Biology subjects

Ludvig, D.

Publications and source records attributed to Ludvig, D..

10 recordsLinked to original sources

bioRxiv2026

Triceps surae and Achilles tendon contributions to ankle stiffness depend on movement state

Ankle stiffness is decreased during movement compared to posture; however, the etiology of this decrease remains unknown. Determining what gives rise to this decrease is critical for understanding how humans successfully interact with their physical world and how that ability is compromised by functional impairments. While the triceps surae and Achilles tendon primarily dictate ankle stiffness, the relative contributions across posture and movement remain unknown. Therefore, our study sought to quantify the relative contributions of the muscle and tendon to ankle stiffness and how those contributions differ between posture and movement. We used our technique, which combines B-mode ultrasound imaging with joint-level perturbations, to quantify ankle, muscle, and tendon stiffness simultaneously. Since ankle, muscle, and tendon stiffness all scale with torque, participants matched torque between posture and movement tasks. During posture, the Achilles tendon is the dominant contributor to ankle stiffness. However, during movement, the triceps surae and Achilles tendon contribute more equally to ankle stiffness, which can be attributed to a significant decrease in muscle stiffness during movement. Here, we provide the first empirical data on how state-dependent properties of the triceps surae and Achilles tendon contribute to ankle stiffness in conditions relevant to locomotion.

bioengineering↗

bioRxiv2024

Effects of Uni- and Bidirectional Interaction During Dyadic Ankle and Wrist Tracking

Haptic human-robot-human interaction allows users to feel and respond to one anothers forces while interfacing with separate robotic devices, providing customizable infrastructure for studying physical interaction during motor tasks (i.e., physical rehabilitation). For both upper- and lower-limb tasks, previous work has shown that virtual interactions with a partner can improve motor performance and enhance individual learning. However, whether the mechanism of these improvements generalizes across different human systems is an open question. In this work, we investigate the effects of haptic interaction between healthy individuals during a trajectory tracking task involving single-joint movements at the wrist and ankle. We compare tracking performance and muscle activation during haptic conditions where pairs of participants were uni- and bidirectionally connected, in order to investigate the contribution of real-time responses from a partner during the interaction. Findings indicate similar improvements in tracking performance during the bidirectional interaction for both the wrist and ankle, despite significant differences in how individuals modulated co-contraction. For each joint, bidirectional and unidirectional interaction resulted in similar improvements for the worse partner in the dyad. For the better partner, bidirectional interaction outperformed unidirectional interaction, likely due to changes in movement planning that were not observed in the unidirectional condition. While these results suggest that unidirectional interaction is sufficient for error correction of less skilled individuals during simple motor tasks, they also highlight the mutual benefits of bidirectional interaction which are consistent across the upper and lower limbs.

bioengineering↗

bioRxiv2024

Along- and cross-muscle fiber shear moduli in skeletal muscle

The material properties of muscle play a central role in how muscle resists joint motion, transmits forces internally, and repairs itself. While many studies have evaluated muscles tensile material properties, few have investigated muscles shear properties. The objective of this study was to quantify the shear moduli of skeletal muscle both along (along-muscle fiber) and perpendicular (cross-muscle fiber) to the direction of muscle fibers. We collected data from the extensor digitorum longus, tibialis anterior, and soleus muscles harvested from both hindlimbs of 12 rats. These muscles were chosen to further evaluate the consistency of shear moduli across muscles with different architectures. We applied strains and measured stress in three configurations: parallel, perpendicular, and across the muscle fibers to characterize the along- and cross-muscle fiber tensile and shear material parameters. We found no significant difference between the shear modulus measured parallel to the fibers (along-muscle fiber) and the shear modulus in the plane perpendicular to the fibers (cross-muscle fiber). Although the shear moduli were not significantly different, there was a greater difference with increasing strain, suggesting that there is greater anisotropy at larger strains. We also found no significant difference in moduli between the muscles with differing muscle architecture. These results characterize the shear behavior of skeletal muscle and are relevant to understanding the role of shear in force transmission and injury.

