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1.
Patients with diabetic peripheral neuropathy are significantly more likely to fall while walking than subjects with intact sensation. While it has been suggested that these patients walk slower to improve locomotor stability, slower speeds are also associated with increased locomotor variability, and increased variability has traditionally been equated with loss of stability. If the latter were true, this would suggest that slowing down, as a locomotor control strategy, should be completely antithetical to the goal of maintaining stability. The present study resolves these seemingly paradoxical findings by using methods from nonlinear time series analysis to directly quantify the sensitivity of the locomotor system to local perturbations that are manifested as natural kinematic variability. Fourteen patients with severe peripheral neuropathy and 12 gender-, age-, height-, and weight-matched non-diabetic controls participated. Sagittal plane angles of the right hip, knee, and ankle joints and tri-axial accelerations of the trunk were measured during 10 min of continuous overground walking at self-selected speeds. Maximum finite-time Lyapunov exponents were computed for each time series to quantify the local dynamic stability of these movements. Neuropathic patients exhibited slower walking speeds and better local dynamic stability of upper body movements in the horizontal plane than did control subjects. The differences in local dynamic stability were significantly predicted by differences in walking speed, but not by differences in sensory status. These results support the hypothesis that reductions in walking speed are a compensatory strategy used by neuropathic patients to maintain dynamic stability of the upper body during level walking.  相似文献   

2.
Falls pose a tremendous risk to those over 65 and most falls occur during locomotion. Older adults commonly walk slower, which many believe helps improve walking stability. While increased gait variability predicts future fall risk, increased variability is also caused by walking slower. Thus, we need to better understand how differences in age and walking speed independently affect dynamic stability during walking. We investigated if older adults improved their dynamic stability by walking slower, and how leg strength and flexibility (passive range of motion (ROM)) affected this relationship. Eighteen active healthy older and 17 healthy younger adults walked on a treadmill for 5min each at each of 5 speeds (80-120% of preferred). Local divergence exponents and maximum Floquet multipliers (FM) were calculated to quantify each subject's inherent local dynamic stability. The older subjects walked with the same preferred walking speeds as the younger subjects (p=0.860). However, these older adults still exhibited greater local divergence exponents (p<0.0001) and higher maximum FM (p<0.007) than the younger adults at all walking speeds. These older adults remained more locally unstable (p<0.04) even after adjusting for declines in both strength and ROM. In both age groups, local divergence exponents decreased at slower speeds and increased at faster speeds (p<0.0001). Maximum FM showed similar changes with speed (p<0.02). Both younger and older adults exhibited decreased instability by walking slower, in spite of increased variability. These increases in dynamic instability might be more sensitive indicators of future fall risk than changes in gait variability.  相似文献   

3.
This study quantified the relationships between local dynamic stabiliht and variabilitr during continuous overground and treadmill walking. Stride-to-stride standard deviations were computed from temporal and kinematic data. Marimum finite-time Lyapunov exponents were estimated to quantify local dynamic stability. Local stability of gait kinematics was shown to be achieved over multiple consecutive strides. Traditional measures of variability poorly predicted local stability. Treadmill walking was associated with significant changes in both variability and local stability. Thus, motorized treadmills may produce misleading or erroneous results in situations where changes in neuromuscular control are likely to affect the variability and/or stability of locomotion.  相似文献   

4.
Individuals with peripheral neuropathy (PN) may compensate for decreased somatosensation by reducing walking speed. Predisposition to falls may therefore arise from an inability to adapt to challenging walking speeds. The purpose of this study was to examine the effects of PN on the magnitude of variability and local instability on walking at different speeds. Twelve individuals with PN and 12 controls completed a 6-min walk test to determine fast walking speed (FWS). Sagittal plane hip, knee, and ankle joint angles were then calculated during 3min of treadmill walking at 100%, 80%, and 60% FWS. The magnitudes of stride duration variability (SDvar), joint angle variability (JTvar), and both short- and long-term Lyapunov exponents (used to estimate local instability) were calculated. The PN group walked slower than the control group (p<.001). With groups combined, walking faster led to increased local instability and increased variability (p<.001). The PN group exhibited increased variability (SDvar, p=.02; JTvar, p=.01) over all speeds, and exaggerated local instability (p<.05) when walking at the fastest speed. PN leads to increased walking variability and local instability, particularly when walking at challenging speeds. These results are important to consider in future patient education and rehabilitation programs.  相似文献   

