Age-related changes of the stretch reflex excitability in human ankle muscles

The purpose of this study was to characterize the effects of aging on the stretch reflex in the ankle muscles, and in particular to compare the effects on the ankle dorsi-flexor (tibialis anterior: TA) and the plantar-flexor (soleus: SOL). Stretch reflex responses were elicited in the TA and SOL at rest and during weak voluntary contractions in 20 elderly and 23 young volunteers. The results indicated that, in the TA muscle, the elderly group had a remarkably larger long-latency reflex (LLR), whereas no aging effect was found in the short latency reflex (SLR). These results were very different from those in the SOL muscle, which showed significant aging effects in the SLR and medium latency reflex (MLR), but not in the LLR. Given the fact that the LLR of the TA stretch reflex includes the cortical pathway, it is probable that the effects of aging on the TA stretch reflex involve alterations not only at the spinal level but also at the cortical level. The present results indicate that the stretch reflexes of each of the ankle antagonistic muscles are affected differently by aging, which might have relevance to the neural properties of each muscle.     

Biomechanical properties of the human umbilical cord

The umbilical cord is a complex and fascinating structure that connects the fetus to the placenta and encases the umbilical vessels. The response of its tissues to mechanical loading due to fetal movements and uterine contractions is not well understood. The aim of this study is the evaluation of the mechanical properties of the main components of the human umbilical cord. Fresh umbilical cord specimens were collected from neonates born at term of the gestation and submitted to compliance tests. Furthermore, uniaxial tensile and stress-relaxation tests were performed on samples of umbilical vein and Wharton's jelly. Both materials exhibited nonlinear stress-strain response with increasing strain, increasing the elastic modulus (E(high) about 10-20 times E(low)) and significant viscoelastic behavior. In addition, anisotropy of the vein was observed. Although the circumferential properties of the vein (mean E(high) about 2.4 MPa) were similar to those after birth, the longitudinal stiffness of both materials was higher (mean E(high) over 10 MPa) and comparable to that of the ligaments. These findings suggest a mechanism of protection acting against excessive elongations of the cord, which could cause undue restriction of the umbilical vessel area and interference with the fetal blood circulation.     

Triceps surae short latency stretch reflexes contribute to ankle stiffness regulation during human running

During human running, short latency stretch reflexes (SLRs) are elicited in the triceps surae muscles, but the function of these responses is still a matter of controversy. As the SLR is primarily mediated by Ia afferent nerve fibres, various methods have been used to examine SLR function by selectively blocking the Ia pathway in seated, standing and walking paradigms, but stretch reflex function has not been examined in detail during running. The purpose of this study was to examine triceps surae SLR function at different running speeds using Achilles tendon vibration to modify SLR size. Ten healthy participants ran on an instrumented treadmill at speeds between 7 and 15 km/h under 2 Achilles tendon vibration conditions: no vibration and 90 Hz vibration. Surface EMG from the triceps surae and tibialis anterior muscles, and 3D lower limb kinematics and ground reaction forces were simultaneously collected. In response to vibration, the SLR was depressed in the triceps surae muscles at all speeds. This coincided with short-lasting yielding at the ankle joint at speeds between 7 and 12 km/h, suggesting that the SLR contributes to muscle stiffness regulation by minimising ankle yielding during the early contact phase of running. Furthermore, at the fastest speed of 15 km/h, the SLR was still depressed by vibration in all muscles but yielding was no longer evident. This finding suggests that the SLR has greater functional importance at slow to intermediate running speeds than at faster speeds.     

Ligament fibre recruitment at the human ankle joint complex in passive flexion

Knowledge of ligament fibre recruitment at the human ankle joint complex is a fundamental prerequisite for analysing mobility and stability. Previous experimental and modelling studies have shown that ankle motion must be guided by fibres within the calcaneofibular and tibiocalcaneal ligaments, which remain approximately isometric during passive flexion. The purpose of this study was to identify these fibres.

Three below-knee amputated specimens were analysed during passive flexion with combined radiostereometry for bone pose estimation and 3D digitisation for ligament attachment area identification. A procedure based on singular value decomposition enabled matching bone pose with digitised data and therefore reconstructing position in space of ligament attachment areas in each joint position. Eleven ordered fibres, connecting corresponding points on origin and insertion curves, were modelled for each of the following ligaments: posterior talofibular, calcaneofibular, anterior talofibular, posterior tibiotalar, tibiocalcaneal, and anterior tibiotalar.

