Medullary expiratory activity: influence of intercostal tendon organs and muscle spindle endings

Studies were conducted to determine the effects of intercostal muscle spindle endings (MSEs) and tendon organs (TOs) on medullary expiratory activity in decerebrate cats. Impeded intercostal muscle contractions, elicited by electrical stimulation of the peripheral cut end of the T6 ventral root, were used to stimulate intercostal TOs without MSEs. Impeded contractions of the intercostal muscles augmented expiratory laryngeal motoneuron activity, and either had no effect on or reduced the activity of bulbospinal expiratory neurons. Vibration was used to stimulate intercostal MSEs. Intercostal MSEs had no effect on medullary expiratory neuron activity. It is concluded that both external and internal intercostal TOs have an excitatory effect on expiratory laryngeal motoneuron activity and an inhibitory effect on a subpopulation of expiratory neurons driving intercostal and/or abdominal muscles, and intercostal MSEs have no direct influence on medullary expiratory activity.     

Medullary inspiratory activity: influence of intercostal tendon organs and muscle spindle endings

Studies were conducted to determine the effects of intercostal muscle spindle endings (MSEs) and tendon organs (TOs) on medullary inspiratory activity in decerebrate and allobarbital-anesthetized cats. Impeded muscle contractions, elicited by electrical stimulation of the peripheral cut end of the T6 ventral root, were used to stimulate external and internal intercostal TOs without MSEs. Impeded contractions of either the external or internal intercostal muscles reduced phrenic and medullary inspiratory neuronal activities. Vibration was used to selectively stimulate external or internal intercostal MSEs (90 and 40 micron amplitude, respectively). Selective stimulation of either external or internal intercostal MSEs did not change phrenic or medullary inspiratory neuronal activities. It is concluded that both external and internal intercostal TOs have a generalized inhibitory effect on medullary inspiratory activity and intercostal MSEs have no effect on medullary inspiratory activity.     

Elastic properties of human Achilles tendon are correlated to muscle strength.

The purpose of this study was to investigate whether the mechanical properties of the Achilles tendon were correlated to muscle strength in the triceps surae in humans. Twenty-four men and twelve women exerted maximal voluntary isometric plantar flexion (MVIP) torque. The elongation (DeltaX) and strain of the Achilles tendon (epsilon), the proximal part of which is the composite of the gastrocnemius tendon and the soleus aponeurosis, at MVIP were determined from the displacement of the distal myotendinous junction of the medial gastrocnemius using ultrasonography. The Achilles tendon force at MVIP (F) was calculated from the MVIP torque and the Achilles tendon moment arm. There were no significant differences in either the F-DeltaX or F-epsilon relationships between men and women. DeltaX and epsilon were 9.8 +/- 2.6 mm and 5.3 +/- 1.6%, respectively, and were positively correlated to F (r = 0.39, P < 0.05; r = 0.39, P < 0.05), which meant that subjects with greater muscle strength could store more elastic energy in the tendon. The regression y-intercepts for the F-DeltaX (P < 0.01) and F-epsilon (P < 0.05) relationship were significantly positive. These results might indicate that the Achilles tendon was stiffer in subjects with greater muscle strength, which may play a role in reducing the probability of tendon strain injuries. It was suggested that the Achilles tendon of subjects with greater muscle strength did not impair the potential for storing elastic energy in tendons and may be able to deliver the greater force supplied from a stronger muscle more efficiently. Furthermore, the difference in the Achilles tendon mechanical properties between men and women seemed to be correlated to the difference in muscle strength rather than gender.     

The regulation of aggrecanase ADAMTS-4 expression in human Achilles tendon and tendon-derived cells