5.
Peripheral sensory feedback is believed to contribute significantly to maintaining walking stability. Patients with diabetic peripheral neuropathy have a greatly increased risk of falling. Previously, we demonstrated that slower walking speeds in neuropathic patients lead to improved local dynamic stability. However, all subjects exhibited significant local instability during walking, even though no subject fell or stumbled during testing. The present study was conducted to determine if and how significant changes in peripheral sensation and walking speed affect orbital stability during walking. Trunk and lower extremity kinematics were examined from two prior experiments that compared patients with significant neuropathy to healthy controls and walking at multiple different speeds in young healthy subjects. Maximum Floquet multipliers were computed for each time series to quantify the orbital stability of these movements. All subjects exhibited orbitally stable walking kinematics, even though these same kinematics were previously shown to be locally unstable. Differences in orbital stability between neuropathic and control subjects were small and, with the exception of knee joint movements (p=0.001), not statistically significant (0.380p0.946). Differences in knee orbital stability were not mediated by differences in walking speed. This was supported by our finding that although orbital stability improved slightly with slower walking speeds, the correlations between walking speed and orbital stability were generally weak (r(2)16.7%). Thus, neuropathic patients do not gain improved orbital stability as a result of slowing down and do not experience any loss of orbital stability because of their sensory deficits.  相似文献   

6.
Several efforts have been made to study gait stability using measures derived from nonlinear time-series analysis. The maximum finite time Lyapunov exponent (λmax) quantifies how a system responds to an infinitesimally small perturbation. Recent studies suggested that slow walking leads to lower λmax values, and thus is more stable than fast walking, but these studies suffer from methodological limitations. We studied the effects of walking speed on the amount of kinematic variability and stability in human walking. Trunk motions of 15 healthy volunteers were recorded in 3D during 2 min of treadmill walking at different speeds. From those time series, maximum Lyapunov exponents, indicating short-term and long-term divergence (λS-stride and λL-stride), and mean standard deviation (MeanSD) were calculated. λS-stride showed a linear decrease with increasing speed for forward–backward (AP) movements and quadratic effects (inverted U-shaped) for medio-lateral (ML) and up–down (VT) movements. λL-stride showed a quadratic effect (inverted U-shaped) of walking speed for AP movements, a linear decrease for ML movements, and a linear increase for VT movements. Moreover, positive correlations between λS and MeanSD were found for all directions, while λL-stride and MeanSD were correlated negatively in the AP direction. The different effects of walking speed on λS-stride and λL-stride for the different planes suggest that slow walking is not necessarily more stable than fast walking. The absence of a consistent pattern of correlations between λL-stride and MeanSD over the three directions suggests that variability and stability reflect, at least to a degree, different properties of the dynamics of walking.  相似文献   

7.
As humans walk or run, external (environmental) and internal (physiological) disturbances induce variability. How humans regulate this variability from stride-to-stride can be critical to maintaining balance. One cannot infer what is “controlled” based on analyses of variability alone. Assessing control requires quantifying how deviations are corrected across consecutive movements. Here, we assessed walking and running, each at two speeds. We hypothesized differences in speed would drive changes in variability, while adopting different gaits would drive changes in how people regulated stepping. Ten healthy adults walked/ran on a treadmill under four conditions: walk or run at comfortable speed, and walk or run at their predicted walk-to-run transition speed. Time series of relevant stride parameters were analyzed to quantify variability and stride-to-stride error-correction dynamics within a Goal-Equivalent Manifold (GEM) framework. In all conditions, participants’ stride-to-stride control respected a constant-speed GEM strategy. At each consecutively faster speed, variability tangent to the GEM increased (p ≤ 0.031), while variability perpendicular to the GEM decreased (p ≤ 0.044). There were no differences (p ≥ 0.999) between gaits at the transition speed. Differences in speed determined how stepping variability was structured, independent of gait, confirming our first hypothesis. For running versus walking, measures of GEM-relevant statistical persistence were significantly less (p ≤ 0.004), but showed minimal-to-no speed differences (0.069 ≤ p ≤ 0.718). When running, people corrected deviations both more quickly and more directly, each indicating tighter control. Thus, differences in gait determined how stride-to-stride fluctuations were regulated, independent of speed, confirming our second hypothesis.  相似文献   

