

Target stepping performance

The stepping error for the targeting tasks was the length of the vector between the foot center and the target center in the horizontal plane (the vertical coordinate of the foot center was ignored). We quantified the root mean squared error (RMSE) of the vector lengths for each task. We quantified the foot placement RMSE separately for the target-shift trials and the no-shift trials for the Anterior and Lateral-shift tasks. For the Baseline task, we computed the standard deviation in the foot center positions.

Uncontrolled manifold (UCM) analysis

In general, the UCM analysis partitions the across-trial variance in the redundant set of input variables into a component that maintains the performance variable (VUCM) and another orthogonal component that changes the performance variables (VORT) [13]. A synergy index is obtained as the normalized excess of the VUCM over VORT.

We performed the UCM analysis to investigate whether the across-trial variance in the GRF and free moments under the two feet (called the input variables) is structured to maintain the resultant forces and moments (called the performance variables) during the double support phase. We considered the vertical, AP and ML components of resultant force and moments as separate performance variables, and therefore, performed six separate UCM analyses (see S1 Appendix for more details). A resultant force component was determined by the GRFs under both feet during double support. A resultant moment component was determined by the GRFs (and free moment for the resultant moment about vertical axis [14]) under both feet. At every 1% of the double support phase, we quantified the synergy index (ΔV) and its z-transformed value (ΔVz) for each performance variable. Finally, we averaged the ΔVz values over the double support phase to obtain a single synergy index for each performance variable and for each task. Each UCM analysis is associated with a discriminating value ΔVz*; Table 2. A ΔVz greater than the corresponding discriminating value indicates existence of a synergy. A greater ΔVz value indicates a stronger synergy, and vice-versa.

Table 2. One-sample t-tests to determine whether the synergy index (ΔVz) is different from the corresponding discriminating value (ΔVz*) for the three resultant force and the three resultant moment components for each task.
Synergy Axis ΔVz* Task mean t(22) p-value Cohen’s d
Resultant force synergy AP 0 Baseline -0.07 -1.91 0.069
Fixed target -0.07 -2.03 0.055
Anterior -0.07 -1.91 0.069
Lateral -0.07 -3.10 0.005 -0.65
ML 0 Baseline 0.12 3.60 0.002 0.75
Fixed target 0.09 4.38 <0.001 0.91
Anterior 0.11 3.73 0.001 0.78
Lateral 0.09 2.99 0.007 0.62
Vertical 0 Baseline 0.06 1.78 0.089
Fixed target 0.08 3.62 0.002 0.75
Anterior 0.07 1.90 0.071
Lateral 0.08 2.02 0.056
Resultant moment synergy AP 0.55 Baseline 2.04 29.67 <0.001 6.19
Fixed target 1.89 35.27 <0.001 7.35
Anterior 1.86 32.90 <0.001 6.86
Lateral 1.86 33.62 <0.001 7.01
ML 0.55 Baseline 1.66 25.02 <0.001 5.22
Fixed target 1.58 19.38 <0.001 4.04
Anterior 1.54 17.00 <0.001 3.55
Lateral 1.53 16.82 <0.001 3.51
Vertical 0.80 Baseline 2.39 39.67 <0.001 8.27
Fixed target 2.34 41.36 <0.001 8.63
Anterior 2.35 34.44 <0.001 7.18
Lateral 2.36 51.81 <0.001 10.80

Trials included in the data analysis

Stepping performance was quantified for six different sets of trials: Baseline, Fixed target, targets that may shift anteriorly but did not, targets that shifted anteriorly, targets that may shift laterally but did not, and targets that shifted laterally. Spatiotemporal and UCM measures were quantified for four of these six sets of trials; the two sets of trials where the targets shifted were not used. Thus, spatiotemporal and UCM measures all had the same foot placement, but the expectation changed across the four conditions. Any differences in these measures are attributable only to the expectation of a maneuver.

2.4 Statistical analysis

To determine if participants adhered to the targeting task, we performed one-way ANOVA (6 levels: Baseline, Fixed target, no shift and shift trials from the Anterior-shift task, no shift and shift trials from the Lateral-shift task) on the RMSE. To determine the presence of anticipatory locomotor adjustments, we performed one-way ANOVAs (4 Tasks: Baseline, Fixed target, Anterior-shift target, Lateral-shift target) on traditional gait measures (gait speed, double support duration, lead and trail foot placement locations in AP and ML directions, step length and step width).

To determine the presence of synergies, we conducted separate two-tailed t-tests for comparing ΔVz to the appropriate discriminating value (ΔVz*; Table 2) for each performance variable and each task. To determine if the synergies were modulated with tasks, we performed the same ANOVA separately on the six synergy indices using the GLIMMIX procedure, with participants as the random effect. We performed all pair-wise post hoc comparisons with Tukey-Kramer adjustments when significant effect of Task was observed. Analyses were performed in SAS (Cary, NC, USA). The level of significance was set at 0.05.

3. Results

3.1 Targeting performance

RMSE was significantly different across tasks (Table 1, Fig 2). Post hoc comparisons revealed that all target tasks had smaller RMSE compared to Baseline, showing that participants performed the targeting task as instructed. The shift trials in Lateral-shift task had larger RMSE compared to the other target tasks.

Table 1. Spatial-temporal gait parameters and associated ANOVA results.

Target stepping accuracy F(5,110) p-value ηp2
RMSE 36.34 <0.001 0.62
Gait measures F (3,66) p-value η p 2
Gait speed 8.46 <0.001 0.28
Double support phase 2.06 0.114
Trail foot placement AP 3.56 0.019 0.14
ML 4.92 0.004 0.18
Lead foot placement AP 48.36 <0.001 0.19
ML 6.05 0.001 0.22
Step length 44.54 <0.001 0.67
Step width 2.52 0.066

Significant effects of Task are bolded.