Several members of the ADAMTS (A Disintegrin And Metalloproteinase with ThromboSpondin motifs) family have been identified as aggrecanases, whose substrates include versican, the principal large proteoglycan in the tendon extracellular matrix. We have characterized the expression of ADAMTS-4 in human Achilles tendon and tendon-derived cells. ADAMTS-4 mRNA levels were higher in ruptured tendon compared with normal tendon or chronic painful tendinopathy. In tissue extracts probed by Western blotting, mature ADAMTS-4 (68 kDa) was detected only in ruptured tendons, while processed ADAMTS-4 (53 kDa) was detected also in chronic painful tendinopathy and in normal tendon. In cultured Achilles tendon cells, transforming growth factor-β (TGF-β) stimulated ADAMTS-4 mRNA expression (typically 20-fold after 24 h), while interleukin-1 induced a smaller, shorter-term stimulation which synergised markedly with that induced by TGF-β. Increased levels of immunoreactive proteins consistent with mature and processed forms of ADAMTS-4 were detected in TGF-β-stimulated cells. ADAMTS-4 mRNA was expressed at higher levels by tendon cells in collagen gels than in monolayer cultures. In contrast, the expression of ADAMTS-1 and -5 mRNA was lower in collagen gels compared with monolayers, and these mRNA showed smaller or opposite responses to growth factors and cytokines compared with that of ADAMTS-4 mRNA. We conclude that both ADAMTS-4 mRNA and ADAMTS-4 protein processing may be differentially regulated in normal and damaged tendons and that both the matrix environment and growth factors such as TGF-β are potentially important factors controlling ADAMTS aggrecanase activities in tendon pathology.     

Structuro-functional organization of the fiber network in the human Achilles tendon

Owing to a complex morphological investigation of the human Achilles tendon, it was possible to distinguish four levels of the structural-functional organization of its fibrous elements and to reveal some regularities of their structure that recur at all the levels. Thus, collagenous molecules, microfibrillae, fibrillae and fibers have a wavy-spiral conformation. This spatial form is stabilized by a complex or a system of transversal connections corresponding to the given level of the organization. In order to maintain integrity (the structural-functional unity) of each level, certain substances of polysaccharide nature take part. Along the course of the long tendinous axis, a re-distribution (branching) of the fibrillar elements is observed at all the levels of the structural-functional organization.     

Effect of habitual running on human Achilles tendon load-deformation properties and cross-sectional area.

Whether the cross-sectional area (CSA) and mechanical properties of the human Achilles tendon change in response to habitual exercise remains largely unexplored. The present study evaluated the CSA and contraction-induced displacement of the aponeurosis-tendon complex of the triceps surae in 11 untrained subjects before (tests 1 and 2) and after (test 3) approximately 9 mo of regular running ( approximately 78 training sessions). Displacement of the tendon-aponeurosis complex obtained by ultrasonography; electromyography of the gastrocnemius, soleus, and dorsiflexor muscles; and joint angular rotation were recorded during graded isometric plantarflexion ramps. Tendon CSA and moment arm were measured by using MRI, and tendon force was calculated from joint moments and tendon moment arm. A treadmill test was used to determine submaximal oxygen consumption (Vo2) at a given speed and maximal Vo2. The total running duration was approximately 43 h, distributed over 34 wk. Maximal Vo2 increased 8.6% (P < 0.01), and submaximal Vo2 decreased 6.2% (P < 0.05). Tendon-aponeurosis displacement during maximal voluntary contraction was unchanged (tests 1-3, 5.2 +/- 0.6, 5.2 +/- 0.5, and 5.3 +/- 0.4 mm, respectively) and yielded a structural stiffness of 365 +/- 50, 358 +/- 40, and 384 +/- 52 N/mm for tests 1-3, respectively (P > 0.05). Tendon CSA also remained unchanged (tests 1-3, 34.2 +/- 2.2, 33.9 +/- 2.2, and 33.8 +/- 2.1 mm2, respectively). In conclusion, a total training stimulus of approximately 9 mo of running in previously untrained subjects was adequate to induce significant cardiovascular improvements, although it did not result in any changes in the mechanical properties of the triceps surea tendon-aponeurosis complex or in the dimensions of Achilles tendon.     

Biological incorporation of human acellular dermal matrix used in Achilles tendon repair

Human acellular dermal matrices (ADMs) are used successfully in a variety of procedures, including sports medicine related, wound repair, and breast reconstructions, but the mechanism of repair is still not fully understood. An opportunity to explore this mechanism presented itself when a patient experienced a rerupture of the native tendon due to a fall that occurred 2 months after undergoing an Achilles tendon repair using Matracell treated ADM. The ADM was removed and an extensive histology analysis was performed on the tissue. Additionally, a literature review was conducted to determine the mechanism of ADM integration into the tendon structure and explore if differences in this mechanism exist for different types of human ADMS. The histology analysis demonstrated that the healing process during a tendon reconstruction procedure is similar to that of wound healing. Furthermore, the literature review showed that differences exist in the mechanism for integration among various human ADMs and that these differences may be due to variances in the methods and technologies that manufactures use to process human ADMs.     

