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1
Ali, Farah, Peter D. Paré e Chun Y. Seow. "Models of contractile units and their assembly in smooth muscle". Canadian Journal of Physiology and Pharmacology 83, n. 10 (1 ottobre 2005): 825–31. http://dx.doi.org/10.1139/y05-052.
Testo completoAbstract (sommario):
It is believed that the contractile filaments in smooth muscle are organized into arrays of contractile units (similar to the sarcomeric structure in striated muscle), and that such an organization is crucial for transforming the mechanical activities of actomyosin interaction into cell shortening and force generation. Details of the filament organization, however, are still poorly understood. Several models of contractile filament architecture are discussed here. To account for the linear relationship observed between the force generated by a smooth muscle and the muscle length at the plateau of an isotonic contraction, a model of contractile unit is proposed. The model consists of 2 dense bodies with actin (thin) filaments attached, and a myosin (thick) filament lying between the parallel thin filaments. In addition, the thick filament is assumed to span the whole contractile unit length, from dense body to dense body, so that when the contractile unit shortens, the amount of overlap between the thick and thin filaments (i.e., the distance between the dense bodies) decreases in exact proportion to the amount of shortening. Assembly of the contractile units into functional contractile apparatus is assumed to involve a group of cells that form a mechanical syncytium. The contractile apparatus is assumed malleable in that the number of contractile units in series and in parallel can be altered to accommodate strains on the muscle and to maintain the muscle's optimal mechanical function.Key words: contraction model, ultrastructure, length adaptation, plasticity.
2
Thomas, C. K., R. S. Johansson e B. Bigland-Ritchie. "Attempts to physiologically classify human thenar motor units". Journal of Neurophysiology 65, n. 6 (1 giugno 1991): 1501–8. http://dx.doi.org/10.1152/jn.1991.65.6.1501.
Testo completoAbstract (sommario):
1. This study was designed to determine whether human thenar motor units can be classified into types by the same physiological criteria used for other mammalian limb motor units and to consider whether such classification is functionally relevant. 2. Contractile responses of 25 human thenar single motor units were examined when their motor axons were stimulated intraneurally at rates from 1 to 100 Hz and intermittently at 40 Hz in a conventional 2-min fatigue test. Twitch and tetanic forces were measured together with various indexes of contractile rate. 3. Twitch contraction times and subtetanic to maximum tetanic force ratios were both distributed continuously. "Sag" in tension was not evident in unfused force profiles. Thus these units could not be divided into fast and slow types by the use of traditional contractile rate criteria. 4. Most units were fatigue resistant, with force fatigue indexes (FI) ranging from 0.33 to 1.14. None could be classified as fatiguable (FI less than 0.25). Seven units (28%) fell into the fatigue-intermediate (FI = 0.25-0.75) category, whereas 18 units (72%) had FI greater than 0.75, i.e., they were fatigue-resistant units. However, these units could not be classified by conventional FI and contractile rate criteria, because fatigue-resistant and fatigue-intermediate units had similar contractile rates. 5. Additional FI were calculated to describe changes in contractile rate. During the fatigue test, units behaved in one of three ways, showing 1) little change in either force or rate; 2) contractile slowing during the contraction and relaxation phases, with little or no force loss; or 3) both force and rate reduction.(ABSTRACT TRUNCATED AT 250 WORDS)
3
Mateika, J. H., E. G. Essif, C. Dellorusso e R. F. Fregosi. "Contractile Properties of Human Nasal Dilator Motor Units". Journal of Neurophysiology 79, n. 1 (1 gennaio 1998): 371–78. http://dx.doi.org/10.1152/jn.1998.79.1.371.
Testo completoAbstract (sommario):
Mateika, J. H., E. G. Essif, C. DelloRusso, and R. F. Fregosi. Contractile properties of human nasal dilator motor units. J. Neurophysiol. 79: 371–378, 1998. The technique of intramuscular microstimulation was used to activate facial nerve fibers while acquiring simultaneous twitch force measurements to measure the contractile properties and force-frequency responses of human nasal dilator (ND) motor units. Twitch force amplitude (TF), contraction time (CT), half-relaxation time (HRT), and the maximal rate of rise of force normalized to the peak force (maximum contraction rate, MCR) were recorded from 98 ND motor units in 37 subjects. The average CT, HRT, MCR, and TF were 47.9 ± 1.8 ms, 42.6 ± 2.1 ms, 28.6 ± 1.8 s−1, and 1.06 ± 0.1 mN, respectively. Neither CT nor HRT were significantly correlated with TF. The average CT and HRT were similar to values recorded for small muscles of the hand but were faster than the values recorded from human toe extensor motor units. However the lack of an association between twitch force and CT or HRT was similar to the findings obtained for both human hand and foot muscles. Force-frequency curves were recorded from eight ND motor units. The force produced by the eight motor units was recorded in response to stimuli delivered at 1, 5, 10, 15, 20, 25, 30, 35, and 40 Hz to assess force-frequency relationships. The mean twitch force of the eight motor units was 0.91 ± 0.3 mN and the average tetanic force was 8.1 ± 1.8 mN. Therefore the average twitch force was equal to 12.7% of the tetanic force. Fifty percent of the unit tetanic force was achieved at an average frequency of 16.4 ± 1.7 Hz, which is greater than the value recorded for human toe extensor motor units (9.6 Hz). Thus the force produced by the ND motor units was more sensitive to changes in discharge frequency over the range of ∼10–30 Hz and less sensitive to changes in the range of 0–10 Hz because of their fast contractile properties. The mean slope of the regression lines that were fit to the steep portion of each force-frequency curve was 5.15 ± 0.5% change in force/Hz. This value was greater than the slope measured for human toe extensor muscles (4.2% change in force/Hz). These observations suggest that force gradation by ND motor units is more sensitive to changes in stimulation frequency than human toe extensor motor units. We conclude that most ND motor units have fast contractile properties and that rate coding may play a significant role in the gradation of force produced by the ND muscle. Furthermore, the findings of this investigation have demonstrated that contractile speed and TF in a human facial muscle are not correlated. This supports previous findings obtained from human hand and foot muscles and suggests that there may be a fundamental difference in the contractile speed-twitch force relationship between many human muscles and most muscles of other mammals.
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Doherty, Timothy J., e William F. Brown. "Age-related changes in the twitch contractile properties of human thenar motor units". Journal of Applied Physiology 82, n. 1 (1 gennaio 1997): 93–101. http://dx.doi.org/10.1152/jappl.1997.82.1.93.
Testo completoAbstract (sommario):
Doherty, Timothy J., and William F. Brown. Age-related changes in the twitch contractile properties of human thenar motor units. J. Appl. Physiol. 82(1): 93–101, 1997.—The purpose of this study was to examine the effects of aging on the contractile and electrophysiological properties of human thenar motor units (MUs). Percutaneous electrical stimulation of single motor axons within the median nerve was used to isolate and examine the twitch tensions, contractile speeds, and surface-detected MU action potential (S-MUAP) sizes of 48 thenar MUs in 17 younger subjects (25–53 yr) and 44 thenar MUs in 9 older subjects (64–77 yr). A wide range of twitch tensions, contractile speeds, and S-MUAP sizes was observed in both age groups. However, older subjects had significantly larger MU twitch tensions and slower MU twitch contraction and half-relaxation times. These changes were accompanied by increased S-MUAP sizes. These findings suggest that the human thenar MU pool undergoes significant age-related increase in MU size and slowing of contractile speed. Such adaptation may help to overcome previously reported age-related losses of thenar MUs.
5
Foehring, R. C., G. W. Sypert e J. B. Munson. "Properties of self-reinnervated motor units of medial gastrocnemius of cat. I. Long-term reinnervation". Journal of Neurophysiology 55, n. 5 (1 maggio 1986): 931–46. http://dx.doi.org/10.1152/jn.1986.55.5.931.
Testo completoAbstract (sommario):
This work tested whether the membrane electrical properties of cat motoneurons, the contractile properties of their muscle units, and the normal relationships among them would be restored 9 mo after section and resuture of their muscle nerve. Properties of medial gastrocnemius (MG) motor units were examined 9 mo following section and resuture of the MG nerve in adult cats. Motoneuron electrical properties and muscle-unit contractile properties were measured. Motor units were classified on the basis of their contractile properties as type fast twitch, fast fatiguing (FF), fast twitch with intermediate fatigue resistance (FI), fast twitch, fatigue resistant (FR), or slow twitch, fatigue resistant (S) (8, 20). Muscle fibers were classified as type fast glycolytic (FG), fast oxidative glycolytic (FOG), or slow oxidative (SO) on the basis of histochemical staining for myosin adenosine triphosphatase, nicotinamide adenine dinucleotide diaphorase, and alpha-glycerophosphate dehydrogenase (48). Following 9 mo self-reinnervation, the proportions of each motor-unit type were the same as in normal control animals. Motoneuron membrane electrical properties [axonal conduction velocity, afterhyperpolarization (AHP) half-decay time, rheobase, and input resistance] also returned to control levels in those motoneurons that made functional reconnection with the muscle (as determined by ability to elicit measurable tension). The relationships among motoneuron electrical properties were normal in motoneurons making functional reconnection. Approximately 10% of MG motoneurons sampled did not elicit muscle contraction. These cells' membrane electrical properties were different from those that did elicit muscle contraction. Contractile speed and fatigue resistance of reinnervated muscle units had recovered to control levels at 9 mo postoperation. Force generation did not recover fully in type-FF units. The reduced tensions were apparently due to failure of recovery of FG muscle fiber area. Following reinnervation, relationships between motoneuron electrical and muscle-unit contractile properties were similar to controls. This was reflected in a degree of correspondence between motor-unit type and motoneuron type similar to normal units (84 vs. 86%, as defined by Ref. 61). There was a significantly increased proportion of type-SO muscle fibers and a decrease in the fast muscle fibers (especially type FOG) in 9 mo reinnervated MG. Together with the unchanged proportions of motor-unit types, this led to an estimate of average innervation ratios being increased in type-S motor units and decreased in type-FR units.(ABSTRACT TRUNCATED AT 400 WORDS)
6
Sutlive, Thomas G., J. Ross McClung e Stephen J. Goldberg. "Whole-Muscle and Motor-Unit Contractile Properties of the Styloglossus Muscle in Rat". Journal of Neurophysiology 82, n. 2 (1 agosto 1999): 584–92. http://dx.doi.org/10.1152/jn.1999.82.2.584.