Fig 2. Mean and standard error for foot placement RMSE for each task.

Fig 2

RMSE is computed separately for the no-shift and shift trials for the Anterior-shift and Lateral-shift tasks. Means with different letters are significantly different from one another.

3.2 Spatial-temporal gait measures

Significant effect of Task was observed for gait speed (Table 1, Fig 3A). Post hoc comparisons showed that gait speed significantly reduced for the Lateral-shift task compared to the other three tasks. No effect of Task was observed for the double support duration (Table 1, Fig 3B).

Fig 3.

Fig 3

(A) Mean and standard error for gait speed during double support phase, and (B) double support duration for the four tasks. Means with different letters are significantly different from one another.

Even though the target remained at the preferred foot location for the analyzed trials in all targeting tasks, a significant effect of Task was observed for AP and ML foot placements around the curb (Table 1). Post hoc comparisons revealed that the trail foot (on the curb) was placed closer to the curb edge and more lateral (left) for the Lateral-shift task compared to the Baseline task (Fig 4A). The lead foot (on the ground) was placed more forward for the Fixed target task compared to the Baseline task, and more forward for the Anterior- and Lateral- shift tasks compared to Fixed target task. The lead foot was also placed more lateral (right) for the Fixed target and Anterior-shift tasks compared to the Baseline task. As a result, step length was increased for the targeting tasks compared to Baseline, and it increased the most for the Anterior-shift task. Step width did not change across tasks (Table 1).

Fig 4.

Fig 4

(A) Mean and SE of foot placement locations relative to the curb edge (horizontal line) and the midpoint of the walkway for the four tasks. Error bars represent across-participant standard error. The dashed footprints are a zoomed-in view of the two foot placement locations with a common scale for provided on the right. (B, C) Mean and standard error for step length and step width. Means with different letters are significantly different from one another.

3.3 Synergies for the resultant force

The separate t-tests revealed that the synergy index for the resultant ML force was significantly greater than zero for all tasks. The Synergy index for the resultant AP force was significantly lower than zero for the Lateral-shift task, and it was not significantly different from zero for the other tasks. The synergy index for the resultant vertical force was significantly greater than zero for the Fixed target task, and it was not significantly different from zero for the other tasks (Table 2).

None of the ANOVAs on the synergy indices for the resultant force components revealed a significant effect of Task (Fig 5; Table 3).

Fig 5.

Fig 5

The mean and SE for synergy index (ΔVz) for stabilizing resultant force along the (A) AP, (B) ML and (C) vertical axis during the double support phase for the four tasks. The horizontal lines in each plot show the discriminating values for each synergy index.

Table 3. ANOVA results for the six synergy measures.

Synergy Axis F(3,66) p-value ηp2
Resultant force synergy AP 0.01 0.998
ML 0.15 0.926
Vertical 0.11 0.953
Resultant moment synergy AP 6.64 0.001 0.23
ML 1.80 0.156
Vertical 0.56 0.641

Significant effects of Task are bolded.

3.4 Synergies for the resultant moment

The separate t-tests revealed that the synergy index for the resultant moments along all three directions was significantly greater than the corresponding ΔVz* for all tasks (Table 2).

The ANOVA analyses revealed a significant effect of Task for the synergy stabilizing the resultant moment about the AP axis. Post hoc comparisons revealed that the synergy index decreased for all targeting tasks compared to Baseline. The ANOVAs on the synergy indices for resultant moments about the other two axes did not reveal an effect of Task (Fig 6; Table 3).

Fig 6.

Fig 6

Mean and standard error for the synergy index (ΔVz) for stabilizing resultant moment about the (A) AP, (B) ML, and (C) vertical axes during the double support phase. The horizontal lines in each plot show the discriminating values for each synergy index. ΔVz greater than the corresponding discriminating value indicates presence of synergy. Means with different letters are significantly different from one another.

4. Discussion

In the present study, we investigated the effects of future maneuverability demands on the stability ensured by the covariation between the GRFs (and free moments) under the two feet during the double-support phase of stepping down from a curb. We extended the current evaluation of kinetic locomotor synergies by examining the modulation of these synergies in a complex environment with maneuverability demands. Our hypothesis that kinetic synergies will be reduced when there is subsequent precision stepping demand was partially supported. Our hypotheses that (1) kinetic synergies will be reduced when expecting the target to shift versus a stationary target, and (2) synergies will be reduced preferentially depending on the direction of the expected maneuver were not supported.

The mechanics and control of step negotiation are well explored. However, the double support phase, with the feet planted on surfaces at different heights, is understudied, consistent with a relative neglect of this phase in the locomotion literature [21]. When both feet contact the ground, there are more kinetic degrees of freedom, which offers greater control authority over the gait cycle [14, 22]. Therefore, this phase of gait, although brief (lasting 20% of the gait cycle), may be an opportunity for modulating the body’s motion for achieving specific kinematic goals to navigate hazards. The main finding of this study contradicted our expectations: the kinetic synergies during double support of curb descent were minimally influenced by expected subsequent maneuvers. Nevertheless, our results corroborate our earlier findings, validating the novel synergy analysis that we introduced, and highlight the robustness of the ground reaction variable synergies engaged in stabilizing whole-body angular motions during the double support phase of curb descent [14]. We discuss the implications of this finding below.