Habitual loading results in tendon hypertrophy and increased stiffness of the human patellar tendon.

The purpose of this study was to examine patellar tendon (PT) size and mechanical properties in subjects with a side-to-side strength difference of >/=15% due to sport-induced loading. Seven elite fencers and badminton players were included. Cross-sectional area (CSA) of the PT obtained from MRI and ultrasonography-based measurement of tibial and patellar movement together with PT force during isometric contractions were used to estimate mechanical properties of the PT bilaterally. We found that distal tendon and PT, but not mid-tendon, CSA were greater on the lead extremity compared with the nonlead extremity (distal: 139 +/- 11 vs. 116 +/- 7 mm(2); mid-tendon: 85 +/- 5 vs. 77 +/- 3 mm(2); proximal: 106 +/- 7 vs. 83 +/- 4 mm(2); P < 0.05). Distal tendon CSA was greater than proximal and mid-tendon CSA on both the lead and nonlead extremity (P < 0.05). For a given common force, stress was lower on the lead extremity (52.9 +/- 4.8 MPa) compared with the nonlead extremity (66.0 +/- 8.0 MPa; P < 0.05). PT stiffness was also higher in the lead extremity (4,766 +/- 716 N/mm) compared with the nonlead extremity (3,494 +/- 446 N/mm) (P < 0.05), whereas the modulus did not differ (lead 2.27 +/- 0.27 GPa vs. nonlead 2.16 +/- 0.28 GPa) at a common force. These data show that a habitual loading is associated with a significant increase in PT size and mechanical properties.     

Comparative ultrastructure of the inner capsule of the muscle spindle and the tendon organ

The ultrastructure of inner capsule cells of the vertebrate muscle spindle was studied by transmission electron microscopy and compared with that of homologous cells in the tendon organ. Aside from variations in their complexity and pattern of organization, cells of the inner capsule in these two sensory receptors exhibited marked similarities in fine structure. The virtual absence of basal lamina in the region of the nucleated soma as well as on the branching cytoplasmic extensions of these cells was noted. In the inner capsule of both end organs, three kinds of intercellular specialization were encountered. Cell processes were typically linked together at multiple sites by intermediate junctions. In addition, focal points of membrane fusion between two or more cellular profiles were identified as tight junctions. In more extensive regions of plasma membrane overlap, gap junctions were also discerned. It seems probable that these sites along the inner capsule represent areas of mechanical and electrical linkage, enabling contiguous cells to function as a synchronous unit. Tight junctions may also provide the inner capsular sheath with specific permeability-barrier characteristics. Elements of the Golgi complex and associated presecretory vesicles and cytoplasmic granules were prominent. Their presence implicates these cells in the elaboration of the paracellular connective-tissue matrix occupying the intracapsular spaces of both receptors. The close resemblance of these cells to endoneurial fibroblasts of peripheral nerve and to hyalocytes of the vitreous body is emphasized. It is likely that, regardless of species examined, cells of the inner capsule in both receptors play an overall protective role in the formation, maintenance and regulation of their luminal paracellular contents.     

Plasticity of human Achilles tendon mechanical and morphological properties in response to cyclic strain

The purpose of the current study in combination with our previous published data (Arampatzis et al., 2007) was to examine the effects of a controlled modulation of strain magnitude and strain frequency applied to the Achilles tendon on the plasticity of tendon mechanical and morphological properties. Eleven male adults (23.9±2.2 yr) participated in the study. The participants exercised one leg at low magnitude tendon strain (2.97±0.47%), and the other leg at high tendon strain magnitude (4.72±1.08%) of similar frequency (0.5 Hz, 1 s loading, 1 s relaxation) and exercise volume (integral of the plantar flexion moment over time) for 14 weeks, 4 days per week, 5 sets per session. The exercise volume was similar to the intervention of our earlier study (0.17 Hz frequency; 3 s loading, 3 s relaxation) allowing a direct comparison of the results. Before and after the intervention ankle joint moment has been measured by a dynamometer, tendon–aponeurosis elongation by ultrasound and cross-sectional area of the Achilles tendon by magnet resonance images (MRI). We found a decrease in strain at a given tendon force, an increase in tendon–aponeurosis stiffness and tendon elastic modulus of the Achilles tendon only in the leg exercised at high strain magnitude. The cross-sectional area (CSA) of the Achilles tendon did not show any statistically significant (P>0.05) differences to the pre-exercise values in both legs. The results indicate a superior improvement in tendon properties (stiffness, elastic modulus and CSA) at the low frequency (0.17 Hz) compared to the high strain frequency (0.5 Hz) protocol. These findings provide evidence that the strain magnitude applied to the Achilles tendon should exceed the value, which occurs during habitual activities to trigger adaptational effects and that higher tendon strain duration per contraction leads to superior tendon adaptational responses.     