The measured changes in length for the ligament fibres revealed patterns of tightening and slackening. The most anterior fibre of the calcaneofibular and the medio-anterior fibre of the tibiocalcaneal ligament exhibited the most isometric behaviour, as well as the most posterior fibre of the anterior talofibular ligament. Fibres within the calcaneofibular ligament remain parallel in the transverse plane, while those within the tibiocalcaneal ligament become almost parallel in joint neutral position. For both these ligaments, fibres maintain their relative inclination in the sagittal plane throughout the passive flexion range.

The observed significant change in both shape and orientation of the ankle ligaments suggest that this knowledge is fundamental for future mechanical analysis of their response to external forces.     

Biomechanical properties of the human cranium: aging aspects

Biomechanical properties of the human skull affect its dynamic tensility (pliability, compliance) by changes of intracranial volume and pressure (deltaV/deltaP). The goal of this study is to substantiate a possibility of noninvasive and dynamic evaluation of cranial compliance. The transcranial dopplerogram of middle cerebral artery and hemispheric bioimpedance were synchronously recorded, which represent information about pulsative changes of intracranial pressure and volume, respectively. The parameters were recorded at rest and during adequate hemo- and liquorodynamic tests in different age groups--20-30, 40-50, and 70-85 years. As compared with the young group, a decrease of the cranial compliance in the intermediate age group was revealed due to an observed increase if rigidity of skull bones and ligaments, which indicates a decrease of stability of the intracranial circulatory system. However, in the group of 70-85 years the compliance rose again due to an enlargement of intracranial liquor spaces and facilitation of liquor circulation inside the intracranial cavity; this can be suggested to be a compensatory mechanism for supporting the adequate brain circulatory-metabolic state.     

Biomechanical properties of human lumbar spine ligaments.

Biomechanical properties of the six major lumbar spine ligaments were determined from 38 fresh human cadaveric subjects for direct incorporation into mathematical and finite element models. Anterior and posterior longitudinal ligaments, joint capsules, ligamentum flavum, interspinous, and supraspinous ligaments were evaluated. Using the results from in situ isolation tests, individual force-deflection responses from 132 samples were transformed with a normalization procedure into mean force-deflection properties to describe the nonlinear characteristics. Ligament responses based on the mechanical characteristics as well as anatomical considerations, were grouped into T12-L2, L2-L4, and L4-S1 levels maintaining individuality and nonlinearity. A total of 18 data curves are presented. Geometrical measurements of original length and cross-sectional area for these six major ligaments were determined using cryomicrotomy techniques. Derived parameters including failure stress and strain were computed using the strength and geometry information. These properties for the lumbar spinal ligaments which are based on identical definitions used in mechanical testing and geometrical assay will permit more realistic and consistent inputs for analytical models.     

Biomechanical properties of human cranium: Age-relayed aspects

Biomechanical properties of the human skull affect its dynamic tensility (pliability or compliance) at changes of intracranial volume and pressure (ΔV/ΔP). The work substantiates a possibility of noninvasive and dynamic evaluation of cranial compliance by synchronous recording of transcranial dopplerogram of middle cerebral artery and cranial bioimpedance that provides information about pulsative changes of intracranial pressure and volume, respectively, with subsequent computer pattern and phasic analysis of these processes. The characteristic peculiarities of the cranial compliance at rest and during action of functional hemo- and liquorodynamic tests were traced in people of the middle (40–50 years) and elderly (70–85 years) age groups as compared with the young group (20–30 years). A relative decrease of this parameter has been revealed in the middle age group due to an increase of rigidity of skull bones and ligaments, which indicates a decrease of tolerance of the intracranial circulatory system. However, in the group of 70–85 years the compliance parameters rose due to an increase of intracranial liquor volume and activation of liquor circulation inside the craniospinal space, which is a compensatory mechanism for maintenance of the adequate brain circulatory-metabolic activity.     