4.1 Synergies stabilizing resultant forces

The variance structure in the GRFs is quite robust; the experimental manipulations had minimal effect on the synergy indices for the resultant forces. The index for the resultant ML force was positive, and it was not affected by the upcoming targeting requirements. This result is compatible with previous work indicating active control of gait in the ML direction [23, 24]. The vertical and AP forces, on the other hand, showed no synergistic co-variation for most cases (Table 2), corroborating our earlier findings for the Baseline task [14]. The behavior of these forces can be understood based on their roles in stabilizing linear and rotational motion.

During double support, each GRF pair contributes to a resultant force and resultant moments about two axes (Table 4). These contributions are evident from the statics analysis of the body posture in double support, i.e., from the spatial relationships between the GRFs lines of action and the CoM location [14]. For example, the vertical GRFs determine the resultant vertical force at the CoM, but also contribute to the resultant moment about the ML axis. Critically, attempting to stabilize the resultant vertical force will impair the stabilization of resultant ML moment, since the forces must covary negatively to stabilize the resultant vertical force, but they must covary positively to stabilize the resultant ML moment.

Table 4. Directions of covariations in the force pairs required to stabilize each performance variable.

Force pair Performance variable
FResultant-ML FResultant-AP FResultant-V MResultant-ML MResultant-AP MResultant-V
ML Negative Negative Positive
AP Negative Negative Positive
Vertical Negative Positive Positive

Therefore, the lack of synergy in resultant vertical and AP forces indicates that the angular motion in sagittal plane is stabilized at the cost of linear motion in the vertical and AP directions, likely to diminish the risk of a forward fall during curb descent. This interpretation is further supported by a consistent pattern in the data across three studies. The stabilization of the resultant AP and vertical forces diminishes across tasks as the person gains, maintains or loses height: both resultant forces are stabilized for curb ascent [14], only the resultant vertical force is stabilized during level walking [15], and neither resultant force is stabilized while stepping down. The risk and the consequences of a forward fall are higher when stepping down. Therefore, controlling forward pitching motion about the ML axis becomes more important, and the vertical and AP forces are engaged in stabilizing the resultant moment about the ML axis instead of their respective sums, reflecting a shift in control objectives in response to the task.

4.2 Synergies stabilizing resultant moments

The synergy indices for resultant moments about all three axes were higher than the corresponding discriminating values (with large effect sizes; Table 2), corroborating our earlier findings for the Baseline task [14], and extending them to more complex tasks. Subsequent stepping requirements reduced the synergy index for the resultant moment about the AP axis by 12%, i.e., the angular motion in the frontal plane became less stable. In contrast to our expectations, this change was insensitive to the nature of the upcoming task; the decline was consistent when the target was fixed, and when the target was likely to jump in the AP or the ML direction.

The decline in this synergy index could arise from increased variability in the trailing ankle push-off for the targeting tasks. Modulating ankle push-off is a known strategy for maintaining balance during locomotion [25, 26]; modulating ankle contributions may be an effective mechanism for modulating synergies during double support as well. Ankle push-off affects not only AP linear motion, but also angular momentum about the AP and ML axes [27]. In the target shift tasks, modulating ankle push-off may be used to increase step length or step width when needed. Furthermore, for the Fixed target task, the presence of the target constrains both step length and width, and this may also influence ankle push-off, and hence the synergy index. Foot target requirements–even when targets are located at preferred foot locations–are known to affect kinematic locomotor synergies [28]. The speculations regarding the association between ankle push-off and kinetic GRF synergies needs to be tested. Additionally, other mechanisms that could result in a lower synergy index, such as hip torque contributions, need to be explored.

The decline in the synergy index indicates active control of the stability of angular motion in the frontal plane, and the lack of Task effect on other synergy indices suggests that the same level of control was not employed for other resultant forces and moments. It is known that human gait requires more active control along the ML compared to the AP direction, and much work has focused on identifying the mechanisms for ML control [23, 25, 26]. However, this literature focuses mainly on ML linear motion, and most quantitative measures that assess ML balance do not consider the angular motion of the body [29]. Although whole-body angular momentum has been quantified during locomotion [27, 3032], our work shows that kinetic synergy analysis can improve our understanding of how we modulate body rotations in challenging environments. Furthermore, previous work on identifying the direction-specificity of locomotor control has focused on reactive foot placement modulations [3335]; our work extends these ideas to bilateral GRF coordination during double support, and critically, from reactive to proactive modulations for future maneuvers. However, we acknowledge that the insensitivity of the synergy adjustment to the nature of the upcoming targeting task requires further study.

4.3 Robust kinetic synergies during curb descent

Overall, the kinetic synergies during double support of curb descent were robust and minimally influenced by expected maneuvers, with only one of the six synergy indices showing a Task effect. There are two possible explanations for this finding. First, the consequences of failure at the two tasks–stepping down from the curb and subsequent targeting–are not equivalent. Loss of balance is more likely during curb descent, and a fall would have a severe cost. In comparison, failure to step into the visual target has minimal consequences. The main function of the synergies, then, is to maintain angular stability (reflected by the high moment synergies, and low force synergies), and this prerogative was not overcome by the upcoming targeting tasks that we used. This interpretation is compatible with the observation that angular momentum is tightly regulated while walking down stairs or inclines [31, 32]. In fact, our work here and earlier [14] extends this view to moments about all three coordinate axes.

Secondly, participants had additional opportunities to alter their movements and accomplish the targeting task. Slower walking speed for the Lateral-shift task reflected a strategy to wait to see if the target would shift. More forward ground foot placement (11% increase in the distance from the curb edge relative to Baseline) and longer step length (6% increase relative to Baseline) may reflect a strategy where the participant biased earlier foot placements in the direction of the potential target shift to facilitate targeting. Furthermore, although we did not quantify the swing phase, previous research has shown that young adults make reactive adjustments to track a moving foot placement target as late as mid-swing [36, 37]. Given these anticipatory and reactive opportunities to alter foot placement, participants may have preserved the kinetic synergies during double support to offset the higher risk of loss of balance during curb descent.

