Joint torques in a freely walking insect reveal distinct functions of leg joints in propulsion and posture control.
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ABSTRACT:
SUBMITTER: Dallmann CJ
PROVIDER: S-EPMC4795010 | biostudies-literature | 2016 Jan
REPOSITORIES: biostudies-literature
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Project description:A joint moment also causes motion at other joints of the body. This joint coupling-perspective allows more insight into two age-related phenomena during gait. First, whether increased hip kinetic output compensates for decreased ankle kinetic output during positive joint work. Second, whether preserved joint kinetic patterns during negative joint work in older age have any functional implication. Therefore, we examined how age and surface inclination affect joint moment strategies to accelerate and/or decelerate individual leg joints during walking. Healthy young (age: 22.5 ± 4.1 years, n = 18) and older (age: 76.0 ± 5.7 years, n = 22) adults walked at 1.4 m/s on a split-belt instrumented treadmill at three grades (0%, 10%, -10%). Lower-extremity moment-induced angular accelerations were calculated for the hip (0% and 10%) and knee (0% and -10%) joints. During level and uphill walking, both age groups showed comparable ankle moment-induced ipsilateral (p = 0.774) and contralateral (p = 0.047) hip accelerations, although older adults generated lower ankle moments in late stance. However, ankle moment-induced contralateral hip accelerations were smaller (p = 0.001) in an older adult subgroup (n = 13) who showed larger hip extension moments in early stance than young adults. During level and downhill walking, leg joint moment-induced knee accelerations were unaffected by age (all p > 0.05). These findings suggest that during level and uphill walking increased hip flexor mechanical output in older adults does not arise from reduced ankle moments, contrary to increased hip extensor mechanical output. Additionally, results during level and downhill walking imply that preserved eccentric knee extensor function is important in maintaining knee stabilization in older age.
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Project description:Multi-legged locomotion requires appropriate coordination of all legs with coincident ground contact. Whereas behaviourally derived coordination rules can adequately describe many aspects of inter-leg coordination, the neural mechanisms underlying these rules are still not entirely clear. The fact that inter-leg coordination is strongly affected by cut thoracic connectives in tethered walking insects, shows that neural information exchange among legs is important. As yet, recent studies have shown that load transfer among legs can contribute to inter-leg coordination through mechanical coupling alone, i.e., without neural information exchange among legs. Since naturalistic load transfer among legs works only in freely walking animals but not in tethered animals, we tested the hypothesis that connective lesions have less strong effects if mechanical coupling through load transfer among legs is possible. To do so, we recorded protraction/retraction angles of all legs in unrestrained walking stick insects that either had one thoracic connective cut or had undergone a corresponding sham operation. In lesioned animals, either a pro-to-mesothorax or a meso-to-metathorax connective was cut. Overall, our results on temporal coordination were similar to published reports on tethered walking animals, in that the phase relationship of the legs immediately adjacent to the lesion was much less precise, although the effect on mean phase was relatively weak or absent. Lesioned animals could walk at the same speed as the control group, though with a significant sideward bias toward the intact side. Detailed comparison of lesion effects in free-walking and supported animals reveal that the strongest differences concern the spatial coordination among legs. In free walking, lesioned animals, touch-down and lift-off positions shifted significantly in almost all legs, including legs of the intact body side. We conclude that insects with disrupted neural information transfer through one connective adjust to this disruption differently if they experience naturalistic load distribution. While mechanical load transfer cannot compensate for lesion-induced effects on temporal inter-leg coordination, several compensatory changes in spatial coordination occur only if animals carry their own weight.
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Project description:Walking is a complex rhythmic locomotor behavior generated by sequential and periodical contraction of muscles essential for coordinated control of movements of legs and leg joints. Studies of walking in vertebrates and invertebrates have revealed that premotor neural circuitry generates a basic rhythmic pattern that is sculpted by sensory feedback and ultimately controls the amplitude and phase of the motor output to leg muscles. However, the identity and functional roles of the premotor interneurons that directly control leg motoneuron activity are poorly understood. Here we take advantage of the powerful genetic methodology available in Drosophila to investigate the role of premotor inhibition in walking by genetically suppressing inhibitory input to leg motoneurons. For this, we have developed an algorithm for automated analysis of leg motion to characterize the walking parameters of wild-type flies from high-speed video recordings. Further, we use genetic reagents for targeted RNAi knockdown of inhibitory neurotransmitter receptors in leg motoneurons together with quantitative analysis of resulting changes in leg movement parameters in freely walking Drosophila Our findings indicate that targeted down-regulation of the GABAA receptor Rdl (Resistance to Dieldrin) in leg motoneurons results in a dramatic reduction of walking speed and step length without the loss of general leg coordination during locomotion. Genetically restricting the knockdown to the adult stage and subsets of motoneurons yields qualitatively identical results. Taken together, these findings identify GABAergic premotor inhibition of motoneurons as an important determinant of correctly coordinated leg movements and speed of walking in freely behaving Drosophila.
