Selected contractile properties and fatigability of the quadriceps muscle were studied in seven spinal cord–injured (SCI) and 13 able‐bodied control (control) individuals. The SCI muscles demonstrated faster rates of contraction and relaxation than did control muscles and extremely large force oscillation amplitudes in the 10‐Hz signal (65 ± 22% in SCI versus 23 ± 8% in controls). In addition, force loss and slowing of relaxation following repeated fatiguing contractions were greater in SCI compared with controls. The faster contractile properties and greater fatigability of the SCI muscles are in agreement with a characteristic predominance of fast glycolytic muscle fibers. Unexpectedly, the SCI muscles exhibited a force–frequency relationship shifted to the left, most likely as the result of relatively large twitch amplitudes. The results indicate that the contractile properties of large human locomotory muscles can be characterized using the approach described and that the transformation to faster properties consequent upon changes in contractile protein expression following SCI can be assessed. These measurements may be useful to optimize stimulation characteristics for functional electrical stimulation and to monitor training effects induced by electrical stimulation during rehabilitation of paralyzed muscles. © 1999 John Wiley & Sons, Inc. Muscle Nerve 22: 1249–1256, 1999.

Six weeks of FES-LCE training increased the cross-sectional area of large conduit arteries and improved blood flow to the paralyzed legs of individuals with SCI.

Study design: A longitudinal training study. Objectives: To assess if contractile speed and fatigability of paralysed quadriceps muscles in individuals with spinal cord injury (SCI) can be altered by functional electrical stimulation leg cycle ergometry (FES-LCE) training. Settings: The Sint Maartenskliniek rehabilitation centre and the University of Nijmegen, Nijmegen, the Netherlands. Methods: Contractile properties of the quadriceps muscle were studied in seven people with motor-complete SCI who participated in a FES-LCE training program. Subjects trained for 30 min, three times per week for 6 weeks. Contractile speed and fatigue characteristics of electrically stimulated isometric contractions were compared before and after 6 weeks of FES-LCE. Results: Fatigue resistance improved following FES-LCE training as indicated by the higher forces maintained in response to repetitive electrical stimulation. In contrast with an improved fatigue resistance, the maximal rate of force rise was unaected, the speed of relaxation increased and the fusion of a 10 Hz force signal decreased. Furthermore, the force-frequency relationship shifted to the right at low stimulation frequencies, indicated by a decline in the ratio of 1 and 100 Hz force responses as well as the ratio of 10 and 100 Hz force responses. Conclusion: FES-LCE training can change the physiological properties of the quadriceps muscle in people with SCI. Even after a short period of training, the stimulated muscles become more resistant to fatigue. Furthermore, the increased speed of relaxation and associated decreased fusion and altered force-frequency relationship following training may be related to adaptations in the calcium handling processes, which re¯ect an early response of long-term disused muscles. Spinal Cord (2000) 38, 214 ± 223

A spinal cord injury usually leads to an increase in contractile speed and fatigability of the paralysed quadriceps muscles, which is probably due to an increased expression of fast myosin heavy chain (MHC) isoforms and reduced oxidative capacity. Sometimes, however, fatigue resistance is maintained in these muscles and also contractile speed is slower than expected. To obtain a better understanding of the diversity of these quadriceps muscles and to determine the effects of training on characteristics of paralysed muscles, fibre characteristics and whole muscle function were assessed in six subjects with spinal cord lesions before and after a 12-week period of daily low-frequency electrical stimulation. Relatively high levels of MHC type I were found in three subjects and this corresponded with a high degree of fusion in 10-Hz force responses (r=0.88). Fatigability was related to the activity of succinate dehydrogenase (SDH) (r=0.79). Furthermore, some differentiation between fibre types in terms of metabolic properties were present, with type I fibres expressing the highest levels of SDH and lowest levels of a-glycerophosphate dehydrogenase. After training, SDH activity increased by 76€26% but fibre diameter and MHC expression remained unchanged. The results indicate that expression of contractile proteins and metabolic properties seem to underlie the relatively normal functional muscle characteristics observed in some paralysed muscles. Furthermore, training-induced changes in fatigue resistance seem to arise, in part, from an improved oxidative capacity.