Lamellar orientation in human cornea in relation to mechanical properties

We have applied wide-angle X-ray scattering to the human cornea in order to quantify the relative number of stromal collagen fibrils directed along the two preferred corneal lamellar directions: superior-inferior and nasal-temporal. The data suggest that, on average, the two directions are populated in equal proportion at the corneal centre. Here, approximately one-third of the fibrils throughout the stromal depth tend to lie within a 45 degrees sector of the superior-inferior meridian, and similarly for the nasal-temporal direction. However, in some eyes we have measured significant differences between the two preferential fibril populations, with some corneas exhibiting as much as 25% more collagen in one direction than the other. Based on these findings, a mechanical model of the normal cornea may be envisaged, whereby the fibril tension in the underlying "background" of isotropically arranged collagen helps to balance the intraocular pressure; while the extra preferentially aligned fibrils take up the additional tensile stress along the superior-inferior and nasal-temporal meridians exerted by the rectus muscles and the orbicularis. It is possible that, through a direct impact on the elastic modulus of the tissue, an imbalance of superior-inferior and nasal-temporal fibrils in some eyes might affect corneal shape.     

Energetics and passive dynamics of the ankle in downhill walking

This study investigated the energetics of the human ankle during the stance phase of downhill walking with the goal of modeling ankle behavior with a passive spring and damper mechanism. Kinematic and kinetic data were collected on eight male participants while walking down a ramp with inclination varying from 0° to 8°. The ankle joint moment in the sagittal plane was calculated using inverse dynamics. Mechanical energy injected or dissipated at the ankle joint was computed by integrating the power across the duration of the stance phase. The net mechanical energy of the ankle was approximately zero for level walking and monotonically decreased (i.e., became increasingly negative) during downhill walking as the slope decreased. The indication is that the behavior of the ankle is energetically passive during downhill walking, playing a key role in dissipating energy from one step to the next. A passive mechanical model consisting of a pin joint coupled with a revolute spring and damper was fit to the ankle torque and its parameters were estimated for each downhill slope using linear regression. The passive model demonstrated good agreement with actual ankle dynamics as indicated by low root-mean-square error values. These results indicate the stance phase behavior of the human ankle during downhill walking may be effectively duplicated by a passive mechanism with appropriately selected spring and damping characteristics.     

Biomechanical properties of human articular cartilage under compressive loads

The function of articular cartilage is to support and distribute loads and to provide lubrication in the diarthrodial joints. Cartilage function is described by proper mechanical and rheological properties, strain and depth-dependent, which are not completely assessed. Unconfined and confined compression are commonly used to evaluate the Young's modulus (E) and the aggregate modulus (H(A)), respectively. The Poisson's ratio (nu) can be calculated indirectly from the equilibrium compression data, or using the biphasic indentation technique; it has recently been optically evaluated by using video microscopy during unconfined compression. The transient response of articular cartilage during confined compression depends on its permeability k; a constant value of k can be easily identified by a simple analytical model of confined compression tests, whereas more complex models or direct measurements (permeation tests) are needed to study the permeability dependence on deformation. A poroelastic finite element model of articular cartilage was developed for this purpose. The elastic parameters (E,nu) of the model were evaluated performing unconfined compression creep tests on human articular cartilage disks, whereas k was identified from the confined test response. Our combined experimental and computational method can be used to identify the parameters that define the permeability dependence on deformation, as a function of depth from articular surface.     

Isoform diversity of giant proteins in relation to passive and active contractile properties of rabbit skeletal muscles