Testo completoAbstract (sommario):
Investigations of whole muscle and motor-unit contractile properties have provided valuable information for our understanding of the spinal cord and extraocular motor systems. However, no previous investigation has examined these properties in an isolated tongue muscle. The purpose of this study was to determine the contractile properties and muscle fiber types of the rat styloglossus muscle. The styloglossus is one of three extrinsic tongue muscles and serves to retract the tongue within the oral cavity. Adult male Sprague-Dawley rats ( n = 19) were used in these experiments. The contractile characteristics of the whole styloglossus muscle ( n = 9) were measured in response to stimulation of the hypoglossal nerve branch to the muscle. The average twitch tension produced was 3.30 g with a mean twitch contraction time of 13.81 ms. The mean maximum tetanic tension was 19.66 g and occurred at or near the fusion frequency, which averaged 109 Hz. The styloglossus muscle was resistant to fatigue [fatigue index (F. I.) = 0.76]. In separate experiments ( n = 7), the contractile characteristics of 37 single motor units were measured in response to extracellular stimulation of hypoglossal motoneurons. The twitch tension generated by styloglossus motor units averaged 35.7 mg, and the mean twitch contraction time was 12.46 ms. The mean fusion frequency was 92 Hz. Maximum tetanic tension averaged 177.8 mg. Styloglossus single motor units were resistant to fatigue (F. I. = 0.74). The sites of stimulation that yielded a contractile response in the styloglossus muscle were consistent with the location of the styloglossus motoneuron pool reported in earlier anatomy studies. Muscle fiber typing was determined in three animals based on the myofibrillar ATPase reaction at pH 9.8, 4.6, and 4.3. The styloglossus muscle was composed of ≈99% type IIA fibers with a few scattered type I fibers present in the study sample. On the basis of the combined findings of the physiology and histochemistry experiments, the styloglossus muscle appeared to be a homogeneous muscle composed almost exclusively of fast, fatigue-resistant motor units. These properties of the styloglossus muscle and its motor units were compared with findings in other rat skeletal muscles.
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Łochyński, Dawid, Dominik Kaczmarek, Włodzimierz Mrówczyński, Wojciech Warchoł, Joanna Majerczak, Janusz Karasiński, Michał Korostyński, Jerzy A. Zoladz e Jan Celichowski. "Contractile properties of motor units and expression of myosin heavy chain isoforms in rat fast-type muscle after volitional weight-lifting training". Journal of Applied Physiology 121, n. 4 (1 ottobre 2016): 858–69. http://dx.doi.org/10.1152/japplphysiol.00330.2016.
Testo completoAbstract (sommario):
Dynamic resistance training increases the force and speed of muscle contraction, but little is known about modifications to the contractile properties of the main physiological types of motor units (MUs) that contribute to these muscle adaptations. Although the contractile profile of MU muscle fibers is tightly coupled to myosin heavy chain (MyHC) protein expression, it is not well understood if MyHC transition is a prerequisite for modifications to the contractile characteristics of MUs. In this study, we examined MU contractile properties, the mRNA expression of MyHC, parvalbumin, and sarcoendoplasmic reticulum Ca2+pump isoforms, as well as the MyHC protein content after 5 wk of volitional progressive weight-lifting training in the medial gastrocnemius muscle in rats. The training had no effect on MyHC profiling or Ca2+-handling protein gene expression. Maximum force increased in slow (by 49%) and fast (by 21%) MUs. Within fast MUs, the maximum force increased in most fatigue-resistant and intermediate but not most fatigable MUs. Twitch contraction time was shortened in slow and fast fatigue-resistant MUs. Twitch half-relaxation was shortened in fast most fatigue-resistant and intermediate MUs. The force-frequency curve shifted rightward in fast fatigue-resistant MUs. Fast fatigable MUs fatigued less within the initial 15 s while fast fatigue-resistant units increased the ability to potentiate the force within the first minute of the standard fatigue test. In conclusion, at the early stage of resistance training, modifications to the contractile characteristics of MUs appear in the absence of MyHC transition and the upregulation of Ca2+-handling genes.
8
Łochyński, Dawid, Marcin Bączyk, Dominik Kaczmarek, Maria Jolanta Rędowicz, Jan Celichowski e Piotr Krutki. "Adaptations in physiological properties of rat motor units following 5 weeks of whole-body vibration". Applied Physiology, Nutrition, and Metabolism 38, n. 9 (settembre 2013): 913–21. http://dx.doi.org/10.1139/apnm-2012-0478.
Testo completoAbstract (sommario):
The purpose of the study was to determine the effects of 5-week whole-body vibration (WBV) on contractile parameters and force–frequency relationship of functionally isolated motor units of the rat medial gastrocnemius muscle: fast fatigable (FF), fast fatigue-resistant (FR), and slow (S). Moreover, myosin heavy chain isoform content was quantified. Following WBV, the maximum tetanic force of FF units was increased by ∼25%. The twitch half-relaxation time in all types of motor units and the twitch contraction time in FR units were shortened. The twitch-to-tetanus force ratio was decreased and the force–frequency curves were shifted rightwards in S and FR units. Myosin heavy chain distribution was not changed. These findings suggest modifications of the excitation–contraction coupling towards shortening of a twitch contraction. The observed increase in force of FF units may contribute to gains in muscle dynamic strength reported following WBV treatment.
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Klass, M., S. Baudry e J. Duchateau. "Contractile properties of single motor units in elderly". Computer Methods in Biomechanics and Biomedical Engineering 8, sup1 (settembre 2005): 167–68. http://dx.doi.org/10.1080/10255840512331388786.
Testo completo
10
Nelson, J. S., S. J. Goldberg e J. R. McClung. "Motoneuron electrophysiological and muscle contractile properties of superior oblique motor units in cat". Journal of Neurophysiology 55, n. 4 (1 aprile 1986): 715–26. http://dx.doi.org/10.1152/jn.1986.55.4.715.
Testo completoAbstract (sommario):
Intracellular techniques were used to study single motor units of the trochlear nucleus and superior oblique muscle in the cat. Motoneuron electrophysiological properties were correlated with muscle-unit contractile characteristics assessed under isometric conditions. Two distinct motor-unit types were identified and designated as twitch and nontwitch. Nontwitch units made up 5% of the total population studied. They responded only to tetanic stimulation with graded force that increased as stimulus frequency was increased up to 300-400 Hz. These units made up a homogeneous population in that they were innervated by slowly conducting axons, produced weak tetanic tensions, and were extremely fatigue resistant. Twitch units made up the majority (95%) of units studied. These units responded to single pulse stimulation with typical twitch contractions. The contraction speed and tension ranges for these units were comparable with those obtained from other extraocular muscle single units. Superior oblique twitch units, mechanically comparable with multiply innervated conducting units, identified in the cat inferior oblique muscle (31) were not observed. The twitch-unit population was heterogeneous in terms of neuromuscular fatigue resistance. Unit fatigability was inversely related to maximal tetanic tension. Motoneuron conduction velocity was related to muscle-unit contractile properties in a way similar to that seen in extremity motor units. The slowest twitch units were weak, fatigue resistant, and innervated by slow conducting axons. The fastest units were, in general, innervated by faster conducting axons, produced greater tetanic tensions, and were more susceptible to fatigue. Correlations among input resistance, rheobase, and conduction velocity were also observed. At present, subdivisions of the twitch-unit population on the basis of any one or combination of unit properties does not seem appropriate.
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Foehring, R. C., G. W. Sypert e J. B. Munson. "Motor-unit properties following cross-reinnervation of cat lateral gastrocnemius and soleus muscles with medial gastrocnemius nerve. I. Influence of motoneurons on muscle". Journal of Neurophysiology 57, n. 4 (1 aprile 1987): 1210–26. http://dx.doi.org/10.1152/jn.1987.57.4.1210.