Effects of two different training regimens on the contractile properties of the quadriceps muscle were studied in six individuals with spinal cord injury. Each subject had both limbs trained with the two regimens, consisting of stimulation with low frequencies (LF) at 10 HZ or high frequencies (HF) at 50 HZ; one limb of each subject was stimulated with the LF protocol and the other with the HF regimen. Twelve weeks of daily training increased tetanic tension by ∼20%, which was not significantly different between training regimens. Interestingly, after HF but not LF training, the unusual high forces at the low frequency range of the force-frequency relationship decreased, possibly due to a reduced activation per impulse. After LF but not HF training, force oscillation amplitudes declined (by 33%) as relaxation tended to slow, which may have opposed possible effects of reduced activation as seen after HF training. Finally, fatigue resistance also increased rapidly after LF training (by 43%) but not after HF training. These results indicate that different types of training may selectively change different aspects of function in disused muscles. Skeletal muscles have the capacity to adapt to altered functional requirements. When muscle activity is reduced, for instance as a result of immobilization 3 or paralysis after spinal cord injury (SCI), 43 muscle mass decreases, leading to a loss of strength. Reduced neuromuscular activity leads not only to a reduction in the total amount of contractile protein but also to changes in the phenotypic properties of the muscle fibers. In general, decreased neuromuscular activity leads to a transition of slow to fast fiber characteristics, with concomitant adaptations of metabolic properties and the peripheral circulation. 30,39 SCI is an example of almost complete loss of neuromuscular activity, leading to muscle atrophy, 6,8,23,37,43 muscle fiber transformation from oxidative type I to glycolytic type II fibers, 5,37 reduction in oxidative metabolism, 24,35 and limited blood flow. 18,27,42 In the past, various programs of electrical stimulation have been used to recondition the paralyzed muscles of individuals with SCI. 16,17,26,28,33,34 A variety of stimulation regimens have been used, most with relatively short contractions (∼1-5 s) elicited at various frequencies (∼20-60 HZ) with duty cycles ranging from ∼1/1 to ∼1/3. However, comparisons as to how paralyzed muscles respond to different types of training regimens are limited. Furthermore, the effects of training are usually assessed in relation to muscle strength, mass, or fatigue resistance, whereas other contractile properties, such as speed or the force-frequency relationship, have rarely been reported for SCI muscle.The purpose of the present investigation was to compare the effects of training with two patterns of electrical stimulation (repetitive high-frequency stimulation and more continuous low-frequency Abbreviations: FOA, force oscillation amplitude; HF, high frequency; LF, low frequency; MFR, maximal ra...

This study investigated the recruitment of type I, IIA and IIAX fibres after seven isometric contractions at 40, 70 and 100% maximal voluntary knee extension torque (MVC, 1 s on/1 s off). Biopsies of the vastus lateralis muscle were collected from seven subjects at rest and immediately post-exercise. Fibre fragments were dissected from the freeze-dried samples and characterized as type I, IIA and IIAX using mATPase staining. Phosphocreatine (PCr) and creatine (Cr) content were measured in the remaining part of characterized fibres. A decline in the ratio of PCr to Cr (PCr/Cr) was used as an indication of activation. The mean peak torques were, respectively, 39 (2), 72 (2) and 87 (6)% MVC. Cumulative distributions of type I and IIA fibres were significantly shifted to lower PCr/Cr ratios at all intensities (Kolmogorov-Smirnov test, P<0.05). The cumulative distribution of type IIAX fibres showed a significant leftward shift only at 87% MVC ( P<0.05). A hierarchical order of fibre activation with increasing intensity of exercise was found, with some indication of rate coding for type I and IIA fibres. Evidence for activation of type IIAX fibres was only found at 87% MVC.

This study assessed the reproducibility of electrically evoked, isometric quadriceps contractile properties in eight people with spinal cord injury (SCI) and eight able-bodied (AB) individuals. Over all, the pooled coefficients of variation (CVps) in the SCI group were significantly lower (ranging from 0.03 to 0.15) than in the AB group (ranging from 0.08 to 0.21) (P<0.05). Furthermore, in all subjects, the variability of force production increased as stimulation frequency decreased (P<0.01). In subjects with SCI, variables of contractile speed are clearly less reproducible than tetanic tension or resistance to fatigue. Contractile properties of quadriceps muscles of SCI subjects were significantly different from that of AB subjects. Muscles of people with SCI were less fatigue resistant (P<0.05) and produced force-frequency relationships that were shifted to the left, compared with AB controls (P<.01). In addition, fusion of force responses resulting from 10 Hz stimulation was reduced (P<.05) and speed of contraction (but not relaxation) was increased (P<0.05), indicating an increased contractile speed in paralysed muscles compared with non-paralysed muscles. These results correspond with an expected predominance of fast glycolytic muscle fibres in paralysed muscles. It is concluded that quadriceps dynamometry is a useful technique to study muscle function in non-paralysed as well as in paralysed muscles. Furthermore, these techniques can be reliably used, for example, to assess therapeutic interventions on paralysed muscles provided that expected differences in relative tetanic tension and fatigue resistance are larger than approximately 5% and differences in contractile speed are larger than approximately 15%.

Low muscle temperature in paralysed muscles of individuals with spinal cord injury may affect the contractile properties of these muscles. The present study was therefore undertaken to assess the effects of increased muscle temperature on the isometric contractile properties of electrically stimulated paralysed quadriceps muscles. When muscle temperature at a depth of 3 cm was increased from approximately 32 degrees C to approximately 36 degrees C by ultra-short-wave application, the half-relaxation time shortened and low-frequency force responses became less fused, but the maximal rate of increase in force remained unchanged. Heating had no effect upon either force decline or slowing of relaxation during fatiguing contractions. The force-frequency relationship of the paralysed quadriceps muscle was shifted to the right after the muscle was heated. Despite this shift, however, the relationship still resembled that in muscles of non-paralysed individuals, probably due to the unexplained high twitch forces. These results indicate that reduced muscle temperature in spinal-cord-injured individuals may lead to an underestimation of the changes in contractile properties in terms of relaxation rate or the degree of fusion with low-frequency stimulation. In addition, the force-frequency relationship of paralysed muscles does not accurately reflect the magnitude of these changes, even when the muscle is heated, and should therefore be treated with caution.