The active and passive contractile performance of skeletal muscle fibers largely depends on the myosin heavy chain (MHC) isoform and the stiffness of the titin spring, respectively. Open questions concern the relationship between titin-based stiffness and active contractile parameters, and titin's importance for total passive muscle stiffness. Here, a large set of adult rabbit muscles (n = 37) was studied for titin size diversity, passive mechanical properties, and possible correlations with the fiber/MHC composition. Titin isoform analyses showed sizes between approximately 3300 and 3700 kD; 31 muscles contained a single isoform, six muscles coexpressed two isoforms, including the psoas, where individual fibers expressed similar isoform ratios of 30:70 (3.4:3.3 MD). Gel electrophoresis and Western blotting of two other giant muscle proteins, nebulin and obscurin, demonstrated muscle type-dependent size differences of < or =70 kD. Single fiber and single myofibril mechanics performed on a subset of muscles showed inverse relationships between titin size and titin-borne tension. Force measurements on muscle strips suggested that titin-based stiffness is not correlated with total passive stiffness, which is largely determined also by extramyofibrillar structures, particularly collagen. Some muscles have low titin-based stiffness but high total passive stiffness, whereas the opposite is true for other muscles. Plots of titin size versus percentage of fiber type or MHC isoform (I-IIB-IIA-IID) determined by myofibrillar ATPase staining and gel electrophoresis revealed modest correlations with the type I fiber and MHC-I proportions. No relationships were found with the proportions of the different type II fiber/MHC-II subtypes. Titin-based stiffness decreased with the slow fiber/MHC percentage, whereas neither extramyofibrillar nor total passive stiffness depended on the fiber/MHC composition. In conclusion, a low correlation exists between the active and passive mechanical properties of skeletal muscle fibers. Slow muscles usually express long titin(s), predominantly fast muscles can express either short or long titin(s), giving rise to low titin-based stiffness in slow muscles and highly variable stiffness in fast muscles. Titin contributes substantially to total passive stiffness, but this contribution varies greatly among muscles.     

Prestress revealed by passive co-tension at the ankle joint

This study was designed to test the assumption that elastic tissues of the ankle are prestressed, by investigating the presence of simultaneous opposite passive elastic moments and thus, passive co-tension, at the ankle joint. A prestressed two-spring model used to generate qualitative predictions of the effects of stretching the posterior elastic structures of the ankle on the net passive moment of this joint was used. Twenty-seven healthy individuals were subjected to passive evaluation of the net elastic moment of the ankle in the sagittal plane, with the knee positioned at 90°, 60°, 30° and 0° of flexion, in order to change the length of the posterior biarticular elastic structures. The placement of the knee in the more extended positions caused changes in the net passive moment as predicted by the prestressed model. The ankle position in which the net passive moment was equal to zero was shifted to more plantar flexed positions (p<0.001) and there was a global increase in ankle stiffness since both passive dorsiflexion stiffness (p≤0.037) and passive plantar flexion stiffness (p≤0.029) increased. The normalized terminal plantar flexion stiffness also increased (p≤0.047), suggesting that biarticular posterior elastic structures are pre-strained and still under tension when the ankle is maximally plantar flexed and the knee is positioned at 60° of flexion. Resting positions were indicative of equilibrium between opposite passive elastic moments. The results revealed that there is passive co-tension at the ankle, demonstrating the existence of prestress in elastic structures of this joint.     

Length-tension properties of ankle muscles in chronic human spinal cord injury

Contracture, or loss of range of motion (ROM) of a joint, is a common clinical problem in individuals with spinal cord injury (SCI). In order to measure the possible contribution of changes in muscle length to the loss of ankle ROM, the active force vs. angle curves for the tibialis anterior (TA) and gastrocnemiussoleus (GS) were measured in 20 participants, 10 with SCI, and 10 gender and age matched, neurologically intact (NI) individuals. Electrical stimuli were applied to the TA and GS motor nerves at incremented angles of the entire ROM of the ankle and the resulting ankle and knee torques were measured using a multi-axis load cell. The muscle forces of the TA and GS were calculated from the torque measurements using estimates of their respective moment arms and the resulting forces were plotted against joint angle. The force–angle relation for the GS at the ankle (GSA) was significantly shifted into plantar flexion in SCI subjects, compared to NI controls (t-test, p<0.001). Similar results were obtained based upon the GS knee (GSK) force–angle measurements (p<0.05). Conversely, no significant shift in the force–angle relation was found for the TA (p=0.138). Differences in the passive ROM were consistent with the force–angle changes. The ROM in the dorsiflexion direction was significantly smaller in SCI subjects compared to NI controls (p<0.05) while the plantar flexion ROM was not significantly different (p=0.114). Based upon these results, we concluded that muscle shortening is an important component of contracture in SCI.     