4.4 Limitations

The tasks we selected to simulate a challenging environment may not have sufficiently challenged young healthy participants. It is possible that if the task were more challenging–such as the target shifting in multiple directions rather than one direction only–we would have observed decline in the synergy indices. Thus, we should use caution when describing the synergies as robust until further environments are considered.

5. Conclusion

We have demonstrated that the kinetic synergies in the ground reaction forces and moments during the double support phase of curb descent are robust to and minimally influenced by expected foot targeting maneuvers that we used. With both feet contacting the ground during double support, there are additional kinetic degrees of freedom with which to affect the motion of the body. The main function of these kinetic variables is to maintain rotational stability (reflected by the high resultant moment synergies, and low resultant force synergies), and this prerogative was minimally influenced by the subsequent targeting tasks that we used, likely because the cost of losing balance while descending the curb would be higher than the cost of a misstep on a visual target. The synergy stabilizing rotational motion in the frontal plane declined when subsequent targeting was required. Further work is required to identify why the synergy adjustment was independent of the direction of target shift.

Supporting information

S1 Appendix

(DOCX)

Data Availability

All relevant data and code are archived with the Purdue University Research Repository. doi:10.4231/65GZ-K658.

Funding Statement

SA: Purdue University COVID-19 Research Disruption Fund. https://www.purdue.edu/research/oevprp/funding-and-grant-writing/funding/overview.php CC: Purdue University Department of Health and Kinesiology, Templin Graduate Student research award. https://www.purdue.edu/hhs/hk/graduate/scholarships/grants.html#:~:text=The%20Templin%20Graduate%20Student%20Research,research%20and%2For%20travel%20activities. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