Testo completoAbstract (sommario):
This study addresses two questions: is reinnervation of mammalian skeletal muscle selective with respect to motor-unit type? And to what degree may muscle-unit contractile properties be determined by the motoneuron? Properties of individual motor units were examined following cross-reinnervation (X-reinnervation) of lateral gastrocnemius (LG) and soleus muscles by the medial gastrocnemius (MG) nerve in the cat. We examined animals at two postoperative times: 9-10 wk (medX) and 9-11 mo (longX). For comparison, properties of normal LG and soleus motor units were studied. Motor units were classified on the basis of their contractile response as fast contracting fatigable, fast intermediate, fast contracting fatigue resistant, or slow (types FF, FI, FR, or S, respectively) (13,29). Muscle fibers were classified on the basis of histochemical properties as fast glycolytic, fast oxidative glycolytic, or slow oxidative (types FG, FOG, or SO, respectively) (61). Reinnervation of LG and soleus was not selective with respect to motor-unit type. Both muscles were innervated by a full complement of MG motoneuron types, apparently in normal MG proportions. MG motoneurons determined LG muscle fibers' properties to a similar degree as reinnervated MG muscle fibers. In contrast, soleus muscle fibers "resisted" the influence of MG motoneurons. Thus, although longX-reinnervated LG muscle (longX LG) had a motor-unit type distribution similar to normal or self-reinnervated MG, longX soleus contained predominantly type S motor units. Overall mean values for muscle-unit contractile properties reflected this motor-unit type distribution. Muscle units in longX LG and longX soleus had contractile properties typical of the same motor-unit type in normal LG or soleus, respectively. Motor-unit types were recognizable at 10 wk X-reinnervation, although muscle-unit tensions were lower than after 10 mo. The proportions of fast and slow motor units in medX LG were similar to longX LG, although a greater proportion of fast units were resistant to fatigue at 10 wk. There were fewer fast units in medX soleus than longX soleus, which suggested that motor-unit type conversion or innervation of muscle fibers by fast motoneurons is not complete at 10 wk. We conclude that reinnervation of the LG and soleus muscles by MG motoneurons was not selective with respect to motor-unit type. MG motoneurons determined LG muscle properties to a similar degree as self-reinnervated MG muscle fibers. Soleus muscle fibers resisted the influence of MG motoneurons, representing a limit to neural determination of muscle properties.
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Yoshitake, Yasuhide, Minoru Shinohara, Hidetoshi Ue e Toshio Moritani. "Characteristics of surface mechanomyogram are dependent on development of fusion of motor units in humans". Journal of Applied Physiology 93, n. 5 (1 novembre 2002): 1744–52. http://dx.doi.org/10.1152/japplphysiol.00008.2002.
Testo completoAbstract (sommario):
The purpose of this study was to test whether surface mechanomyogram (MMG) recorded on the skin reflects the contractile properties of individual motor units in humans. Eight motor units in the medial gastrocnemius muscle were identified, and trains of stimulation at 5, 10, 15, and 20 Hz were delivered to each isolated motor unit. There was a significant positive correlation between the duration of MMG and twitch duration. MMG amplitude decreased with increasing stimulation frequency. Reductions in MMG amplitude were in parallel with the reductions in force fluctuations, and the rate of change in both was positively correlated across the motor units. Rate of change in MMG amplitude against force was negatively correlated to half relaxation time and twitch duration. Similar negative correlations were found between force fluctuations and contractile properties. These results provide evidence supporting a direct relation between MMG and contractile properties of individual motor units within the gastrocnemius muscle, indicating that surface MMG is dependent on the contractile properties of the activated motor units in humans.
13
Fournier, M., e G. C. Sieck. "Mechanical properties of muscle units in the cat diaphragm". Journal of Neurophysiology 59, n. 3 (1 marzo 1988): 1055–66. http://dx.doi.org/10.1152/jn.1988.59.3.1055.
Testo completoAbstract (sommario):
1. Muscle units in the right sternocostal region of the cat diaphragm (DIA) were isolated in situ by dissecting filaments of the C5 ventral root. Isometric contractile and fatigue properties of DIA units were then measured. Contractile properties included: twitch contraction time (CT), peak twitch tension (Pt), maximum tetanic tension (P0), and the frequency dependence of tension production. Muscle-unit fatigue resistance was estimated using a 2-min fatigue test. 2. DIA muscle units were classified as fast (F) or slow (S) based on the presence or absence of sag in their unfused tetanic force responses. Muscle-unit fatigue indices (FI) were used to further classify DIA units as slow-twitch fatigue-resistant (S), fast-twitch fatigue-resistant (FR) fast-twitch fatigue-intermediate (FInt), or fast-twitch fatigable (FF) types. 3. Based on a total of 47 completely characterized DIA muscle units, 21% were classified as S, 4% as FR, 28% as FInt, and 47% as FF. In contrast to the distribution of unit types in other mixed appendicular muscles, the DIA was composed of a very low proportion of FR units and a relatively high proportion of FInt units. An interval of FIs between 0.50 and 0.75 separated units into fatigue-resistant and fatigable groups. The distribution of FIs for FF and most FInt units was continuous, indicating that they formed a single fatigable group. Relatively few FF units in the DIA had FIs less than 0.10. 4. A wide range of contractile properties was observed for DIA muscle units. Type S units had longer CTs and lower Pt and P0 values than type F units. The mean Pt and P0 of FF and FInt units were comparable, whereas the mean Pt and P0 of the two FR units were lower. Type S units produced a greater proportion of their P0 at lower frequencies of activation than type F units. The lower P0S produced by type F units in the DIA indicated that they were smaller than similar units in appendicular muscles. It was concluded that in meeting most normal ventilatory requirements, adequate force could be generated by the recruitment of only type S and FR units. The recruitment of the more fatigable FF and FInt units may occur only during more forceful respiratory and nonrespiratory behaviors of the DIA.
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McAndrew, D., M. Gorelick e J. M. M. Brown. "MUSCLES WITHIN MUSCLES: A MECHANOMYOGRAPHIC ANALYSIS OF MUSCLE SEGMENT CONTRACTILE PROPERTIES WITHIN HUMAN GLUTEUS MAXIMUS". Journal of Musculoskeletal Research 10, n. 01 (marzo 2006): 23–35. http://dx.doi.org/10.1142/s0218957706001704.
Testo completoAbstract (sommario):
The aim of this investigation was to determine the contractile properties of motor units within 3 segments of the gluteus maximus utilizing a laser-based mechanomyographic (MMG) technique. The intention was to determine whether there were segmental differences in motor unit contractile properties and whether these differences may be related to the muscle segment's function and its fibre type composition. Ten subjects were recruited from the student population at the University of Wollongong. Maximal percutaneous neuromuscular stimulation (PNS) was delivered to the medial and lateral portions of three (cranial, middle, caudal) muscle segments of the gluteus maximus by an MMG stimulator. An MMG laser sensor measured the lateral displacement of the muscle segment belly resulting from the development of maximal isometric tension. Parameters characterizing the MMG waveforms were statistically compared to determine variations in contractile properties both within (medial to lateral) and between segments. Our results indicated that the contractile properties of motor units varied significantly (p < 0.05) between, but not within (medial to lateral), the three segments of the gluteus maximus. Most the gluteus maximus. Most notably, segment contraction time (t c ) decreased significantly (p < 0.05) in a cranio to caudal direction suggesting a variation in muscle fibre type composition within the three segments of the muscle. Even when corrected for differences in muscle belly displacement between subjects, the cranial segment was found to have a significantly (p < 0.05) longer contraction time than the two more caudal segments. The results suggest that the gluteus maximus was composed of muscle segments that were physiologically, as well as anatomically, designed to fulfil particular roles during everyday motor tasks. Based upon these results, the MMG technique appears to have considerable utility for the non-invasive assessment of muscle segment contractile properties for both laboratory and clinical applications.
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Carp, J. S., P. A. Herchenroder, X. Y. Chen e J. R. Wolpaw. "Sag During Unfused Tetanic Contractions in Rat Triceps Surae Motor Units". Journal of Neurophysiology 81, n. 6 (1 giugno 1999): 2647–61. http://dx.doi.org/10.1152/jn.1999.81.6.2647.
Testo completoAbstract (sommario):
Sag during unfused tetanic contractions in rat triceps surae motor units. Contractile properties and conduction velocity were studied in 202 single motor units of intact rat triceps surae muscles activated by intra-axonal (or intra-myelin) current injection in L5 or L6 ventral root to assess the factors that determine the expression of sag (i.e., decline in force after initial increase during unfused tetanic stimulation). Sag was consistently detected in motor units with unpotentiated twitch contraction times <20 ms. However, the range of frequencies at which sag was expressed varied among motor units such that there was no single interstimulus interval (ISI), with or without adjusting for twitch contraction time, at which sag could be detected reliably. Further analysis indicated that using the absence of sag as a criterion for identifying slow-twitch motor units requires testing with tetani at several different ISIs. In motor units with sag, the shape of the force profile varied with tetanic frequency and contractile properties. Simple sag force profiles (single maximum reached late in the tetanus followed by monotonic decay) tended to occur at shorter ISIs and were observed more frequently in fatigue-resistant motor units with long half-relaxation times and small twitch amplitudes. Complex sag profiles reached an initial maximum early in the tetanus, tended to occur at longer ISIs, and were more common in fatigue-sensitive motor units with long half-relaxation times and large twitch amplitudes. The differences in frequency dependence and force maximum location suggested that these phenomena represented discrete entities. Successive stimuli elicited near-linear increments in force during tetani in motor units that never exhibited sag. In motor units with at least one tetanus displaying sag, tetanic stimulation elicited large initial force increments followed by rapidly decreasing force increments. That the latter force envelope pattern occurred in these units even in tetani without sag suggested that the factors responsible for sag were expressed in the absence of overt sag. The time-to-peak force (TTP) of the individual contractions during a tetanus decreased in tetani with sag. Differences in the pattern of TTP change during a tetanus were consistent with the differences in force maximum location between tetani exhibiting simple and complex sag. Tetani from motor units that never exhibited sag did not display a net decrease in TTP during successive contractions. These data were consistent with the initial force decrement of sag resulting from a transient reduction in the duration of the contractile state.