8.
It is widely accepted that humans and animals minimize energetic cost while walking. While such principles predict average behavior, they do not explain the variability observed in walking. For robust performance, walking movements must adapt at each step, not just on average. Here, we propose an analytical framework that reconciles issues of optimality, redundancy, and stochasticity. For human treadmill walking, we defined a goal function to formulate a precise mathematical definition of one possible control strategy: maintain constant speed at each stride. We recorded stride times and stride lengths from healthy subjects walking at five speeds. The specified goal function yielded a decomposition of stride-to-stride variations into new gait variables explicitly related to achieving the hypothesized strategy. Subjects exhibited greatly decreased variability for goal-relevant gait fluctuations directly related to achieving this strategy, but far greater variability for goal-irrelevant fluctuations. More importantly, humans immediately corrected goal-relevant deviations at each successive stride, while allowing goal-irrelevant deviations to persist across multiple strides. To demonstrate that this was not the only strategy people could have used to successfully accomplish the task, we created three surrogate data sets. Each tested a specific alternative hypothesis that subjects used a different strategy that made no reference to the hypothesized goal function. Humans did not adopt any of these viable alternative strategies. Finally, we developed a sequence of stochastic control models of stride-to-stride variability for walking, based on the Minimum Intervention Principle. We demonstrate that healthy humans are not precisely “optimal,” but instead consistently slightly over-correct small deviations in walking speed at each stride. Our results reveal a new governing principle for regulating stride-to-stride fluctuations in human walking that acts independently of, but in parallel with, minimizing energetic cost. Thus, humans exploit task redundancies to achieve robust control while minimizing effort and allowing potentially beneficial motor variability.  相似文献   

9.
Lower limb amputees have decreased balance during daily ambulation compared to nonamputees. An optimally compliant torsion adapter, which enables transverse plane rotation at the socket–pylon junction may reduce limb asymmetries and improve comfort leading to increased confidence and stability during gait. The purpose of this study was to determine if the presence of a torsion adapter affects amputee sensitivity to local perturbations (local dynamic stability) during straight-line walking and during a turning task. Ten unilateral transtibial amputees were fit with a torsion and rigid adapter in random order and blinded to the condition. After a 3-week acclimation period, kinematic data were collected while subjects walked in a straight-line on a treadmill and around a 1-m radius circular path at constant speed. Maximum finite-time Lyapunov exponents (λ), an estimator of local dynamic stability, were calculated for the amputee’s sagittal plane hip, knee and ankle angles for each condition. The prosthetic limb λ was greater during a turn compared to straight-line walking, suggesting amputees are less stable while turning. There were no statistically significant differences found in λ between adapters during both walking conditions, suggesting the torsion adapter had no effect on amputee stability; however, high inter-subject variability due to the examined population and turning task may have masked a small decrease in prosthetic limb hip and knee stability for the torsion adapter during straight-line gait. Therefore, the torsion adapter’s added degree of freedom may have a small adverse effect on prosthetic limb stability during straight-line walking and no effect on turning.  相似文献   

10.
Currently there is no commonly accepted way to define, much less quantify, locomotor stability. In engineering, "orbital stability" is defined using Floquet multipliers that quantify how purely periodic systems respond to perturbations discretely from one cycle to the next. For aperiodic systems, "local stability" is defined by local divergence exponents that quantify how the system responds to very small perturbations continuously in real time. Triaxial trunk accelerations and lower extremity sagittal plane joint angles were recorded from ten young healthy subjects as they walked for 10 min over level ground and on a motorized treadmill at the same speed. Maximum Floquet multipliers (Max FM) were computed at each percent of the gait cycle (from 0% to 100%) for each time series to quantify the orbital stability of these movements. Analyses of variance comparing Max FM values between walking conditions and correlations between Max FM values and previously published local divergence exponent results were computed. All subjects exhibited orbitally stable walking kinematics (i.e., magnitudes of Max FM < 1.0), even though these same kinematics were previously found to be locally unstable. Variations in orbital stability across the gait cycle were generally small and exhibited no systematic patterns. Walking on the treadmill led to small, but statistically significant improvements in the orbital stability of mediolateral (p = 0.040) and vertical (p = 0.038) trunk accelerations and ankle joint kinematics (p = 0.002). However, these improvements were not exhibited by all subjects (p < or = 0.012 for subject x condition interaction effects). Correlations between Max FM values and previously published local divergence exponents were inconsistent and 11 of the 12 comparisons made were not statistically significant (r2 < or = 19.8%; p > or = 0.049). Thus, the variability inherent in human walking, which manifests itself as local instability, does not substantially adversely affect the orbital stability of walking. The results of this study will allow future efforts to gain a better understanding of where the boundaries lie between locally unstable movements that remain orbitally stable and those that lead to global instability (i.e., falling).  相似文献   