The effects of Achilles tendon compliance on triceps surae mechanics and energetics in walking

Achilles tendon (AT) compliance can affect the generation and transmission of triceps surae muscle forces, and thus has important biomechanical consequences for walking performance. However, the uniarticular soleus (SOL) and the biarticular (GAS) function differently during walking, with in vivo evidence suggesting that their associated fascicles and tendinous structures exhibit unique kinematics during walking. Given the strong association between muscle fiber length, velocity and force production, we conjectured that SOL and GAS mechanics and energetic behavior would respond differently to altered AT compliance. To test this, we characterized GAS and SOL muscle and tendon mechanics and energetics due to systematic changes in tendon compliance using musculoskeletal simulations of walking. Increased tendon compliance enlarged GAS and SOL tendon excursions, shortened fiber operation lengths and affected muscle excitation patterns. For both muscles, an optimal tendon compliance (tendon strains of approximately 5% with maximum isometric force) existed that minimized metabolic energy consumption. However, GAS muscle-tendon mechanics and energetics were significantly more sensitive to changes in tendon compliance than were those for SOL. In addition, GAS was not able to return stored tendon energy during push-off as effectively as SOL, particularly for larger values of tendon compliance. These fundamental differences between GAS and SOL sensitivity to altered tendon compliance seem to arise from the biarticular nature of GAS. These insights are potentially important for understanding the functional consequences of altered Achilles tendon compliance due to aging, injury, or disease.     

Effect of muscle contraction levels on the force-length relationship of the human Achilles tendon during lengthening of the triceps surae muscle-tendon unit

Findings from animal experiments are sometimes contradictory to the idea that the tendon structure is a simple elastic spring in series with muscle fibers, and suggest influence of muscle contraction on the tendon mechanical properties. The purpose of the present study was to investigate the influence of muscle contraction levels on the force-length relationship of the human Achilles tendon during lengthening of the triceps surae muscle-tendon unit. For seven subjects, ankle dorsiflexion was performed without (passive condition) and with contraction of plantar flexor muscles (eccentric conditions, at 3 contraction levels) on an isokinetic dynamometer. Deformation of the Achilles tendon during each trial was measured using ultrasonography. The Achilles tendon force corresponding to the tendon elongation of 10mm in the passive condition was significantly smaller than those in the eccentric conditions (p<0.05 or p<0.01). Within the eccentric conditions, the Achilles tendon force corresponding to the tendon elongation of 10mm was significantly greater in the maximal contraction level than those in submaximal eccentric conditions (p<0.05 or p<0.01). In addition, the tendon stiffness was greater in higher contraction levels (p<0.05 or p<0.01). Present results suggest that the human tendon structure is not a simple elastic spring in series with muscle fibers.     

The innervation pattern of the human Achilles tendon: studies of the normal and tendinosis tendon with markers for general and sensory innervation

Pain-free normal Achilles tendons and chronic painful Achilles tendons were examined by the use of antibodies against a general nerve marker (protein gene-product 9.5, PGP9.5), sensory markers (substance P, SP; calcitonin gene-related peptide, CGRP), and immunohistochemistry. In the normal tendons, immunoreactions against PGP9.5 and against SP/CGRP were encountered in the paratendinous loose connective tissue, being confined to nerve fascicles and to nerve fibers located in the vicinity of blood vessels. To some extent, these immunoreactions also occurred in the tendon tissue proper. Immunoreaction against PGP9.5 and against SP/CGRP was also observed in the tendinosis samples and included immunoreactive nerve fibers that were intimately associated with small blood vessels. In conclusion, Mechanoreceptors (sensory corpuscles) were occasionally observed, nerve-related components are present in association with blood vessels in both the normal and the tendinosis tendon.     

The effect of exogenous TSH and Achilles tendon reflex time in man.