References

  • 1.McFadyen BJ, Magnan GA, Boucher JP. Anticipatory locomotor adjustments for avoiding visible, fixed obstacles of varying proximity. Human Movement Science. 1993;12: 259–272. doi: 10.1016/0167-9457(93)90019-L [DOI] [Google Scholar]
  • 2.McFadyen BJ, Carnahan H. Anticipatory locomotor adjustments for accommodating versus avoiding level changes in humans. Exp Brain Res. 1997;114: 500–506. doi: 10.1007/pl00005659 [DOI] [PubMed] [Google Scholar]
  • 3.Rietdyk S. Anticipatory locomotor adjustments of the trail limb during surface accommodation. Gait & Posture. 2006;23: 268–272. doi: 10.1016/j.gaitpost.2005.03.006 [DOI] [PubMed] [Google Scholar]
  • 4.Acasio J, Wu M, Fey NP, Gordon KE. Stability-maneuverability trade-offs during lateral steps. Gait & Posture. 2017;52: 171–177. doi: 10.1016/j.gaitpost.2016.11.034 [DOI] [PubMed] [Google Scholar]
  • 5.Nestico J, Novak A, Perry SD, Mansfield A. Does increased gait variability improve stability when faced with an expected balance perturbation during treadmill walking? Gait & Posture. 2021;86: 94–100. doi: 10.1016/j.gaitpost.2021.03.014 [DOI] [PubMed] [Google Scholar]
  • 6.Wu M, Matsubara JH, Gordon KE. General and Specific Strategies Used to Facilitate Locomotor Maneuvers. PLOS ONE. 2015;10: e0132707. doi: 10.1371/journal.pone.0132707 [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 7.Reimann H, Fettrow T, Jeka JJ. Strategies for the Control of Balance During Locomotion. Kinesiology Review. 2018;7: 18–25. doi: 10.1123/kr.2017-0053 [DOI] [Google Scholar]
  • 8.Dickinson MH. How Animals Move: An Integrative View. Science. 2000;288: 100–106. doi: 10.1126/science.288.5463.100 [DOI] [PubMed] [Google Scholar]
  • 9.Webb PW, Weihs D. Stability versus Maneuvering: Challenges for Stability during Swimming by Fishes. Integrative and Comparative Biology. 2015;55: 753–764. doi: 10.1093/icb/icv053 [DOI] [PubMed] [Google Scholar]
  • 10.Tillman M, Ambike S. Cue-induced changes in the stability of finger force-production tasks revealed by the uncontrolled manifold analysis. Journal of Neurophysiology. 2018;119: 21–32. doi: 10.1152/jn.00519.2017 [DOI] [PubMed] [Google Scholar]
  • 11.Piscitelli D, Falaki A, Solnik S, Latash ML. Anticipatory postural adjustments and anticipatory synergy adjustments: preparing to a postural perturbation with predictable and unpredictable direction. Exp Brain Res. 2017;235: 713–730. doi: 10.1007/s00221-016-4835-x [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 12.Latash ML, Scholz JP, Schoner G. Motor Control Strategies Revealed in the Structure of Motor Variability: Exercise and Sport Sciences Reviews. 2002;30: 26–31. doi: 10.1097/00003677-200201000-00006 [DOI] [PubMed] [Google Scholar]
  • 13.Scholz JP, Schöner G. The uncontrolled manifold concept: identifying control variables for a functional task. Exp Brain Res. 1999;126: 289–306. doi: 10.1007/s002210050738 [DOI] [PubMed] [Google Scholar]
  • 14.Cui C, Kulkarni A, Rietdyk S, Barbieri FA, Ambike S. Synergies in the ground reaction forces and moments during double support in curb negotiation in young and older adults. Journal of Biomechanics. 2020;106: 109837. doi: 10.1016/j.jbiomech.2020.109837 [DOI] [PubMed] [Google Scholar]
  • 15.Toney ME, Chang Y-H. Humans robustly adhere to dynamic walking principles by harnessing motor abundance to control forces. Exp Brain Res. 2013;231: 433–443. doi: 10.1007/s00221-013-3708-9 [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 16.Elias LJ, Bryden MP, Bulman-Fleming MB. Footedness is a better predictor than is handedness of emotional lateralization. Neuropsychologia. 1998;36: 37–43. doi: 10.1016/s0028-3932(97)00107-3 [DOI] [PubMed] [Google Scholar]
  • 17.Hoogkamer W, Potocanac Z, Duysens J. Quick foot placement adjustments during gait: direction matters. Exp Brain Res. 2015;233: 3349–3357. doi: 10.1007/s00221-015-4401-y [DOI] [PubMed] [Google Scholar]
  • 18.da Silva Costa AA, dos Santos LO, Moraes R. Effect of a cognitive task on online adjustments when avoiding stepping on an obstacle and stepping on a target during walking in young adults. Exp Brain Res. 2018;236: 2387–2397. doi: 10.1007/s00221-018-5310-7 [DOI] [PubMed] [Google Scholar]
  • 19.Sparrow W, Begg R, Parker S. Aging effects on visual reaction time in a single task condition and when treadmill walking. Motor control. 2006. doi: 10.1123/MCJ.10.3.201 [DOI] [PubMed] [Google Scholar]
  • 20.Desailly E, Daniel Y, Sardain P, Lacouture P. Foot contact event detection using kinematic data in cerebral palsy children and normal adults gait. Gait & Posture. 2009;29: 76–80. doi: 10.1016/j.gaitpost.2008.06.009 [DOI] [PubMed] [Google Scholar]
  • 21.Vlutters M, Asseldonk EHF van, Kooij H van der. Center of mass velocity-based predictions in balance recovery following pelvis perturbations during human walking. Journal of Experimental Biology. 2016;219: 1514–1523. doi: 10.1242/jeb.129338 [DOI] [PubMed] [Google Scholar]
  • 22.Vielemeyer J, Müller R, Staufenberg N-S, Renjewski D, Abel R. Ground reaction forces intersect above the center of mass in single support, but not in double support of human walking. Journal of Biomechanics. 2021;120: 110387. doi: 10.1016/j.jbiomech.2021.110387 [DOI] [PubMed] [Google Scholar]
  • 23.Bauby CE, Kuo AD. Active control of lateral balance in human walking. Journal of Biomechanics. 2000;33: 1433–1440. doi: 10.1016/s0021-9290(00)00101-9 [DOI] [PubMed] [Google Scholar]
  • 24.O’Connor SM, Kuo AD. Direction-Dependent Control of Balance During Walking and Standing. J Neurophysiol. 2009;102: 1411–1419. doi: 10.1152/jn.00131.2009 [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 25.Hof AL, van Bockel RM, Schoppen T, Postema K. Control of lateral balance in walking: Experimental findings in normal subjects and above-knee amputees. Gait & Posture. 2007;25: 250–258. doi: 10.1016/j.gaitpost.2006.04.013 [DOI] [PubMed] [Google Scholar]
  • 26.Reimann H, Fettrow T, Thompson ED, Jeka JJ. Neural Control of Balance During Walking. Front Physiol. 2018;9. doi: 10.3389/fphys.2018.01271 [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 27.Neptune RR, McGowan CP. Muscle contributions to frontal plane angular momentum during walking. Journal of Biomechanics. 2016;49: 2975–2981. doi: 10.1016/j.jbiomech.2016.07.016 [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 28.Rosenblatt NJ, Hurt CP, Latash ML, Grabiner MD. An apparent contradiction: increasing variability to achieve greater precision? Exp Brain Res. 2014;232: 403–413. doi: 10.1007/s00221-013-3748-1 [DOI] [PubMed] [Google Scholar]
  • 29.Nott CR, Neptune RR, Kautz SA. Relationships between frontal-plane angular momentum and clinical balance measures during post-stroke hemiparetic walking. Gait & Posture. 2014;39: 129–134. doi: 10.1016/j.gaitpost.2013.06.008 [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 30.Herr H, Popovic M. Angular momentum in human walking. Journal of Experimental Biology. 2008;211: 467–481. doi: 10.1242/jeb.008573 [DOI] [PubMed] [Google Scholar]
  • 31.Silverman AK, Neptune RR, Sinitski EH, Wilken JM. Whole-body angular momentum during stair ascent and descent. Gait & Posture. 2014;39: 1109–1114. doi: 10.1016/j.gaitpost.2014.01.025 [DOI] [PubMed] [Google Scholar]
  • 32.Silverman AK, Wilken JM, Sinitski EH, Neptune RR. Whole-body angular momentum in incline and decline walking. Journal of Biomechanics. 2012;45: 965–971. doi: 10.1016/j.jbiomech.2012.01.012 [DOI] [PubMed] [Google Scholar]
  • 33.Collins SH, Kuo AD. Two independent contributions to step variability during over-ground human walking. PloS one. 2013;8: e73597. doi: 10.1371/journal.pone.0073597 [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 34.McAndrew Young PM, Wilken JM, Dingwell JB. Dynamic margins of stability during human walking in destabilizing environments. Journal of Biomechanics. 2012;45: 1053–1059. doi: 10.1016/j.jbiomech.2011.12.027 [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 35.Vlutters M, Van Asseldonk EHF, van der Kooij H. Foot Placement Modulation Diminishes for Perturbations Near Foot Contact. Front Bioeng Biotechnol. 2018;6. doi: 10.3389/fbioe.2018.00048 [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 36.Reynolds RF, Day BL. Visual guidance of the human foot during a step. The Journal of Physiology. 2005;569: 677–684. doi: 10.1113/jphysiol.2005.095869 [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 37.Young WR, Hollands MA. Evidence for age-related decline in visuomotor function and reactive stepping adjustments. Gait & Posture. 2012;36: 477–481. doi: 10.1016/j.gaitpost.2012.04.009 [DOI] [PubMed] [Google Scholar]