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Häger-Ross, C. K., C. S. Klein e C. K. Thomas. "Twitch and Tetanic Properties of Human Thenar Motor Units Paralyzed by Chronic Spinal Cord Injury". Journal of Neurophysiology 96, n. 1 (luglio 2006): 165–74. http://dx.doi.org/10.1152/jn.01339.2005.
Testo completoAbstract (sommario):
Little is known about how human motor units respond to chronic paralysis. Our aim was to record surface electromyographic (EMG) signals, twitch forces, and tetanic forces from paralyzed motor units in the thenar muscles of individuals ( n = 12) with chronic (1.5–19 yr) cervical spinal cord injury (SCI). Each motor unit was activated by intraneural stimulation of its motor axon using single pulses and trains of pulses at frequencies between 5 and 100 Hz. Paralyzed motor units ( n = 48) had small EMGs and weak tetanic forces ( n = 32 units) but strong twitch forces, resulting in half-maximal force being achieved at a median of only 8 Hz. The distributions for cumulative twitch and tetanic forces also separated less for paralyzed units than for control units, indicating that increases in stimulation frequency made a smaller relative contribution to the total force output in paralyzed muscles. Paralysis also induced slowing of conduction velocities, twitch contraction times and EMG durations. However, the elevated ratios between the twitch and the tetanic forces, but not contractile speed, correlated significantly with the extent to which unit force summated in response to different frequencies of stimulation. Despite changes in the absolute values of many electrical and mechanical properties of paralyzed motor units, most of the distributions shifted uniformly relative to those of thenar units obtained from control subjects. Thus human thenar muscles paralyzed by SCI retain a population of motor units with heterogeneous contractile properties because chronic paralysis influenced all of the motor units similarly.
17
Dutka, T. L., C. R. Lamboley, M. J. McKenna, R. M. Murphy e G. D. Lamb. "Effects of carnosine on contractile apparatus Ca2+ sensitivity and sarcoplasmic reticulum Ca2+ release in human skeletal muscle fibers". Journal of Applied Physiology 112, n. 5 (1 marzo 2012): 728–36. http://dx.doi.org/10.1152/japplphysiol.01331.2011.
Testo completoAbstract (sommario):
There is considerable interest in potential ergogenic and therapeutic effects of increasing skeletal muscle carnosine content, although its effects on excitation-contraction (EC) coupling in human muscle have not been defined. Consequently, we sought to characterize what effects carnosine, at levels attained by supplementation, has on human muscle fiber function, using a preparation with all key EC coupling proteins in their in situ positions. Fiber segments, obtained from vastus lateralis muscle of human subjects by needle biopsy, were mechanically skinned, and their Ca2+ release and contractile apparatus properties were characterized. Ca2+ sensitivity of the contractile apparatus was significantly increased by 8 and 16 mM carnosine (increase in pCa50 of 0.073 ± 0.007 and 0.116 ± 0.006 pCa units, respectively, in six type I fibers, and 0.063 ± 0.018 and 0.103 ± 0.013 pCa units, respectively, in five type II fibers). Caffeine-induced force responses were potentiated by 8 mM carnosine in both type I and II fibers, with the potentiation in type II fibers being entirely explicable by the increase in Ca2+ sensitivity of the contractile apparatus caused by carnosine. However, the potentiation of caffeine-induced responses caused by carnosine in type I fibers was beyond that expected from the associated increase in Ca2+ sensitivity of the contractile apparatus and suggestive of increased Ca2+-induced Ca2+ release. Thus increasing muscle carnosine content likely confers benefits to muscle performance in both fiber types by increasing the Ca2+ sensitivity of the contractile apparatus and possibly also by aiding Ca2+ release in type I fibers, helping to lessen or slow the decline in muscle performance during fatiguing stimulation.
18
Gielen, Stan C. A. M. "What Does EMC Tell Us about Muscle Function?" Motor Control 3, n. 1 (gennaio 1999): 9–11. http://dx.doi.org/10.1123/mcj.3.1.9.
Testo completoAbstract (sommario):
EMG recordings are frequently used to obtain a better understanding in the coordination of movements. However, EMG activity is made up by the weighted summation of activity of many motor units with different contractile properties. Recent studies have revealed that different motor units contribute to muscle force in different motor tasks. The flexible recruitment of motor units with various contractile properties allows a flexible tuning of muscle properties, but also complicates the interpretation of EMG activity.
19
Botterman, B. R., e T. C. Cope. "Motor-unit stimulation patterns during fatiguing contractions of constant tension". Journal of Neurophysiology 60, n. 4 (1 ottobre 1988): 1198–214. http://dx.doi.org/10.1152/jn.1988.60.4.1198.
Testo completoAbstract (sommario):
1. Through computer feedback control, muscle-unit tension was maintained by altering the stimulation rate of a functionally isolated motor axon. The required stimulation patterns and fatigue properties of motor units from the flexor carpi radialis (FCR), flexor digitorum longus (FDL), and medial gastrocnemius (MG) muscles of the cat were studied when tension was maintained or "clamped" at a constant average level (25% of maximum tetanic tension). 2. In each muscle, two distinct stimulation patterns were observed during constant-tension contractions, one associated with slow-twitch units and the other with fast-twitch units. Once target tension was reached, slow-twitch units required fairly constant rates in order to maintain a constant force, whereas fast-twitch units displayed a marked decline in rate during the early phases of the contraction, averaging between 42 and 54% for the three muscles. The decline in rate most likely represented potentiation of the contractile response and slowing of contractile speed. In general, slow-twitch units responded with lower mean rates (approximately 14 pps less), averaged over the course of the contraction, than fast-twitch units. 3. For fast-twitch units of each muscle, resistance to fatigue varied continuously and over a wide range. The duration that tension could be maintained at 25% of maximum, defined as endurance time, ranged between 16 and 2063 s. No categorization of fast-twitch units into groups could be made on the basis of endurance time. Of the 5 slow-twitch units followed beyond 2700 s, only one failed to maintain tension during the observation period. 4. For hindlimb fast-twitch units, endurance was independent of the stimulation rate needed to maintain tension during the contraction. By contrast, there was a significant tendency for an inverse relation between endurance time and mean stimulation rate for FCR fast-twitch units. 5. Recovery of maximum tension was evaluated at 30 s, 1 min, 2 min, and 5 min following a constant-tension contraction. After a 5-min rest, fast-twitch units were able to produce an average of 80-85% of their maximum tetanic tension. By using the median endurance time (approximately 100 s) to divide the fast-twitch population into "low" and "high" endurance groups, recovery of tension was found not to be uniform among the two groups. High endurance units were able to recover a greater percentage of their original maximum tetanic tension. No difference was found between force recovery for low and high endurance units at 30 s.(ABSTRACT TRUNCATED AT 400 WORDS)
20
Lambert, R. K., P. D. Paré e C. Y. Seow. "Mathematical description of geometric and kinematic aspects of smooth muscle plasticity and some related morphometrics". Journal of Applied Physiology 96, n. 2 (febbraio 2004): 469–76. http://dx.doi.org/10.1152/japplphysiol.00736.2003.
Testo completoAbstract (sommario):
Despite considerable investigation, the mechanisms underlying the functional properties of smooth muscle are poorly understood. This can be attributed, at least in part, to a lack of knowledge about the structure and organization of the contractile apparatus inside the muscle cell. Recent observations of the plasticity of smooth muscle and of morphometry of the cell have provided enough information for us to propose a quantitative, although highly simplified, model for the geometric arrangement of contractile units and their collective kinematic functions in smooth muscle, particularly airway smooth muscle. We propose that, to a considerable extent, contractile machinery restructures upon activation of the muscle and adapts to cell geometry at the time of activation. We assume that, under steady-state conditions, the geometric arrangement of contractile units and the filaments within these units determines the kinematic characteristics of the muscle. The model successfully predicts the results of experiments on airway smooth muscle plasticity relating to maximal force generation, maximal velocity of shortening, and the variation of compliance with adapted length. The model is also concordant with morphometric observations that show an increase in myosin filament density when muscle is adapted to a longer length. The model provides a framework for design of experiments to quantitatively test various aspects of smooth muscle plasticity in terms of geometric arrangement of contractile units and the muscle's mechanical properties.
21
Munson, J. B., R. C. Foehring, S. A. Lofton, J. E. Zengel e G. W. Sypert. "Plasticity of medial gastrocnemius motor units following cordotomy in the cat". Journal of Neurophysiology 55, n. 4 (1 aprile 1986): 619–34. http://dx.doi.org/10.1152/jn.1986.55.4.619.