The effect of exogenous thyrotropin (TSH) -- 3 X 10 IU daily -- on Achilles tendon reflex time (ART) was investigated in the group of 38 in-patients, treated for thyroid cancer. A significant shortening of ART was recorded in a subgroup of hypothyroid persons after the 2nd and 3rd dose of TSH. Moreover, a significant correlation was proved between initial values of ART and their changes after the 2nd and 3rd TSH injection in this group. No significant interrelationship was observed between changes of thyroid function and ART after TSH administration. In interpretation of this finding the possibility of direct effect of TSH on muscle metabolism in hypothyroid man should be considered.     

Medial gastrocnemius muscle fascicle active torque-length and Achilles tendon properties in young adults with spastic cerebral palsy

Individuals with spastic cerebral palsy (CP) typically experience muscle weakness. The mechanisms responsible for muscle weakness in spastic CP are complex and may be influenced by the intrinsic mechanical properties of the muscle and tendon. The purpose of this study was to investigate the medial gastrocnemius (MG) muscle fascicle active torque-length and Achilles tendon properties in young adults with spastic CP. Nine relatively high functioning young adults with spastic CP (GMFCS I, 17±2 years) and 10 typically developing individuals (18±2 years) participated in the study. Active MG torque-length and Achilles tendon properties were assessed under controlled conditions on a dynamometer. EMG was recorded from leg muscles and ultrasound was used to measure MG fascicle length and Achilles tendon length during maximal isometric contractions at five ankle angles throughout the available range of motion and during passive rotations imposed by the dynamometer. Compared to the typically developing group, the spastic CP group had 33% lower active ankle plantarflexion torque across the available range of ankle joint motion, partially explained by 37% smaller MG muscle and 4% greater antagonistic co-contraction. The Achilles tendon slack length was also 10% longer in the spastic CP group. This study confirms young adults with mild spastic CP have altered muscle–tendon mechanical properties. The adaptation of a longer Achilles tendon may facilitate a greater storage and recovery of elastic energy and partially compensate for decreased force and work production by the small muscles of the triceps surae during activities such as locomotion.     

Presence of substance P and the neurokinin-1 receptor in tenocytes of the human Achilles tendon

Nerve signal substances, such as the tachykinin substance P (SP), may be involved in the changes that occur in response to tendinopathy (tendinosis). It is previously known that the level of SP innervation within tendon tissue is limited, but results of experimental studies have suggested that SP may have stimulatory, angiogenetic and healing effects in injured tendons. Therefore, it would be of interest to know if there is a local SP-supply in tendon tissue. In the present study, the patterns of expression of SP and its preferred receptor, the neurokinin-1 receptor (NK-1 R), in normal and tendinosis human Achilles tendons were analyzed by use of both immunohistochemistry and in situ hybridization. We found that there was expression of SP mRNA in tenocytes, and that tenocytes showed expression of NK-1 R at protein as well as mRNA levels. The observations concerning both SP and NK-1 R were most evident for tenocytes in tendinosis tendons. Our findings suggest that SP is produced in tendinosis tendons, and furthermore that SP has marked effects on the tenocytes via the NK-1 R. It cannot be excluded that the SP effects are of importance concerning the processes of reorganization and healing that occur for tendon tissue in tendinosis. In conclusion, it appears as if SPergic autocrine/paracrine effects occur in tendon tissue during the processes of tendinosis, hitherto unknown effects for human tendons.     

The risk of sports activities--the injuries of the Achilles tendon in sportsmen

The aim of this study was to determine whether professional sports activities and intensive training practice affect pathological changes of the Achilles tendon. We also aimed to investigate whether these injuries primarily occur earlier or later in life. This was a cross sectional study of 214 athletes and non-athletes between the ages of 17-66. Participants were divided into four groups: 63 active young athletes, 64 young persons not involved in sports, 6 elderly athletes and 58 elderly people who did not participate in intensive athletic activities during their lifetimes. Ultrasound measurements of the Achilles tendon were performed on all participants. All participants also completed a self-administered survey to collect demographic information and data on athletic participation. Data were analyzed using the chi-square test and Fisher's exact test. Overall, those who were active athletes sought medical care more often than those who did not participate regularly in sports. Twenty-seven (33%) of young athletes had some Achilles tendon pathology, compared to 9 (14%) younger non-athletes. A total of 36 (64%) of elderly athletes were diagnosed with Achilles tendon disorders, compared to 15 (26%) of elderly people who were not active in sports. Microtears, acute tendonitis and peritendonitis were associated with younger age. By contrast, calcifications, scar tissue, and partial or complete rupture were only seen in the elderly