Decision Letter 0

Hugh Cowley

2 Sep 2022

PONE-D-22-11648Locomotion control during curb descent: Bilateral ground reaction variables covary consistently during the double support phase regardless of future foot placement constraintsPLOS ONE

Dear Dr. Ambike,

Thank you for submitting your manuscript to PLOS ONE. After careful consideration, we feel that it has merit but does not fully meet PLOS ONE’s publication criteria as it currently stands. Therefore, we invite you to submit a revised version of the manuscript that addresses the points raised during the review process.

Please see the reviewers' comments below. Please note that while Reviewer #2 was quite critical regarding the overall significance of this study compared to previous work in the field, and we respect that opinion, it is our assessment that this does not fail our criteria for publication; novelty (in the sense of significance or impact) is not a requirement for publication in PLOS ONE for original research that otherwise meets our requirements for quality of execution, reporting, and ethical standards. As such, while you may wish to clarify the specific contributions this work brings to the field, we require you only to address the other comments raised by each reviewer.

Please submit your revised manuscript by Oct 17 2022 11:59PM. If you will need more time than this to complete your revisions, please reply to this message or contact the journal office at plosone@plos.org. When you're ready to submit your revision, log on to https://www.editorialmanager.com/pone/ and select the 'Submissions Needing Revision' folder to locate your manuscript file.

Please include the following items when submitting your revised manuscript:

  • A rebuttal letter that responds to each point raised by the academic editor and reviewer(s). You should upload this letter as a separate file labeled 'Response to Reviewers'.

  • A marked-up copy of your manuscript that highlights changes made to the original version. You should upload this as a separate file labeled 'Revised Manuscript with Track Changes'.

  • An unmarked version of your revised paper without tracked changes. You should upload this as a separate file labeled 'Manuscript'.

If you would like to make changes to your financial disclosure, please include your updated statement in your cover letter. Guidelines for resubmitting your figure files are available below the reviewer comments at the end of this letter.

If applicable, we recommend that you deposit your laboratory protocols in protocols.io to enhance the reproducibility of your results. Protocols.io assigns your protocol its own identifier (DOI) so that it can be cited independently in the future. For instructions see: https://journals.plos.org/plosone/s/submission-guidelines#loc-laboratory-protocols. Additionally, PLOS ONE offers an option for publishing peer-reviewed Lab Protocol articles, which describe protocols hosted on protocols.io. Read more information on sharing protocols at https://plos.org/protocols?utm_medium=editorial-email&utm_source=authorletters&utm_campaign=protocols.

We look forward to receiving your revised manuscript.

Kind regards,

Hugh Cowley

Staff Editor

PLOS ONE

Journal Requirements:

When submitting your revision, we need you to address these additional requirements.

1. Please ensure that your manuscript meets PLOS ONE's style requirements, including those for file naming. The PLOS ONE style templates can be found at 

https://journals.plos.org/plosone/s/file?id=wjVg/PLOSOne_formatting_sample_main_body.pdf and 

https://journals.plos.org/plosone/s/file?id=ba62/PLOSOne_formatting_sample_title_authors_affiliations.pdf

2. We note that you have stated that you will provide repository information for your data at acceptance. Should your manuscript be accepted for publication, we will hold it until you provide the relevant accession numbers or DOIs necessary to access your data. If you wish to make changes to your Data Availability statement, please describe these changes in your cover letter and we will update your Data Availability statement to reflect the information you provide.

3. Please review your reference list to ensure that it is complete and correct. If you have cited papers that have been retracted, please include the rationale for doing so in the manuscript text, or remove these references and replace them with relevant current references. Any changes to the reference list should be mentioned in the rebuttal letter that accompanies your revised manuscript. If you need to cite a retracted article, indicate the article’s retracted status in the References list and also include a citation and full reference for the retraction notice.

[Note: HTML markup is below. Please do not edit.]

Reviewers' comments:

Reviewer's Responses to Questions

Comments to the Author

1. Is the manuscript technically sound, and do the data support the conclusions?

The manuscript must describe a technically sound piece of scientific research with data that supports the conclusions. Experiments must have been conducted rigorously, with appropriate controls, replication, and sample sizes. The conclusions must be drawn appropriately based on the data presented.

Reviewer #1: Yes

Reviewer #2: Yes

**********

2. Has the statistical analysis been performed appropriately and rigorously?

Reviewer #1: Yes

Reviewer #2: Yes

**********

3. Have the authors made all data underlying the findings in their manuscript fully available?

The PLOS Data policy requires authors to make all data underlying the findings described in their manuscript fully available without restriction, with rare exception (please refer to the Data Availability Statement in the manuscript PDF file). The data should be provided as part of the manuscript or its supporting information, or deposited to a public repository. For example, in addition to summary statistics, the data points behind means, medians and variance measures should be available. If there are restrictions on publicly sharing data—e.g. participant privacy or use of data from a third party—those must be specified.