Kinematic and passive resistive properties of human elbow complex

In recent years, owing to their versatility and reduced cost of operation, multisegmented mathematical models of the total human body have gained increased attention in gross biodynamic motion studies. This, in turn, has stimulated the need for a proper biomechanical data base for the major human articulating joints. The lack of such a database for the humero-elbow complex is the impetus for this study. The total angular range of motion permitted by the complex and the passive resistive properties beyond the full elbow extension were studied. Results obtained on ten normal male subjects were utilized to establish a statistical data base for the humero-elbow complex. Results are also expressed in functional expansion form suitable for incorporation into the existing multisegmented models.     

In vivo passive mechanical properties of the human gastrocnemius muscle belly

The purpose of the present study was to determine the in vivo passive mechanical properties, including the length below the slack length, of the gastrocnemius muscle (GAS) belly in humans. Transverse ultrasound images of the medial head of the GAS were taken in 11 subjects during passive knee extension from 80 degrees to 5 degrees with a constant ankle joint angle of 10 degrees (0 degrees is the neutral ankle position: positive values for dorsiflexion). The change in passive ankle joint moment (Mp), which is produced only by the GAS length change, was also measured during passive knee extension. The onset of Mp during passive knee extension was found to be 43+/-8 degrees (mean+/-SD) when the baseline of the Mp was set at the average Mp in the range of 55-60 degrees where the Mp was almost constant (SD<0.03 Nm). At this onset, the muscle fascicle length of the GAS (Lf) was 46+/-7 mm (slack length; Lfs). Lf at 80 degrees was 6+/-4 mm (13+/-6%) less than the Lfs, and Lf at 5 degrees was 12+/-5 mm (27+/-11%) greater than the Lfs. The passive force-resisting compression of the GAS did not produce a dorsiflexion moment in the joint angle range adopted. The passive ankle joint moment increased linearly with Lf (coefficient of determination (R2)=0.85-0.96), and the slopes of the relationships between Lf and Mp, and between the relative Lf to Lfs and Mp were 0.093+/-0.038 Nm/mm and 0.043+/-0.021 Nm/%Lfs. The findings of the present study can be implemented in musculoskeletal modeling, which would provide a more accurate evaluation of the passive mechanical properties of muscle during movement.     

Dynamic properties of stretch reflex

Dynamic aspects of stretch and unloading reflexes were investigated in the hindlimb extensor muscles of decerebrate cats. A complex transformation of non-linear effects inherent in the dynamics of the deafferented muscle was seen to occur under reflex control without hysteris (underlying non-linear static qualities of the muscle) being suppressed. Hysteresis of stretch reflex is responsible for uncertainty in muscle length equilibrium level — a factor in the origin of the movement — as well as the close connection between muscle stiffness and coordinates of the point of change in movement direction. The functional significance of non-linear aspects of the stretch reflex system is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 21, No. 5, pp. 589–597, September–October, 1989.     

Aging effects on posture-related modulation of stretch reflex excitability in the ankle muscles in humans

The purpose of this study was to examine the effects of aging on posture-related changes of the stretch reflex excitability in the ankle extensor, soleus (SOL), and flexor, tibialis anterior (TA) muscles. Fourteen neurologically normal elderly (mean 68 ± 6 years) and 12 young (mean 27 ± 3 years) subjects participated. Under two postural conditions, upright standing (STD) and sitting (SIT), stretch reflex electromyographic (EMG) responses in the SOL/TA muscle were elicited by imposing rapid ankle dorsi-/plantar-flexion. Under the SIT condition, subjects were asked to keep the SOL background EMG level, which is identical to that under the STD condition. In the SOL muscle, both groups showed significant enhancement of the short-latency stretch reflex (SLR) response when the posture changed from SIT to STD. In the TA muscle, the young group showed significant enhancement of the middle- (MLR) and long-latency stretch reflex (LLR) when the posture changed from SIT to STD; no such modulation was observed in the elderly group. Since the TA stretch reflex responses under the STD condition were comparable in the young and elderly groups, the lack of posture-related modulation of the TA muscle in the elderly group might be explained by augmented stretch reflex excitability under the SIT condition. The present results suggest that the (1) SOL SLR responses are modulated both in the young and elderly subjects when the posture is changed from SIT to STD, (2) TA MLR and LLR responses are not modulated in the elderly subjects when the posture is changed from SIT to STD, while each response is same between the young and elderly in STD, and (3) the effect of aging on the posture-related stretch reflex differs in the SOL and TA muscles.