11.
Lower limb amputation substantially disrupts motor and proprioceptive function. People with lower limb amputation experience considerable impairments in walking ability, including increased fall risk. Understanding the biomechanical aspects of the gait of these patients is crucial in improving their gait function and their quality of life. In the present study, 9 persons with unilateral transtibial amputation and 13 able-bodied controls walked on a large treadmill in a Computer Assisted Rehabilitation Environment (CAREN). While walking, subjects were either not perturbed, or were perturbed either by continuous mediolateral platform movements or by continuous mediolateral movements of the visual scene. Means and standard deviations of both step lengths and step widths increased significantly during both perturbation conditions (all p<0.001) for both groups. Measures of variability, local and orbital dynamic stability of trunk movements likewise exhibited large and highly significant increases during both perturbation conditions (all p<0.001) for both groups. Patients with amputation exhibited greater step width variability (p=0.01) and greater trunk movement variability (p=0.04) during platform perturbations, but did not exhibit greater local or orbital instability than healthy controls for either perturbation conditions. Our findings suggest that, in the absence of other co-morbidities, patients with unilateral transtibial amputation appear to retain sufficient sensory and motor function to maintain overall upper body stability during walking, even when substantially challenged. Additionally, these patients did not appear to rely more heavily on visual feedback to maintain trunk stability during these walking tasks.  相似文献   

12.
Successful community and household ambulation require the ability to navigate corners and maneuver around obstacles, posing unique challenges compared to straight-line walking. The challenges associated with turning may contribute to an increased incidence of falling and the occurrence of fall-related injuries. A measure of stability applied to turning gait may be able to quantify a system's response to naturally occurring disturbances associated with turning and identify subjects at greater risk of falling. An index of stability has been used previously to assess the rate of kinematic separation (local dynamic stability) during straight-line gait. The purpose of this study was to determine if local dynamic stability during constant speed turning is reduced compared to straight-line treadmill walking. Maximum finite-time Lyapunov exponents (λ) were used to estimate the local stability of able-bodied subjects’ (n=19) sagittal plane hip, knee, and ankle trajectories for turning compared to straight-line walking at two different walking speeds. Turning λ was greater than straight λ for the hip, right knee, and ankle (p<0.05). Turning λ for the left knee angle was similar to straight λ. There were no differences in λ between left and right limbs for the hip and ankle and also no differences between the inside and outside limbs during turning for all joints. These findings indicate able-bodied subjects’ hip, right knee, and ankle kinematics are less locally stable while turning than walking in a straight line and may be used as a comparative tool for determining the efficacy of therapeutic interventions for mobility-impaired populations.  相似文献   

13.
Healthy humans display a preference for walking at a stride frequency dependent on the inertial properties of their legs. Walking at preferred stride frequency (PSF) is predicted to maximize local dynamic stability, whereby sensitivity to intrinsic perturbations arising from natural variability inherent in biological motion is minimized. Previous studies testing this prediction have employed different variability measures, but none have directly quantified local dynamic stability by computing maximum finite-time Lyapunov exponent (λMax), which quantifies the rate of divergence of nearby trajectories in state space. Here, ten healthy adults walked 45 m overground while sagittal motion of both knees was recorded via electrogoniometers. An auditory metronome prescribed 7 different frequencies relative to each individual's PSF (PSF; ±5, ±10, ±15 strides/min). Stride frequencies were performed under both freely adopted speed (FS) and controlled speed (CS: set at the speed of PSF trials) conditions. Local dynamic stability was maximal (λMax was minimal) at the PSF, becoming less stable for higher and lower stride frequencies. This occurred under both FS and CS conditions, although controlling speed further reduced local dynamic stability at non-preferred stride frequencies. In contrast, measures of variability revealed effects of stride frequency and speed conditions that were distinct from λMax. In particular, movement regularity computed by approximate entropy (ApEn) increased for slower walking speeds, appearing to depend on speed rather than stride frequency. The cadence freely adopted by humans has the benefit of maximizing local dynamic stability, which can be interpreted as humans tuning to their resonant frequency of walking.  相似文献   