Reviewer #1: Yes

Reviewer #2: Yes

**********

4. Is the manuscript presented in an intelligible fashion and written in standard English?

PLOS ONE does not copyedit accepted manuscripts, so the language in submitted articles must be clear, correct, and unambiguous. Any typographical or grammatical errors should be corrected at revision, so please note any specific errors here.

Reviewer #1: Yes

Reviewer #2: Yes

**********

5. Review Comments to the Author

Please use the space provided to explain your answers to the questions above. You may also include additional comments for the author, including concerns about dual publication, research ethics, or publication ethics. (Please upload your review as an attachment if it exceeds 20,000 characters)

Reviewer #1: Thank you for this well-written and interesting paper. Also, your description of UCM was well done-- terse but complete.

I have two comments for things that you need to address before publication: 1) where is the "foot center" line 170 p8 and throughout. Specifically, how do you define the foot center? I can imagine that the target center is the geometric center of the box. Obviously, this is a critical component of your results. 2) Were you participants shod or unshod? Please specify and if shod, discuss what kind of shoes and how that might have effected the results.

Reviewer #2: The paper is an extension of the “synergy index” line of work on which Dr. Ambike has developed and published extensively. The current focus is whether a targeted stepping requirement subsequent to a step up or down on a curb influence the synergy indices during the “curb step” compared to a condition in which no targeted stepping is required. The results showed minor effects, primarily in gait parameters, of the targeted stepping task. The authors conclude that performance on the targeting task had “low cost” relative to the task of maintaining upright stability while stepping up or down. Therefore, the targeting task had little effect on the synergy indices.

I agree with the authors’ interpretation of the results. There were small adjustments in gait parameters during the targeting conditions compared to baseline. But clearly, the targeting tasks were not demanding enough to threaten the stability of the upright body while stepping up or down.

From this perspective, the paper reproduces many of the results found in previous work, such as synergy indices related to free moments are related to upright stability moreso than ground reaction forces. The attempt to modify this essential finding was not successful. Such results contribute to the synergy index line of work in showing the nervous system prioritizes constraints while moving through the environment. Not a dramatic contribution.

Minor Comments

1. P 10 1st PP confusing sentences - please clarify. “All remaining measures included only the trials where the target did not shift. Thus, in the remaining measures, the foot placements during curb descent as well as the next step were the same in every task, and the expectation of a target shift differs across the tasks.”

2. P 10, line 221 – “…the same the ANOVA…” remove “the” before ANOVA.

3. Figure 2 caption – What are “…dissimilar letters…”?

**********

6. PLOS authors have the option to publish the peer review history of their article (what does this mean?). If published, this will include your full peer review and any attached files.

If you choose “no”, your identity will remain anonymous but your review may still be made public.

Do you want your identity to be public for this peer review? For information about this choice, including consent withdrawal, please see our Privacy Policy.

Reviewer #1: No

Reviewer #2: No

**********

[NOTE: If reviewer comments were submitted as an attachment file, they will be attached to this email and accessible via the submission site. Please log into your account, locate the manuscript record, and check for the action link "View Attachments". If this link does not appear, there are no attachment files.]

While revising your submission, please upload your figure files to the Preflight Analysis and Conversion Engine (PACE) digital diagnostic tool, https://pacev2.apexcovantage.com/. PACE helps ensure that figures meet PLOS requirements. To use PACE, you must first register as a user. Registration is free. Then, login and navigate to the UPLOAD tab, where you will find detailed instructions on how to use the tool. If you encounter any issues or have any questions when using PACE, please email PLOS at figures@plos.org. Please note that Supporting Information files do not need this step.

PLoS One. 2022 Oct 5;17(10):e0268090. doi: 10.1371/journal.pone.0268090.r002

Author response to Decision Letter 0


15 Sep 2022

We thank the reviewers for taking the time to review our manuscript and for their valuable feedback. Below is our point-by-point response to the editor’s comments and then the reviewers’ comments.

Editor’s comment:

Please note that while Reviewer #2 was quite critical regarding the overall significance of this study compared to previous work in the field, and we respect that opinion, it is our assessment that this does not fail our criteria for publication; novelty (in the sense of significance or impact) is not a requirement for publication in PLOS ONE for original research that otherwise meets our requirements for quality of execution, reporting, and ethical standards. As such, while you may wish to clarify the specific contributions this work brings to the field, we require you only to address the other comments raised by each reviewer.

RESPONSE: Thank you for the comment. We respect this comment by Reviewer 2. We have responded to it in this document below.

The main contributions of this work are: (1) the robust nature of the synergies, and (2) the replication of the synergy results from our previous work (Cui et al 2018). We think that the point about robust synergies is quite salient – in the title, abstract, discussion and conclusion sections. We adjusted some language to make the point about replication more prominent (Lines 328 - 333).

Comments by Reviewer 1:

Thank you for this well-written and interesting paper. Also, your description of UCM was well done-- terse but complete.

RESPONSE: We thank the reviewer for the encouragement.

I have two comments for things that you need to address before publication:

1) where is the "foot center" line 170 p8 and throughout. Specifically, how do you define the foot center? I can imagine that the target center is the geometric center of the box. Obviously, this is a critical component of your results.

RESPONSE: Foot center was defined as the average (geometric center) of the digitized toe and heel positions. The foot center was computed only when the foot was fully on the ground. For computing stepping error, the vertical coordinate of the foot center was ignored and only the AP and ML coordinates were utilized. We added text (Lines 173 and 181) to clarify this point.

2) Were you participants shod or unshod? Please specify and if shod, discuss what kind of shoes and how that might have affected the results.