Testo completoAbstract (sommario):
Experiments were performed in adult cats to determine the effects of lumbar cordotomy on synaptic potentials, motoneuron membrane electrical properties, muscle-unit contractile properties, and whole-muscle histochemical properties of a heterogeneous skeletal muscle. Medial gastrocnemius (MG) motor units were examined 1 wk to 7 mo following complete transection of the lumbar spinal cord (cordotomy). Motor units were classified on the basis of their contractile properties as type FF, FI, FR, or S (8, 68). Muscle fibers were classified as type FG, FOG, or SO on the basis of histochemical staining (59). Motoneuron electrical properties (axonal conduction velocity, action-potential amplitude, rheobase, input resistance, afterhyperpolarization), group I EPSPs, and muscle-unit contractile properties (unpotentiated and potentiated twitch, unfused and fused tetanus, fatigability) were measured. Reduced numbers of type FR motor units and increased numbers of types FI + FF motor units were found in electrophysiological experiments 2 wk to 7 mo following cordotomy. Corroborative data were obtained from histochemical studies of the same MG muscles. Electrical properties of the motoneurons of each motor-unit type were normal following cordotomy. The close correspondence between motoneuron electrical properties and muscle-unit contractile properties found in normal MG muscle (68) was preserved following cordotomy. Contractile strength of muscle units of all types was severely reduced following cordotomy; partial recovery occurred 4-7 mo following cordotomy. Cross-sectional area of muscle fibers was reduced at all times investigated (2 wk to 7 mo). In three cats, homonymous group Ia single-fiber-motoneuron EPSPs were studied 1 or 2 mo following cordotomy at spinal level L4-5 or L5. EPSP amplitude and afferent-to-motoneuron projection frequency were normal. In 12 other cats, composite heteronymous group I EPSPs were studied 2 wk to 7 mo following cordotomy at various levels. Amplitude of these EPSPs was increased, dependent upon level of cordotomy and postoperative time. Hypotheses concerning the influence of motoneurons on muscle, and of muscle on motoneurons, are presented as possible mechanisms whereby the close relation between motoneuron electrical and muscle-unit contractile properties is preserved in the face of redistributed motor-unit populations.
22
Fine, Michael L., Barbara Bernard e Thomas M. Harris. "Functional morphology of toadfish sonic muscle fibers: relationship to possible fiber division". Canadian Journal of Zoology 71, n. 11 (1 novembre 1993): 2262–74. http://dx.doi.org/10.1139/z93-318.
Testo completoAbstract (sommario):
Sexually dimorphic sonic muscles, which vibrate the swimbladder for sound production in the oyster toadfish (Opsanus tau), are among the fastest vertebrate muscles. Previous work has shown that sonic muscle fibers are smaller in males, have an unusual morphology, and increase in number and size for life. We now report evidence consistent with the hypothesis that mature, presumably postmitotic, sonic fibers divide, and suggest that division, which returns fibers to small energy-efficient units, is necessary because mitochondria are excluded from the fiber's contractile cylinder. Large fibers, potential candidates for division, develop fragments of contractile cylinder separated by channels of an expanded sarcoplasmic reticulum; these channels can assume the appearance of the sarcoplasm (glycogen granules and mitochondria) beneath the sarcolemma. Measurements indicate that contractile cylinder diameter does not increase with fish size and that diameters are approximately 21% larger in females (p < 0.0001). Fiber fragmentation, possible division, and the presence of smaller fibers with smaller diameter contractile cylinders in males are seen as adaptations for repeated rapid contraction and fatigue resistance during production of the male's courtship boatwhistle call.
23
Noble, E. G., e F. P. Pettigrew. "Appearance of "transitional" motor units in overloaded rat skeletal muscle". Journal of Applied Physiology 67, n. 5 (1 novembre 1989): 2049–54. http://dx.doi.org/10.1152/jappl.1989.67.5.2049.
Testo completoAbstract (sommario):
The contractile characteristics of single motor units, isolated from rat plantaris muscles subjected to short-term (30 days) compensatory overload, were assessed to determine whether motor units in transition could be detected. In the control plantaris 88% of the motor units were classified as fast. After overload, a large decline (26.5%) in the proportion of typical fast motor units was noted. The estimated contribution of fast fatigable units to whole muscle tetanic tension (Po 200) also declined (from 55 to 25%), whereas that of fast intermediate motor units increased (from 33 to 55%). In the overloaded plantaris, motor units that exhibited unusual “sag” and contraction time characteristics were detected. These motor units, which could be further subdivided into two distinct types by a variety of indexes, exhibited characteristics intermediate to fast and slow units and therefore were termed “transitional.” Transitional units accounted for 12% of the estimated whole muscle Po200 after overload. These experiments characterize novel classifications of motor units undergoing transformation and further detail the motor unit shift that accompanies compensatory overload.
24
Horcholle-Bossavit, G., L. Jami, J. Petit, R. Vejsada e D. Zytnicki. "Effects of muscle shortening on the responses of cat tendon organs to unfused contractions". Journal of Neurophysiology 59, n. 5 (1 maggio 1988): 1510–23. http://dx.doi.org/10.1152/jn.1988.59.5.1510.
Testo completoAbstract (sommario):
1. The discharges from individual Golgi tendon organs of peroneus tertius and brevis muscles were recorded in anesthetized cats. Responses to unfused isometric contractions of single motor units and combinations of motor units were compared with responses to contractions eliciting muscle shortening (i.e., shortening contractions). 2. In 75% of the examined instances, the effect of muscle shortening during unfused contractions was a slight decrease in tendon organ activation, in keeping with the reduction of contractile tension recorded at the muscle tendon. In other instances there was either no change in tendon organ response or, in less than 10% of instances, a slight increase For two motor units eliciting similar activation of a given tendon organ under isometric conditions, the effect of shortening contraction was not necessarily the same. 3. The reductions observed in tendon organ discharges upon muscle shortening were less than proportional to the reductions of contractile tension and difficult to correlate with the properties of motor units, as determined under isometric conditions. The present observations suggest three main reasons for this lack of relation. 4. The first reason depended on the properties of motor units, in that the relation between length changes and tension changes was not the same for all units. Two motor units developing similar isometric tensions did not necessarily produce the same degree of muscle shortening. Some units produced relatively significant shortening without much loss of tension. 5. Second, the dynamic sensitivity of tendon organs is known to exert a major influence on their responses to isometric unfused contractions, accounting for 1:1 driving of discharge during tension oscillations and high frequency bursts upon abrupt increase of tension. Although less tension was produced and the rate of tension development was slower in shortening contractions, similar manifestations of the dynamic sensitivity of tendon organs were observed. In such cases, the responses of tendon organs were the same whether or not the muscle shortened during contraction. 6. Third, when several motor units were stimulated in combination, the unloading influences of in-parallel units were facilitated by muscle shortening so that unloading effects, which were hardly visible under isometric conditions became evident during shortening contractions.
25
Bigland-Ritchie, Brenda, Andrew J. Fuglevand e Christine K. Thomas. "Contractile Properties of Human Motor Units: Is Man a Gat?" Neuroscientist 4, n. 4 (luglio 1998): 240–49. http://dx.doi.org/10.1177/107385849800400413.
Testo completoAbstract (sommario):
A major goal in neuroscience is to understand how the CNS controls posture and movement in humans. This requires an understanding of individual human motor unit properties and how they interact within the muscle to perform different tasks. This article describes differences and similarities between the contractile properties of human motor units and those of the cat prototype medial gastrocnemius (MG) muscle, on which so many studies have been conducted. The article describes the methods available for measuring human motor unit properties and their limitations, and it discusses how far the behavior of whole muscles can be predicted from their histochemistry. It questions the extent to which human motor units conform to the conventional criteria by which S (slow, fatigue resistant), FR (fast but fatigue resistant) and FF (fast, fatigable) unit types are usually classified. An important difference between human and cat MG data is that weak human motor units are not necessarily slow, nor strong ones fast; that is, generally, human unit force is not correlated with contractile speed. Also, unlike cat MG, the few human muscles studied so far contain few if any FF units but a high proportion of units with intermediate fatigue resistance (Flnt). These apparently aberrant human properties, however, are also found in other cat and rat muscles. Thus, cat MG may not be the best model for motor unit behavior generally. Finally, the influence of human motor unit properties on force output by recruitment and/or rate coding is discussed.
26
Desypris, G., e D. J. Parry. "Relative efficacy of slow and fast alpha-motoneurons to reinnervate mouse soleus muscle". American Journal of Physiology-Cell Physiology 258, n. 1 (1 gennaio 1990): C62—C70. http://dx.doi.org/10.1152/ajpcell.1990.258.1.c62.
Testo completoAbstract (sommario):
Contractile and histochemical properties of reinnervated motor units in soleus muscles of C57BL/6J mice were examined 1 mo after sectioning the soleus nerve. Fifty-one motor units were isolated by the technique of ventral root splitting. Their sizes ranged from 0.4 to 13.6% of whole muscle tetanic tension (Po) with a mean size of 5.3% Po corresponding to 19 motor units. In control unoperated mice, the range was 2.2-8.6% Po, with a mean size of 4.8% Po corresponding to 22 motor units. Although no clear relationship between unit time to peak tension and size was seen in control units, it appeared that in the reinnervated muscle the large units were also slow contracting, whereas the smaller units were predominantly fast contracting. Adenosinetriphosphatase (ATPase) staining revealed an increase in the proportion of muscle area occupied by type I fibers in reinnervated soleus compared with control soleus. Immunohistochemical staining of reinnervated soleus with monoclonal antibodies against type I and IIa myosin showed the presence of hybrid fibers containing both myosins. It is concluded that during reinnervation most motoneurons reinnervate the soleus muscle of the mouse. The hypothesis that slow motoneurons are more adept at expanding their innervating field than fast motoneurons is also supported by the data.
27
Chamberlain, S., e D. M. Lewis. "Contractile characteristics and innervation ratio of rat soleus motor units." Journal of Physiology 412, n. 1 (1 maggio 1989): 1–21. http://dx.doi.org/10.1113/jphysiol.1989.sp017601.