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Project description:BACKGROUND:Older adults are more susceptible to balance perturbations during walking than young adults. However, we lack an individual joint-level understanding of how aging affects the neuromechanical strategies used to accommodate balance perturbations. RESEARCH QUESTION:We investigated gait phase-dependence in and aging effects on leg joint kinematic variability during walking with balance perturbations. We hypothesized that leg joint variability would: 1) vary across the gait cycle and 2) increase with balance perturbations. We also hypothesized that perturbation effects on leg joint kinematic variability would be larger and more pervasive in older versus young adults. METHODS:We collected leg joint kinematics in young and older adults walking with and without mediolateral optical flow perturbations of different amplitudes. RESULTS:We first found that leg joint variability during walking is gait phase-dependent, with step-to-step adjustments occurring predominantly during push-off and early swing. Second, young adults accommodated perturbations almost exclusively by increasing coronal plane hip joint variability, likely to adjust step width. Third, perturbations elicited larger and more pervasive increases in all joint kinematic outcome measures in older adults. Finally, we also provide insight into which joints contribute more to foot placement variability in walking, adding that variability in sagittal plane knee and coronal plane hip joint angles contributed most to that in step length and step width, respectively. SIGNIFICANCE:Taken together, our findings may be highly relevant to identifying specific joint-level therapeutic targets to mitigate balance impairment in our aging population.
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Project description:IntroductionAdvanced age brings a distal-to-proximal redistribution of positive joint work during walking that is relevant to walking performance and economy. It is unclear whether negative joint work is similarly redistributed in old age. Negative work can affect positive work through elastic energy return in gait. We determined the effects of age, walking speed, and grade on positive and negative joint work in young and older adults.MethodsBilateral ground reaction force and marker data were collected from healthy young (age = 22.5 yr, n = 18) and older (age = 76.0 yr, n = 22) adults walking on a split-belt instrumented treadmill at 1.1, 1.4, and 1.7 m·s at each of three grades (0%, 10%, and -10%). Subjects also performed maximal voluntary eccentric, isometric, and concentric contractions for the knee extensors (120°·s, 90°·s, and 0°·s) and plantarflexors (90°·s, 30°·s, and 0°·s).ResultsCompared with young adults, older adults exhibited a distal-to-proximal redistribution of positive leg joint work during level (P < 0.001) and uphill (P < 0.001) walking, with larger differences at faster walking speeds. However, the distribution of negative joint work was unaffected by age during level (P = 0.150) and downhill (P = 0.350) walking. Finally, the age-related loss of maximal voluntary knee extensor (P < 0.001) and plantarflexor (P = 0.001) strength was smaller during an eccentric contraction versus concentric contraction for the knee extensors (P < 0.001) but not for the plantarflexors (P = 0.320).ConclusionThe distal-to-proximal redistribution of positive joint work during level and uphill walking is absent for negative joint work during level and downhill walking. Exercise prescription should focus on improving ankle muscle function while preserving knee muscle function in older adults trying to maintain their independence.
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Project description:We sought to evaluate the effects of the application of torque pulses to the hip and knee joint via a robotic exoskeleton in the context of training propulsion during walking. Based on our previous study, we formulated a set of conditions of torque pulses applied to the hip and knee joint associated with changes in push-off posture, a component of propulsion. In this work, we quantified the effects of hip/knee torque pulses on metrics of propulsion, specifically hip extension (HE) and normalized propulsive impulse (NPI), in two experiments. In the first experiment, we exposed 16 participants to sixteen conditions of torque pulses during single strides to observe the immediate effects of pulse application. In the second experiment, we exposed 16 participants to a subset of those conditions for 200 strides to quantify short-term adaptation effects. During pulse application, NPI aligned with the expected modulation of push-off posture, while HE was modulated in the opposite direction. The timing of the applied pulses, early or late stance, was crucial, as the effects were often in the opposite direction when changing timing condition. Extension torque applied at late stance increased HE in both experiments - range of change in HE: (2.9 ± 0.4 deg, 7.7 ± 1.0 deg), . The same conditions resulted in a negative change in NPI only in the single pulse experiment - change in NPI for knee torque: -3.0 ± 0.4 ms, - and no significant change for hip torque. Also, knee extension and flexion torque during early and late stance, respectively, increased NPI during single pulse application - range of change in NPI: (3.8, 4.6) ± 0.8 ms, . During repeated pulse application, NPI increased for late stance flexion torque - range of change in NPI: (4.5 ± 0.7 ms, 4.8 ± 0.8 ms), , but not late stance extension torque. After exposure, we observed positive after-effects in HE in three conditions - range of change in HE: (2.0 ± 0.3 deg, 3.7 ± 0.7 deg) - and significant positive after-effects in NPI for early stance flexion torques - change in NPI: (2.7 ± 0.6 ms, ). These results indicate that positive propulsive after-effects can be achieved through repeated exposure to torque pulses.