bioengineering↗

bioRxiv2023

At matched loads, aging does not alter ankle, muscle, or tendon stiffness

Older adults have difficulty maintaining balance when faced with postural disturbances, a task that is influenced by the stiffness of the triceps surae and Achilles tendon. Age-related changes in Achilles tendon stiffness have been reported at matched levels of effort, but measures typically have not been made at matched loads, which is important due to age-dependent changes in strength. Moreover, age-dependent changes in muscle stiffness have yet to be tested. Here, we investigate how age alters muscle and tendon stiffness and their influence on ankle stiffness. We hypothesized that age-related changes in muscle and tendon contribute to reduced ankle stiffness in older adults and evaluated this hypothesis when either load or effort were matched. We used B-mode ultrasound with joint-level perturbations to quantify ankle, muscle, and tendon stiffness across a range of loads and efforts in seventeen healthy younger and older adults. At matched loads, there was no significant difference in ankle, muscle, or tendon stiffness between groups (all p>0.13). However, at matched effort, older adults exhibited a significant decrease in ankle (27%; p=0.008), muscle (37%; p=0.02), and tendon stiffness (22%; p=0.03) at 30% of maximum effort. This is consistent with our finding that older adults were 36% weaker than younger adults in plantarflexion (p=0.004). Together these results indicate that, at the loads tested in this study, there are no age-dependent changes in the mechanical properties of muscle or tendon, only differences in strength that result in altered ankle, muscle, and tendon stiffness at matched levels of effort. NO_SCPLOWEWC_SCPLOWO_SCPCAP C_SCPCAPO_SCPLOWANDC_SCPLOW NO_SCPLOWOTEWORTHYC_SCPLOWWe provide the first simultaneous estimates of ankle, muscle, and tendon stiffness in younger and older adults. In contrast to earlier conclusions, we found that muscle and tendon mechanical properties are unaffected by age when compared at matched loads. However, due to age-related decreases in strength, mechanical properties do differ at matched efforts. As such, it is important to assess the relevance of the comparisons being made relative to the functional tasks under consideration.

bioengineering↗

bioRxiv2022

The nervous system leverages the non-linear properties of the Achilles tendon to regulate ankle impedance during postural control

Regulating ankle impedance is essential for controlled interactions with the environment and rejecting unexpected disturbances. Ankle impedance in the sagittal plane depends strongly on the triceps surae and Achilles tendon, but their relative contributions remain unknown. It is commonly assumed that ankle impedance is controlled by changing muscle activation and, thereby, muscle impedance, but this ignores the fact that tendon impedance also changes with loading caused by increases in muscle activation. Thus, we sought to determine the relative contributions from the triceps surae and Achilles tendon during conditions relevant to postural control. We used a novel technique that combines B-mode ultrasound imaging with joint-level perturbations to quantify ankle, muscle, and tendon impedance simultaneously across activation levels from 0 - 30% of maximum voluntary contraction. We found that muscle and tendon stiffness, the static component of impedance, increased with voluntary plantarflexion contractions, but that muscle stiffness exceeded tendon stiffness at very low loads (21 {+/-} 7 N). Above these loads, corresponding to 1.3% of maximal strength for an average participant in our study, ankle stiffness was determined predominately by Achilles tendon stiffness. Hence, the nervous system leverages the non-linear properties of the Achilles tendon to increase ankle stiffness during postural conditions.

physiology↗

bioRxiv2021

Simultaneous quantification of ankle, muscle, and tendon impedance in humans

ObjectiveRegulating the impedance of our joints is essential for the effective control of posture and movement. The impedance of a joint is governed mainly by the mechanical properties of the muscle-tendon units spanning it. Many studies have quantified the net impedance of joints but not the specific contributions from the muscles and tendons. The inability to quantify both muscle and tendon impedance limits the ability to determine the causes underlying altered movement control associated with aging, neuromuscular injury, and other conditions that have different effects on muscle and tendon properties. Therefore, we developed a technique to quantify joint, muscle, and tendon impedance simultaneously and evaluated this technique at the human ankle. MethodsWe used a single degree of freedom actuator to deliver pseudorandom rotations to the ankle while measuring the corresponding torques. We simultaneously measured the displacement of the medial gastrocnemius muscle-tendon junction with B-mode ultrasound. From these experimental measurements, we were able to estimate ankle, muscle, and tendon impedance using non-parametric system identification. ResultsWe validated our estimates by comparing them to previously reported muscle and tendon stiffness, the position-dependent component of impedance, to demonstrate that our technique generates reliable estimates of these properties. ConclusionOur approach can be used to clarify the respective contributions from the muscle and tendon to the net mechanics of a joint. SignificanceThis is a critical step forward in the ultimate goal of understanding how muscles and tendons govern ankle impedance during posture and movement.

bioengineering↗

bioRxiv2021

Short latency stretch reflexes depend on the balance of activity in agonist and antagonist muscles during ballistic elbow movements

The spinal stretch reflex is a fundamental building block of motor function, with a sensitivity that varies continuously during movement and when changing between movement and posture. Many have investigated task-dependent reflex sensitivity, but few have provided simple, quantitative analyses of the relationship between the volitional control and stretch reflex sensitivity throughout tasks that require coordinated activity of several muscles. Here we develop such an analysis and use it to test the hypothesis that modulation of reflex sensitivity during movement can be explained by the balance of activity within agonist and antagonist muscles better than by activity only in the muscle homonymous with the reflex. Subjects completed hundreds of flexion and extension movements as small, pseudo-random perturbations of elbow angle were applied to obtain estimates of stretch reflex amplitude throughout the movement. A subset of subjects performed a postural control task at muscle activities matched to those during movement. We found that reflex modulation during movement can be described by background activity in antagonist muscles about the elbow much better than by activity only in the muscle homonymous to the reflex (p<0.001). Agonist muscle activity enhanced reflex sensitivity whereas antagonist activity suppressed it. Surprisingly, the magnitude of these effects was similar, suggesting a balance of control between agonists and antagonists very different from the dominance of sensitivity to homonymous activity during posture. This balance is due to a large decrease in sensitivity to homonymous muscle activity during movement rather than substantial changes in the influence of antagonistic muscle activity. New and NoteworthyThis study examined the sensitivity of the stretch reflexes elicited in elbow muscles to the background activity in these same muscles during movement and postural tasks. We found a heightened reciprocal control of reflex sensitivity during movement that was not present during maintenance of posture. These results help explain previous discrepancies in reflex sensitivity measured during movement and posture and provide a simple model for assessing their contributions to muscle activity in both tasks.