Testo completo
28
Dubose, Laniel, Teresa B. Schelhorn e H. Peter Clamann. "Changes in contractile speed of cat motor units during activity". Muscle & Nerve 10, n. 8 (ottobre 1987): 744–52. http://dx.doi.org/10.1002/mus.880100811.
Testo completo
29
Fenix, Aidan M., Nilay Taneja, Carmen A. Buttler, John Lewis, Schuyler B. Van Engelenburg, Ryoma Ohi e Dylan T. Burnette. "Expansion and concatenation of nonmuscle myosin IIA filaments drive cellular contractile system formation during interphase and mitosis". Molecular Biology of the Cell 27, n. 9 (maggio 2016): 1465–78. http://dx.doi.org/10.1091/mbc.e15-10-0725.
Testo completoAbstract (sommario):
Cell movement and cytokinesis are facilitated by contractile forces generated by the molecular motor, nonmuscle myosin II (NMII). NMII molecules form a filament (NMII-F) through interactions of their C-terminal rod domains, positioning groups of N-terminal motor domains on opposite sides. The NMII motors then bind and pull actin filaments toward the NMII-F, thus driving contraction. Inside of crawling cells, NMIIA-Fs form large macromolecular ensembles (i.e., NMIIA-F stacks), but how this occurs is unknown. Here we show NMIIA-F stacks are formed through two non–mutually exclusive mechanisms: expansion and concatenation. During expansion, NMIIA molecules within the NMIIA-F spread out concurrent with addition of new NMIIA molecules. Concatenation occurs when multiple NMIIA-Fs/NMIIA-F stacks move together and align. We found that NMIIA-F stack formation was regulated by both motor activity and the availability of surrounding actin filaments. Furthermore, our data showed expansion and concatenation also formed the contractile ring in dividing cells. Thus interphase and mitotic cells share similar mechanisms for creating large contractile units, and these are likely to underlie how other myosin II–based contractile systems are assembled.
30
Nelson, A. G., e W. J. Thompson. "Contractile properties and myosin phenotype of single motor units from neonatal rats". American Journal of Physiology-Cell Physiology 266, n. 4 (1 aprile 1994): C919—C924. http://dx.doi.org/10.1152/ajpcell.1994.266.4.c919.
Testo completoAbstract (sommario):
The unloaded shortening velocity (Vus), twitch time to peak force (TTP), and myosin heavy chain (MHC) composition of motor units from soleus muscles of neonatal rats (7-8 and 16-18 days) were compared. The Vus range was from 2.2 to 7.1 fiber lengths (fl).s-1 and was unchanged from 7-8 to 16-18 days. TTP shifted from 7-8 (range, 66-120) to 16-18 days (range, 41-95). MHC-specific antibody staining of motor units revealed a correlation between MHC and Vus of individual motor units. Slowest motor units contained type I MHC. Intermediate Vus motor units contained both embryonic, neonatal, and/or type IIa MHCs. Fastest motor units contained neonatal and/or type IIa MHCs. These findings demonstrate that an individual motor unit of a neonatal muscle contains a nonrandom distribution of fiber types. The range of myosins present within the motor units of the soleus translates into a range of possible Vus.
31
Devasahayam, S. R., e T. G. Sandercock. "Velocity of shortening of single motor units from rat soleus". Journal of Neurophysiology 67, n. 5 (1 maggio 1992): 1133–45. http://dx.doi.org/10.1152/jn.1992.67.5.1133.
Testo completoAbstract (sommario):
1. The force-velocity relationship of a motor unit can provide insight into the contractile proteins of its constituent fibers as well as fundamental information about the function and use of the motor unit. Although the force-velocity profiles of whole muscle and skinned mammalian fibers have been studied, technical difficulties have prevented similar studies on motor units. A technique is presented to directly measure the velocity of shortening of individual motor units from in vivo rat soleus muscle. 2. The soleus muscles of anesthetized rats were dissected free of surrounding tissue while their nerve and blood supplies were preserved. Both tendons were cut, and the distal tendon was attached to a servomechanism to control muscle length, whereas the proximal tendon was attached to a force transducer. Single motor units were stimulated via the ventral roots. 3. The major problem encountered in measuring the force-velocity profile of a motor unit was that the force from the large number of passive fibers and connective tissue in the soleus confounded the force produced by the small number of active fibers in the motor unit. This problem was minimized by measuring active motor unit tension during an isovelocity ramp. This allowed experimental measurement of the passive tension by shortening the muscle with an identical isovelocity ramp without, however, stimulating the motor unit. Active tension was estimated by subtracting the passive tension waveform from the waveform recorded when the motor unit was active. 4. The method substantially reduced the noise from the passive fibers; however, problems remained. The probable sources of error are discussed, with the most significant being the elasticity associated with the blood and nerve connections to surrounding tissue. The elasticity prevents uniform shortening velocities along the length of the active fibers, thereby introducing a systematic bias to measurements made at high velocities. These errors are most pronounced when the data are extrapolated to determine the maximum velocity of shortening (Vmax). Determination of velocity at peak power (Vpp) is a more robust measure; however, of the 34 motor units studied, only 19 exhibited a distinct peak in the power-force curve, indicating residual noise. 5. To assess the validity of using twitch contraction time as an index of the velocity of shortening, when possible, Vmax and Vpp of each motor unit were correlated with the inverse of its twitch contraction time. The correlation was poor (r less than 0.2), indicating that, although widely used, twitch contraction time is a poor index of contractile speed.
32
Meredith, M. A., e S. J. Goldberg. "Contractile differences between muscle units in the medial rectus and lateral rectus muscles in the cat". Journal of Neurophysiology 56, n. 1 (1 luglio 1986): 50–62. http://dx.doi.org/10.1152/jn.1986.56.1.50.
Testo completoAbstract (sommario):
Conjugate eye movements in the horizontal plane are accomplished by the coactivation of the medial rectus (MR) muscle of one orbit and the lateral rectus (LR) muscle of the other. While control of these excursions has been thought to be effected by identical inputs to these muscles, recent studies have demonstrated that MR motoneurons receive different inputs than LR motoneurons. This raises the question of whether the character of the muscles they control are different. The present study evaluated the contractile properties of MR and LR muscle units in the cat. Based on the mechanical aspects of their contractile properties, only two physiological types of muscle units were identified within the MR and LR muscles: twitch and non-twitch muscle units. Twitch muscle units represented over 90% of the units sampled in each muscle. Significant differences in the rate-related and the tension-related contractile properties were demonstrated between MR and LR twitch muscle units. MR muscle units exhibited significantly faster twitch contractions than did LR units. The rate of stimulation at which MR units exhibited fused tetany was significantly higher than for LR units, although units from both muscles demonstrated similar rates of rise of tension at fusion. The rate of rise of tension was closely correlated to tension production (twitch and tetanus) in each muscle. However, MR muscle units demonstrated significantly weaker maximum tetanic tensions and lower tetanus-to-twitch ratios than LR units. These data indicate that while similar physiological types of muscle fibers are present within the MR and LR, MR muscle units are adapted for faster rate-related properties, whereas LR units are adapted for greater tetanic tensions. These distinctions between MR and LR muscle units, coupled with differences between the afferent inputs to their respective motoneurons, suggest that the preservation of conjugacy during horizontal gaze shifts may require a complex interaction of peripheral and central factors.
33
Fuglevand, Andrew J., Vaughan G. Macefield e Brenda Bigland-Ritchie. "Force-Frequency and Fatigue Properties of Motor Units in Muscles That Control Digits of the Human Hand". Journal of Neurophysiology 81, n. 4 (1 aprile 1999): 1718–29. http://dx.doi.org/10.1152/jn.1999.81.4.1718.
Testo completoAbstract (sommario):
Force-frequency and fatigue properties of motor units in muscles that control digits of the human hand. Modulation of motor unit activation rate is a fundamental process by which the mammalian nervous system encodes muscle force. To identify how rate coding of force may change as a consequence of fatigue, intraneural microstimulation of motor axons was used to elicit twitch and force-frequency responses before and after 2 min of intermittent stimulation (40-Hz train for 330 ms, 1 train/s) in single motor units of human long finger flexor muscles and intrinsic hand muscles. Before fatigue, two groups of units could be distinguished based on the stimulus frequency needed to elicit half-maximal force; group 1 ( n = 8) required 9.1 ± 0.5 Hz (means ± SD), and group 2 ( n = 5) required 15.5 ± 1.1 Hz. Twitch contraction times were significantly different between these two groups (group 1 = 66. 5 ms; group 2 = 45.9 ms). Overall 18% of the units were fatigue resistant [fatigue index (FI) > 0.75], 64% had intermediate fatigue sensitivity (0.25 ≤ FI ≤ 0.75), and 18% were fatigable (FI < 0.25). However, fatigability and tetanic force were not significantly different among groups. Therefore unlike findings in some other mammals, fast-contracting motor units were neither stronger nor more susceptible to fatigue than slowly contracting units. Fatigue, however, was found to be greatest in those units that initially exerted the largest forces. Despite significant slowing of contractile responses, fatigue caused the force-frequency relation to become displaced toward higher frequencies (44 ± 41% increase in frequency for half-maximal force). Moreover, the greatest shift in the force-frequency relation occurred among those units exhibiting the largest force loss. A selective deficit in force at low frequencies of stimulation persisted for several minutes after the fatigue task. Overall, these findings suggest that with fatigue higher activation rates must be delivered to motor units to maintain the same relative level of force. Questions regarding classification of motor units and possible mechanisms by which fatigue-related slowing might coexist with a shift in the force-frequency curve toward higher frequencies are discussed.