RESPONSE: All participants walked with their own shoes. We instructed the participants to wear comfortable athletic shoes to the experiment. Walking in their own shoes adds to the ecological validity of the gait task. Compared to walking over level ground, there is greater impact on the foot while stepping down from a curb, and this may influence behavior. We decided to study shod walking because young adults likely spend more time walking with shoes than without, especially over uneven terrain.

We added text in the Methods section (Line 111) to clarify this point.

As for the effect of performing our tasks with and without shoes: There would likely only be minor differences across the shod/no-shod condition, if any. Specific values of the outcomes might change, but we expect the same significant effects in our statistical comparisons. There is no good reason why the synergy indices – the focus of this work – would change differently for expectation of shifting targets when wearing shoes versus barefoot.

Comments by Reviewer 2:

The paper is an extension of the “synergy index” line of work on which Dr. Ambike has developed and published extensively. The current focus is whether a targeted stepping requirement subsequent to a step up or down on a curb influence the synergy indices during the “curb step” compared to a condition in which no targeted stepping is required. The results showed minor effects, primarily in gait parameters, of the targeted stepping task. The authors conclude that performance on the targeting task had “low cost” relative to the task of maintaining upright stability while stepping up or down. Therefore, the targeting task had little effect on the synergy indices.

I agree with the authors’ interpretation of the results. There were small adjustments in gait parameters during the targeting conditions compared to baseline. But clearly, the targeting tasks were not demanding enough to threaten the stability of the upright body while stepping up or down.

From this perspective, the paper reproduces many of the results found in previous work, such as synergy indices related to free moments are related to upright stability more so than ground reaction forces. The attempt to modify this essential finding was not successful. Such results contribute to the synergy index line of work in showing the nervous system prioritizes constraints while moving through the environment. Not a dramatic contribution.

RESPONSE: We respectfully suggest that although not dramatic, the contributions of this work are significant. We show that, at least to the set of expected maneuvers that we used, these synergies are robust. This was not known before. Furthermore, our earlier paper was the first to introduce the UCM analysis of all ground reaction variables. Given the novelty of this analysis, it is quite important to demonstrate the replicability of the results, which we have achieved here.

Minor Comments

1. P 10 1st PP confusing sentences - please clarify. “All remaining measures included only the trials where the target did not shift. Thus, in the remaining measures, the foot placements during curb descent as well as the next step were the same in every task, and the expectation of a target shift differs across the tasks.”

RESPONSE: By analyzing only the trials where the target did not shift in the Anterior- and Lateral-shift conditions, we are certain that any difference in the outcomes is attributable to only to the expectation of a maneuver. We have expanded the section on how the data were analyzed to improve clarity (Lines 208-214).

2. P 10, line 221 – “…the same the ANOVA…” remove “the” before ANOVA.

RESPONSE: Thank you for pointing out the typo. We edited the text as suggested (Line 227).

3. Figure 2 caption – What are “…dissimilar letters…”?

RESPONSE: We replaced “dissimilar letters” with “…different letters…” in all the captions (Lines 564, Line 567, Line 574, and Line 585)

Decision Letter 1

Hugh Cowley

19 Sep 2022

Locomotion control during curb descent: Bilateral ground reaction variables covary consistently during the double support phase regardless of future foot placement constraints

PONE-D-22-11648R1

Dear Dr. Ambike,

We’re pleased to inform you that your manuscript has been judged scientifically suitable for publication and will be formally accepted for publication once it meets all outstanding technical requirements.

Within one week, you’ll receive an e-mail detailing the required amendments. When these have been addressed, you’ll receive a formal acceptance letter and your manuscript will be scheduled for publication.

An invoice for payment will follow shortly after the formal acceptance. To ensure an efficient process, please log into Editorial Manager at http://www.editorialmanager.com/pone/, click the 'Update My Information' link at the top of the page, and double check that your user information is up-to-date. If you have any billing related questions, please contact our Author Billing department directly at authorbilling@plos.org.

If your institution or institutions have a press office, please notify them about your upcoming paper to help maximize its impact. If they’ll be preparing press materials, please inform our press team as soon as possible -- no later than 48 hours after receiving the formal acceptance. Your manuscript will remain under strict press embargo until 2 pm Eastern Time on the date of publication. For more information, please contact onepress@plos.org.

Kind regards,

Hugh Cowley

Staff Editor

PLOS ONE

Additional Editor Comments (optional):

Reviewers' comments:

Acceptance letter

Hugh Cowley

26 Sep 2022

PONE-D-22-11648R1

Locomotion control during curb descent: Bilateral ground reaction variables covary consistently during the double support phase regardless of future foot placement constraints

Dear Dr. Ambike:

I'm pleased to inform you that your manuscript has been deemed suitable for publication in PLOS ONE. Congratulations! Your manuscript is now with our production department.

If your institution or institutions have a press office, please let them know about your upcoming paper now to help maximize its impact. If they'll be preparing press materials, please inform our press team within the next 48 hours. Your manuscript will remain under strict press embargo until 2 pm Eastern Time on the date of publication. For more information please contact onepress@plos.org.

If we can help with anything else, please email us at plosone@plos.org.

Thank you for submitting your work to PLOS ONE and supporting open access.

Kind regards,

PLOS ONE Editorial Office Staff

on behalf of

Mr Hugh Cowley

Staff Editor

PLOS ONE

Associated Data

    This section collects any data citations, data availability statements, or supplementary materials included in this article.

    Supplementary Materials

    S1 Appendix

    (DOCX)

    Data Availability Statement

    All relevant data and code are archived with the Purdue University Research Repository. doi:10.4231/65GZ-K658.


    Articles from PLoS ONE are provided here courtesy of PLOS

    RESOURCES