14.
Repetitive trunk flexion elicits passive tissue creep, which has been hypothesized to compromise spine stability. The current investigation determined if increased spine flexion angle at the onset of flexion relaxation (FR) in the lumbar extensor musculature was associated with altered dynamic stability of spine kinematics. Twelve male participants performed 125 consecutive cycles of full forward trunk flexion. Spine kinematics and lumbar erector spinae (LES) electromyographic (EMG) activity were obtained throughout the repetitive trunk flexion trial. Dynamic stability was evaluated with maximum finite-time Lyapunov exponents over five sequential blocks of 25 cycles. Spine flexion angle at FR onset, and peak LES EMG activity were determined at baseline and every 25th cycle. Spine flexion angle at FR increased on average by 1.7° after baseline with significant increases of 1.7° and 2.4° at the 50th and 100th cycles. Maximum finite-time Lyapunov exponents demonstrated a transient, non-statistically significant, increase between cycles 26 and 50 followed by a recovery to baseline over the remainder of the repetitive trunk flexion cycles. Recovery of dynamic stability may be the consequence of increased active spine stiffness demonstrated by the non-significant increase in peak LES EMG that occurred as the repetitive trunk flexion progressed.  相似文献   

15.
Investigating inter-joint coordination at different walking speeds in young and elderly adults could provide insights to age-related changes in neuromuscular control of gait. We examined effects of walking speed and age on the pattern and variability of inter-joint coordination. Gait analyses of 10 young and 10 elderly adults were performed with different self-selected speeds, including a preferred, faster, and slower speed. Continuous relative phase (CRP), derived from phase planes of two adjacent joints, was used to assess the inter-joint coordination. CRP patterns were examined with cross-correlation measures and root-mean-square (RMS) differences when comparing ensemble mean curves of the faster or slower speed to preferred speed walking. Variability of coordination for each participant was assessed with the average value of all standard deviations calculated for each data point over a gait cycle from all CRP curves, namely the deviation phase (DP). For hip-knee CRP pattern, RMS differences were significantly greater between the slower and preferred walking speeds than between the faster and preferred walking speeds in young adults, but this was not found in elderly adults. Significant group differences in RMS differences and cross-correlation measures were detected in hip-knee CRP patterns between the slower and preferred walking speeds. No significant walking speed or age effects were detected for the knee-ankle CRP. Significant walking speed effects were also detected in hip-knee DP values. However, no significant group differences were detected for all three speeds. These findings suggested that young and elder adults compromise changes of walking speed with different neuromuscular control strategies.  相似文献   

16.
When walking at a given speed, humans often appear to prefer gait patterns that minimize metabolic rate, thereby maximizing metabolic economy. However, recent experiments have demonstrated that humans do not maximize economy when walking downhill. The purpose of this study was to investigate whether this non-metabolically optimal behavior is the result of a trade-off between metabolic economy and gait stability. We hypothesized that humans have the ability to modulate their gait strategy to increase either metabolic economy or stability, but that increase in one measure will be accompanied by decrease in the other. Subjects walked downhill using gait strategies ranging from risky to conservative, which were either prescribed by verbal instructions or induced by the threat of perturbations. We quantified spatiotemporal gait characteristics, metabolic rate and several indicators of stability previously associated with fall risk: stride period variability; step width variability; Lyapunov exponents; Floquet multipliers; and stride period fractal index. When subjects walked using conservative gait strategies, stride periods and lengths decreased, metabolic rate increased, and anteroposterior maximum Lyapunov exponents increased, which has previously been interpreted as an indicator of decreased stability. These results do not provide clear support for the proposed trade-off between economy and stability, particularly when stability is approximated using complex metrics. However, several gait pattern changes previously linked to increased fall risk were observed when our healthy subjects walked with a conservative strategy, suggesting that these changes may be a response to, rather than a cause of, increased fall risk.  相似文献   