34
Cope, T. C., C. B. Webb, A. K. Yee e B. R. Botterman. "Nonuniform fatigue characteristics of slow-twitch motor units activated at a fixed percentage of their maximum tetanic tension". Journal of Neurophysiology 66, n. 5 (1 novembre 1991): 1483–92. http://dx.doi.org/10.1152/jn.1991.66.5.1483.
Testo completoAbstract (sommario):
1. The endurance of slow-twitch motor units from the soleus (SOL) and medial gastrocnemius (MG) muscles of the cat were tested by determining the length of time (endurance time, Et) that a unit could maintain its tension output at 85% of maximum. Motor-unit tension was clamped at the target level by altering the stimulation rate of a unit's motor axon through computer feedback control. Tested in this way, units of both muscles displayed a wide range of Ets, approximately 40- to 50-fold. 2. Electromyographic (EMG) waveforms of motor units subjected to force-clamp contractions were analyzed to access whether any activity-dependent changes in their waveform shape might predict Et. Three measurements of waveform shape were determined: baseline-to-baseline duration, peak-to-peak amplitude, and area. Typically, amplitude decreased and duration increased as a contraction proceeded, whereas area remained fairly constant. Because changes in each measure were very similar for units of widely different Ets, it was concluded that neuromuscular junction failure and changes in the excitability of the sarcolemma (excluding the t-tubule system) play a minor role in determining Et. 3. Et was highly correlated with the mean stimulation rate (Et/number of stimuli) used during the force-clamp contractions. Mean rate was seen to progressively decrease with increasing Et. This correlation could not be explained by measures of isometric contractile speed or relaxation (e.g., twitch contraction time or half-relaxation time) measured before the force-clamp contractions. Both contraction time and half-relaxation time were found to be unrelated to both Et and the rate used to stimulate the unit during the force-clamp contraction. 4. Among type S units of SOL and MG, maximum tetanic tension and Et were not related. A significant relation (r = -0.49) was found between axonal conduction velocity and Et for SOL units (n = 38). In addition, a significant correlation (r = 0.47) was found between conduction velocity and tetanic tension for SOL units. Perhaps because of the small sample of type S units from MG (n = 10), conduction velocity was found not be related to either Et or tetanic tension. 5. Others have shown that a motor unit's maximum tetanic tension and axonal conduction velocity are correlated with its order of recruitment among motoneurons innervating a muscle. Recent work has further shown that among type F units the order in which a motoneuron is recruited is highly correlated with the fatigue resistance of its muscle unit.(ABSTRACT TRUNCATED AT 400 WORDS)
35
Petit, J., e M. Gioux. "Properties of motor units after immobilization of cat peroneus longus muscle". Journal of Applied Physiology 74, n. 3 (1 marzo 1993): 1131–39. http://dx.doi.org/10.1152/jappl.1993.74.3.1131.
Testo completoAbstract (sommario):
Changes in contractile properties of cat peroneus longus motor units were studied 2, 5, and 8 wk after selective immobilization of this muscle, which was achieved by fixing the distal tendon of the peroneus longus to the fibula either at the muscle minimal physiological length ("short" length) or at the length for a 90 degree ankle joint ("neutral" length). In each muscle, 75–90% of the units [slow (S), fast resistant to fatigue (FR), fast intermediate (FI), and fast fatigable (FF)] were studied. Immobilization elicited a permanent decrease in tetanic force developed by single motor units, which was larger for resistant-to-fatigue units (S, FR). In most instances this decrease was not related to the immobilization length. In all units, twitch contraction and half-relaxation times underwent a transient increase, the extent and time course of which were influenced by immobilization length. The relationship between the frequency of motor units activation and the ratio of unfused to maximal tetanic force was studied. For fast units, there was a transient shift of the relation toward low frequencies after 2 and 5 wk of immobilization at neutral and short length, respectively.
36
Nelson, A. G. "995 Series compliance varies among motor units of differing contractile speeds". Medicine & Science in Sports & Exercise 25, Supplement (maggio 1993): S177. http://dx.doi.org/10.1249/00005768-199305001-00998.
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Elek, J. M., M. Schubert, K. Wohlfahrt, H. Woldag e R. Dengler. "Contractile properties of motor units in patients with chronic partial denervation". Electroencephalography and Clinical Neurophysiology 87, n. 2 (agosto 1993): S49. http://dx.doi.org/10.1016/0013-4694(93)91055-6.
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Chan, K. Ming, Timothy J. Doherty e William F. Brown. "Contractile properties of human motor units in health, aging, and disease". Muscle & Nerve 24, n. 9 (2001): 1113–33. http://dx.doi.org/10.1002/mus.1123.
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Leitch, Michael, e Vaughan G. Macefield. "Comparison of the contractile responses to irregular and regular trains of stimuli during microstimulation of single human motor axons". Journal of Neurophysiology 111, n. 7 (1 aprile 2014): 1499–506. http://dx.doi.org/10.1152/jn.00835.2013.
Testo completoAbstract (sommario):
During voluntary contractions, human motoneurons discharge with a physiological variability of ∼20%. However, studies that have measured the contractile responses to microstimulation of single motor axons have used regular trains of stimuli with no variability. We tested the hypothesis that irregular (physiological) trains of stimuli produce greater contractile responses than regular (nonphysiological) trains of identical mean frequency but zero variability. High-impedance tungsten microelectrodes were inserted into the common peroneal nerve and guided into fascicles supplying a toe extensor muscle. Selective microstimulation was achieved for 14 single motor axons. Contractile responses were measured via an angular displacement transducer over the relevant toe. After the responses to regular trains of 10 stimuli extending from 2 to 100 Hz were recorded, irregular trains of 10 stimuli, based on the interspike intervals recorded from single motor units during voluntary contractions, were delivered. Finally, the stimulation sequences were repeated following a 2-min period of continuous stimulation at 10 Hz to induce muscle fatigue. Regular trains of stimuli generated a sigmoidal increase in displacement with frequency, whereas irregular trains, emulating the firing of volitionally driven motoneurons, displayed significantly greater responses over the same frequency range (8–24 Hz). This was maintained even in the presence of fatigue. We conclude that physiological discharge variability, which incorporates short and long interspike intervals, offers an advantage to the neuromuscular system by allowing motor units to operate on a higher level of the contraction-frequency curve and taking advantage of catch-like properties in skeletal muscle.
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Doherty, T. J., A. A. Vandervoort, A. W. Taylor e W. F. Brown. "Effects of motor unit losses on strength in older men and women". Journal of Applied Physiology 74, n. 2 (1 febbraio 1993): 868–74. http://dx.doi.org/10.1152/jappl.1993.74.2.868.
Testo completoAbstract (sommario):
The influence of age-associated motor unit loss on contractile strength was investigated in a representative sample of healthy, active young and older men and women. In 24 younger subjects (22–38 yr) and 20 older subjects (60–81 yr) spike-triggered averaging was employed to extract a sample of surface-recorded single motor unit action potentials (S-MUAPs) from the biceps brachii and brachialis muscles. The amplitude of the maximum compound muscle action potential of the biceps brachii and brachialis muscles was divided by the mean S-MUAP amplitude to estimate the numbers of motor units present. The maximum isometric twitch contraction (MTC) and maximum voluntary contraction (MVC) of the elbow flexors were also recorded in 18 of the younger subjects and in all older subjects. The estimated numbers of motor units were significantly reduced (47%, P < 0.001) in older subjects with a mean value of 189 +/- 77 compared with a mean of 357 +/- 97 in younger subjects. The sizes of the S-MUAPs, however, were significantly larger in older subjects (23%, P < 0.01). Significant but less marked age-associated reductions in the MTC (33%, P < 0.05) and MVC (33%, P < 0.001) were also found and were similar for both men and women. These results suggest that motor unit losses, even in healthy active individuals, are a primary factor in the age-associated reductions in contractile strength.
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Macefield, V. G., A. J. Fuglevand e B. Bigland-Ritchie. "Contractile properties of single motor units in human toe extensors assessed by intraneural motor axon stimulation". Journal of Neurophysiology 75, n. 6 (1 giugno 1996): 2509–19. http://dx.doi.org/10.1152/jn.1996.75.6.2509.