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Project description:Individual joint deviations are often identified in the analysis of cerebral palsy (CP) gait. However, knowledge is limited as to how these deviations affect the control of the locomotor system as a whole when striving to meet the demands of walking. The current study aimed to bridge the gap by describing the control of the locomotor system in children with diplegic CP in terms of their leg stiffness, both skeletal and muscular components, and associated joint stiffness during gait. Twelve children with spastic diplegia CP and 12 healthy controls walked at a self-selected pace in a gait laboratory while their kinematic and forceplate data were measured and analyzed during loading response, mid-stance, terminal stance and pre-swing. For calculating the leg stiffness, each of the lower limbs was modeled as a non-linear spring, connecting the hip joint center and the corresponding center of pressure, with varying stiffness that was calculated as the slope (gradient) of the axial force vs. the deformation curve. The leg stiffness was further decomposed into skeletal and muscular components considering the alignment of the lower limb. The ankle, knee and hip of the limb were modeled as revolute joints with torsional springs whose stiffness was calculated as the slope of the moment vs. the angle curve of the joint. Independent t-tests were performed for between-group comparisons of all the variables. The CP group significantly decreased the leg stiffness but increased the joint stiffness during stance phase, except during terminal stance where the leg stiffness was increased. They appeared to rely more on muscular contributions to achieve the required leg stiffness, increasing the muscular demands in maintaining the body posture against collapse. Leg stiffness plays a critical role in modulating the kinematics and kinetics of the locomotor system during gait in the diplegic CP.
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Project description:Understanding how humans adapt gait mechanics for a wide variety of locomotor tasks is important for inspiring the design of robotic, prosthetic and wearable assistive devices. We aimed to elicit the mechanical adjustments made to leg joint functions that are required to generate accelerative walking and running, using metrics with direct relevance to device design. Twelve healthy male participants completed constant speed (CS) walking and running and emulated acceleration (ACC) trials on an instrumented treadmill. External force and motion capture data were combined in an inverse dynamics analysis. Ankle, knee and hip joint mechanics were described and compared using angles, moments, powers and normalized functional indexes that described each joint as relatively more: spring, motor, damper or strut-like. To accelerate using a walking gait, the ankle joint was switched from predominantly spring-like to motor-like, while the hip joint was maintained as a motor, with an increase in hip motor-like function. Accelerating while running involved no change in the primary function of any leg joint, but involved high levels of spring and motor-like function at the hip and ankle joints. Mechanical adjustments for ACC walking were achieved primarily via altered limb positioning, but ACC running needed greater joint moments.
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Project description: Not available
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Project description:We investigated the effect of simultaneous changes in body-weight-support level and walking speed on mean peak internal joint moments at the ankle, knee and hip. We hypothesized that observed changes in these joint moments would be approximately linear with both body-weight-support and walking speed and would be similar across joints. Kinematic and kinetic data were collected from 8 unimpaired adult subjects walking on an instrumented treadmill while wearing a dynamically controlled overhead support harness. Subjects walked with four levels of body-weight-support (0%, 20%, 40%, and 60% of bodyweight) at three walking speeds (0.4, 0.6, and 0.8 statures/s, ranging on average from 0.7 to 1.4m/s). Data were used to calculate mean peak joint moments across subjects for each condition. In general, subjects' mean peak joint moments decreased linearly with decreasing walking speed and with increasing body-weight-support, except the knee extension moment, which showed a quadratic relationship with walking speed and no significant change with body-weight-support. All joint moments, with the exception of knee extension, showed a significant interaction effect between walking speed and body-weight-support, indicating that the sensitivity of these joint moments to changes in these variables was interdependent. In most cases, the ankle and hip extension moments showed the largest sensitivity to walking speed. The ankle moment was observed to have the greatest sensitivity to body-weight-support. This finding, that altering walking speed and body-weight-support level results in non-uniform changes in peak moments across joints, suggests that further research is warranted to establish the set of combined speed and support conditions that produce motor patterns supportive of normal gait retraining.
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