17.
Humans do not generally walk at constant speed, except perhaps on a treadmill. Normal walking involves starting, stopping and changing speeds, in addition to roughly steady locomotion. Here, we measure the metabolic energy cost of walking when changing speed. Subjects (healthy adults) walked with oscillating speeds on a constant-speed treadmill, alternating between walking slower and faster than the treadmill belt, moving back and forth in the laboratory frame. The metabolic rate for oscillating-speed walking was significantly higher than that for constant-speed walking (6–20% cost increase for ±0.13–0.27 m s−1 speed fluctuations). The metabolic rate increase was correlated with two models: a model based on kinetic energy fluctuations and an inverted pendulum walking model, optimized for oscillating-speed constraints. The cost of changing speeds may have behavioural implications: we predicted that the energy-optimal walking speed is lower for shorter distances. We measured preferred human walking speeds for different walking distances and found people preferred lower walking speeds for shorter distances as predicted. Further, analysing published daily walking-bout distributions, we estimate that the cost of changing speeds is 4–8% of daily walking energy budget.  相似文献   

18.
To facilitate stable walking, humans must generate appropriate motor patterns and effective corrective responses to perturbations. Yet most EMG analyses do not address the continuous nature of muscle activation dynamics over multiple strides. We compared muscle activation dynamics in young and older adults by defining a multivariate state space for muscle activity. Eighteen healthy older and 17 younger adults walked on a treadmill for 2 trials of 5 min each at each of 5 controlled speeds (80–120% of preferred). EMG linear envelopes of v. lateralis, b. femoris, gastrocnemius, and t. anterior of the left leg were obtained. Interstride variability, local dynamic stability (divergence exponents), and orbital stability (maximum Floquet multipliers; FM) were calculated. Both age groups exhibited similar preferred walking speeds (p=0.86). Amplitudes and variability of individual EMG linear envelopes increased with speed (p<0.01) in all muscles but gastrocnemius. Older adults also exhibited greater variability in b. femoris and t. anterior (p<0.004). When comparing continuous multivariate EMG dynamics, older adults demonstrated greater local and orbital instability of their EMG patterns (p<0.01). We also compared how muscle activation dynamics were manifested in kinematics. Local divergence exponents were strongly correlated between kinematics and EMG, independent of age and walking speed, while variability and max FM were not. These changes in EMG dynamics may be related to increased neuromotor noise associated with aging and may indicate subtle deterioration of gait function that could lead to future functional declines.  相似文献   

19.
This study tested whether the lower economy of walking in healthy elderly subjects is due to greater gait instability. We compared the energy cost of walking and gait instability (assessed by stride to stride changes in the stride time) in octogenarians (G80, n = 10), 65-yr-olds (G65, n = 10), and young controls (G25, n = 10) walking on a treadmill at six different speeds. The energy cost of walking was higher for G80 than for G25 across the different walking speeds (P < 0.05). Stride time variability at preferred walking speed was significantly greater in G80 (2.31 +/- 0.68%) and G65 (1.93 +/- 0.39%) compared with G25 (1.40 +/- 0.30%; P < 0.05). There was no significant correlation between gait instability and energy cost of walking at preferred walking speed. These findings demonstrated greater energy expenditure in healthy elderly subjects while walking and increased gait instability. However, no relationship was noted between these two variables. The increase in energy cost is probably multifactorial, and our results suggest that gait instability is probably not the main contributing factor in this population. We thus concluded that other mechanisms, such as the energy expenditure associated with walking movements and related to mechanical work, or neuromuscular factors, are more likely involved in the higher cost of walking in elderly people.  相似文献   

20.
Research has shown that individuals have an optimal walking speed–a speed which minimizes energy expenditure for a given distance. Because the optimal walking speed varies with mass and lower limb length, it also varies with sex, with males in any given population tending to have faster optimal walking speeds. This potentially creates an energetic dilemma for mixed-sex walking groups. Here we examine speed choices made by individuals of varying stature, mass, and sex walking together. Individuals (N = 22) walked around a track alone, with a significant other (with and without holding hands), and with friends of the same and opposite sex while their speeds were recorded every 100 m. Our findings show that males walk at a significantly slower pace to match the females’ paces (p = 0.009), when the female is their romantic partner. The paces of friends of either same or mixed sex walking together did not significantly change (p>0.05). Thus significant pace adjustment appears to be limited to romantic partners. These findings have implications for both mobility and reproductive strategies of groups. Because the male carries the energetic burden by adjusting his pace (slowing down 7%), the female is spared the potentially increased caloric cost required to walk together. In energetically demanding environments, we will expect to find gender segregation in group composition, particularly when travelling longer distances.  相似文献   

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