neuroscience↗

bioRxiv2021

Frontal plane ankle stiffness increases with axial load independent of muscle activity

Ankle sprains are the most common musculoskeletal injury, typically resulting from excessive inversion of the ankle. One way to prevent excessive inversion and maintain ankle stability is to generate a stiffness that is sufficient to resist externally imposed rotations. Frontal-plane ankle stiffness increases as participants place more weight on their ankle, but whether this effect is due to muscle activation or axial loading of the ankle is unknown. Identifying whether and to what extent axial loading affects ankle stiffness is important in understanding what role the passive mechanics of the ankle joint play in maintaining its stability. The objective of this study was to determine the effect of passive axial load on frontal-plane ankle stiffness. We had subjects seated in a chair as an axial load was applied to the ankle ranging from 10% to 50% body weight. Small rotational perturbations were applied to the ankle in the frontal plane to estimate stiffness. We found a significant, linear, 3-fold increase in ankle stiffness with axial load from the range of 0% bodyweight to 50% bodyweight. This increase could not be due to muscle activity as we observed no significant axial-load-dependent change in any of the recorded muscle activations. These results demonstrate that axial loading is a significant contributor to maintaining frontal-plane ankle stability, and that disruptions to the mechanism mediating this sensitivity of stiffness to axial loading may result in pathological cases of ankle instability.

bioengineering↗

bioRxiv2021

Multidirectional measures of shear modulus in skeletal muscle

The material properties of muscle play a central role in how muscle resists joint motion, transmits forces internally, and repairs itself. While many studies have evaluated muscles tensile material properties, few have investigated muscles shear properties. None of which have taken into account muscles anisotropic structure or investigated how different muscle architecture affect muscles shear properties. The objective of this study was to quantify the shear moduli of skeletal muscle in three orientations relevant to the function of whole muscle. We collected data from the extensor digitorum longus, tibialis anterior, and soleus harvested from both hindlimbs of 12 rats. These muscles were chosen to further evaluate the consistency of shear moduli across muscles with different architectures. We calculated the shear modulus in three orientations: parallel, perpendicular, and across with respect to muscle fiber alignment; while the muscle was subjected to increasing shear strain. For all muscles and orientations, the shear modulus increased with increasing strain. The shear modulus measured perpendicular to fibers was greater than in any other orientation. Despite architectural differences between muscles, we did not find a significant effect of muscle type on shear modulus. Our results show that in rat, muscles shear moduli vary with respect to fiber orientation and are not influenced by architectural differences in muscles.

physiology↗

bioRxiv2021

Leveraging joint mechanics simplifies the neural control of movement

Behaviors we perform every day, such as manipulating an object or walking, require precise control of interaction forces between our bodies and the environment. These forces are generated by active muscle contractions, specified by the nervous system, and by joint mechanics, determined by the intrinsic properties of the musculoskeletal system. Depending on behavioral goals, joint mechanics might simplify or complicate control of movement by the nervous system. However, whether humans can exploit joint mechanics to simplify neural control remains unclear. Here we evaluated if leveraging joint mechanics can simplify neural control by comparing performance in three tasks that required subjects to generate specified torques about the ankle during imposed sinusoidal movements; only one task required torques that could be generated by leveraging the intrinsic mechanics of the joint. We developed a novel approach that used continuous estimates of impedance, a quantitative description of joint mechanics, and measures of muscle activity to determine the mechanical and neural contributions to the net ankle torque generated in each motor task. We found that the torque resulting from changes in neural control was reduced when ankle impedance was consistent with the task being performed, resulting in a task that required less muscular effort. Subjects perceived this task to be easier than those that were not consistent with the impedance of the ankle and were able to perform it with the highest level of consistency. These results demonstrate that leveraging the mechanical properties of a joint can simplify task completion and improve performance. KEY POINTSO_LIInteracting with our environment requires production of interaction forces, which are generated by muscle contractions, specified by the nervous system, and by joint mechanics, determined by the intrinsic properties of the musculoskeletal system. C_LIO_LIWe assessed whether leveraging joint mechanics can simplify neural control by having subjects complete 3 tasks, only one of which could be accomplished by leveraging joint mechanics. C_LIO_LIWe found that subjects reduced their muscular effort, perceived the task to be easier and completed the task more consistently when the mechanics of the ankle were consistent with the task. C_LIO_LIThese results highlight the importance of considering limb mechanics when interpreting measures of neural control related to movement and may benefit the design of mechanical interfaces that optimize human performance. C_LI

neuroscience↗