Testo completoAbstract (sommario):
1. Single motor axons innervating human toe extensor muscles were selectively stimulated through a tungsten microelectrode inserted percutaneously into the peroneal nerve. Twitch and tetanic forces were measured from a strain gauge over the proximal phalanx of the toe generating the greatest force. Twitch data were obtained from 19 single motor units in nine subjects: 8 motor units supplied extensor hallucis longus (EHL), 5 motor units supplied extensor digitorum longus (EDL), and 6 motor units supplied extensor digitorum brevis (EDB). Unpotentiated twitch forces ranged from 6.3 to 78.1 mN (20.0 +/- 4.0 mN, mean +/- SE), with the distribution highly skewed toward small forces. Twitch contraction and half-relaxation times were 74.8 +/- 3.9 and 78.6 +/- 6.0 ms, respectively. Compared with motor units in human thenar muscles, those in human toe extensor muscles were stronger but slower. However, as in thenar motor units, twitch force and contraction time were not related. 2. Force-frequency relationships were determined for 13 units (5 EDL, 5 EHL, 3 EDB) by stimulating each unit with short trains (1.0-5.0 s) of constant frequency (2-100 Hz). Peak force was related to stimulus frequency in a sigmoid fashion. The steep region of the curve extended from 5.5 +/- 0.7 (SE) Hz to 16.3 +/- 1.1 Hz for all units, and the stimulus frequency required to generate half-maximal force (9.6 +/- 0.6 Hz) was close to the center of the steep range. This frequency, which was inversely related to twitch contraction time, was lower than the frequency required to develop half-maximal force of human thenar motor units (12 +/- 4 Hz, mean +/- SD). The slopes of the regression lines relating force to frequency, computed over the steep range for each unit, were also lower for the toe extensors (3.7 +/- 0.7 mN/Hz) than for the thenar muscles (6 +/- 1 mN/Hz). 3. Maximal tetanic forces ranged from 29.9 to 188.1 mN (89.0 +/- 16.5 mN, mean +/- SE), and were generated at stimulus frequencies from 15 to 100 Hz (median 50 Hz). The stimulation frequency required for fused tetani (absence of noticeable force fluctuation) was generally less than that required for maximum tetanic force. The mean twitch-tetanus ratio, calculated for unpotentiated twitches, was 0.22 +/- 0.02 (range 0.15-0.41). This ratio was higher than for human thenar motor units (0.14 +/- 0.06, mean +/- SE). After twitch potentiation of 10 units, the mean twitch-tetanus ratio increased to 0.28 +/- 0.04. 4. The effects of preceding each stimulus train with a short interstimulus interval (10 ms) on force production at each frequency were examined in nine motor units. Peak forces at the onset of each contraction were higher when such an “initial doublet” preceded stimulus trains of < or = 20 Hz, but the mean force at the end of each stimulus train was not significantly affected at any frequency. 5. Eight units were stimulated with a train that increased in frequency continuously from 2 to 80 Hz, and then decreased symmetrically. This pattern resulted in peak forces that were higher on the descending limb of the stimulus train, the force-frequency relationship tracing a hysteresis loop. Hysteresis was exhibited because damping in the neuromuscular system causes the mechanical output of muscle to lag behind neural input. Thus, in non-steady-state conditions (as in most forms of natural activity), somewhat higher firing rates may be required to attain a particular level of force; once attained, force output will be transiently unresponsive to diminution of firing rate. 6. We conclude that there are differences in the contractile properties of single motor units in human toe extensor muscles (involved in posture and locomotion) and thenar muscles (involved in prehension and manipulation). Twitch-tetanus ratios were greater for motor units in the toe extensors, and this property accounted for the lower force sensitivity of these units to increases in frequency. (ABSTRACT TRUNCATED)
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Cope, T. C., C. B. Webb e B. R. Botterman. "Control of motor-unit tension by rate modulation during sustained contractions in reinnervated cat muscle". Journal of Neurophysiology 65, n. 3 (1 marzo 1991): 648–56. http://dx.doi.org/10.1152/jn.1991.65.3.648.
Testo completoAbstract (sommario):
1. The aim of this study was to describe the control of tension by rate modulation of single motor units in reinnervated muscle. 2. Single fast-twitch motor units were isolated from medial gastrocnemius (MG) muscles in two groups of anesthetized adult cats: one in which the MG nerve was left untreated and another in which that nerve was sectioned and immediately sutured together 10-33 mo before study. Together with conventional measures of isometric contractile properties, units were tested with the use of computer-controlled feedback regulation of stimulation rate to maintain tension during continuous isometric contraction at a constant submaximal level [25% of maximal tension (Pmax)]. 3. For motor units from both groups, stimulation rate began to decline after target tension was attained and then settled at lower values for variable durations before rapidly increasing, usually within the last 5% of the contraction's duration, until reaching the experimentally selected limit of 100 pulses/s (pps). 4. Measures of the declining phase in stimulation rate occurring at the beginning of sustained contraction were not significantly different in comparison of untreated versus reinnervated muscles. These measures included 1) the magnitude of the decrease in rate, 2) the minimum rate attained, and 3) the time taken to reach minimum stimulation rate expressed as a fraction of endurance time (Et, total duration of the sustained contraction). 5. Most fast-twitch units from reinnervated muscles fell within normal limits for both endurance time and the number of stimuli applied over that period.(ABSTRACT TRUNCATED AT 250 WORDS)
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Kuo, Kuo-Hsing, Ana M. Herrera, Lu Wang, Peter D. Paré, Lincoln E. Ford, Newman L. Stephens e Chun Y. Seow. "Structure-function correlation in airway smooth muscle adapted to different lengths". American Journal of Physiology-Cell Physiology 285, n. 2 (agosto 2003): C384—C390. http://dx.doi.org/10.1152/ajpcell.00095.2003.
Testo completoAbstract (sommario):
Airway smooth muscle is able to adapt and maintain a nearly constant maximal force generation over a large length range. This implies that a fixed filament lattice such as that found in striated muscle may not exist in this tissue and that plastic remodeling of its contractile and cytoskeletal filaments may be involved in the process of length adaptation that optimizes contractile filament overlap. Here, we show that isometric force produced by airway smooth muscle is independent of muscle length over a twofold length change; cell cross-sectional area was inversely proportional to cell length, implying that the cell volume was conserved at different lengths; shortening velocity and myosin filament density varied similarly to length change: increased by 69.4% ± 5.7 (SE) and 76.0% ± 9.8, respectively, for a 100% increase in cell length. Muscle power output, ATPase rate, and myosin filament density also have the same dependence on muscle cell length: increased by 35.4% ± 6.7, 34.6% ± 3.4, and 35.6% ± 10.6, respectively, for a 50% increase in cell length. The data can be explained by a model in which additional contractile units containing myosin filaments are formed and placed in series with existing contractile units when the muscle is adapted at a longer length.
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Bishop, Keith N., J. Ross McClung, Stephen J. Goldberg e Mary S. Shall. "Anatomic and physiological characteristics of the ferret lateral rectus muscle and abducens nucleus". Journal of Applied Physiology 103, n. 5 (novembre 2007): 1706–14. http://dx.doi.org/10.1152/japplphysiol.00580.2007.
Testo completoAbstract (sommario):
The ferret has become a popular model for physiological and neurodevelopmental research in the visual system. We believed it important, therefore, to study extraocular whole muscle as well as single motor unit physiology in the ferret. Using extracellular stimulation, 62 individual motor units in the ferret abducens nucleus were evaluated for their contractile characteristics. Of these motor units, 56 innervated the lateral rectus (LR) muscle alone, while 6 were split between the LR and retractor bulbi (RB) muscle slips. In addition to individual motor units, the whole LR muscle was evaluated for twitch, tetanic peak force, and fatigue. The abducens nucleus motor units showed a twitch contraction time of 15.4 ms, a mean twitch tension of 30.2 mg, and an average fusion frequency of 154 Hz. Single-unit fatigue index averaged 0.634. Whole muscle twitch contraction time was 16.7 ms with a mean twitch tension of 3.32 g. The average fatigue index of whole muscle was 0.408. The abducens nucleus was examined with horseradish peroxidase conjugated with the subunit B of cholera toxin histochemistry and found to contain an average of 183 motoneurons. Samples of LR were found to contain an average of 4,687 fibers, indicating an LR innervation ratio of 25.6:1. Compared with cat and squirrel monkeys, the ferret LR motor units contract more slowly yet more powerfully. The functional visual requirements of the ferret may explain these fundamental differences.
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Fitts, R. H., e C. J. Brimmer. "Recovery in skeletal muscle contractile function after prolonged hindlimb immobilization". Journal of Applied Physiology 59, n. 3 (1 settembre 1985): 916–23. http://dx.doi.org/10.1152/jappl.1985.59.3.916.
Testo completoAbstract (sommario):
Contractile properties of slow-twitch soleus (SOL), fast-twitch extensor digitorum longus (EDL), and fast-twitch superficial region of the vastus lateralis were determined in vitro (22 degrees C) in rats remobilized after prolonged (3 mo) hindlimb immobilization (IM). For all muscles the muscle-to-body weight ratio was significantly depressed by IM, and the ratios failed to completely recover even after 90 days. The contractile properties of the fast-twitch muscles were less affected by IM than the slow-twitch SOL. The IM shortened the SOL isometric twitch duration due to a reduced contraction and half-relaxation time. These parameters returned to control levels by the 14th day of recovery. Peak tetanic tension (Po, g/cm2) declined with IM by 46% in the SOL but showed no significant change in the fast-twitch muscles. After IM the SOL Po (g/cm2) recovered to control values by 28 days. The recovery of Po in absolute units (g) was considerably slower and did not return to control levels until 60 (SOL) to 90 (EDL) days. The maximum shortening velocity was not altered by IM in any of the muscles studied. These results demonstrate that both fast- and slow-twitch skeletal muscles possess the ability to completely recover normal contractile function following prolonged periods of hindlimb IM.
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Out, Lia, Tanja G. M. Vrijkotte, Arthur J. van Soest e Maarten F. Bobbert. "Influence of the Parameters of a Human Triceps Surae Muscle Model on the Isometric Torque-Angle Relationship". Journal of Biomechanical Engineering 118, n. 1 (1 febbraio 1996): 17–25. http://dx.doi.org/10.1115/1.2795940.
Testo completoAbstract (sommario):
This study investigates the influence of parameter values of the human triceps surae muscle on the torque-angle relationship. The model used consisted of three units, each containing a contractile, a series elastic and a parallel elastic element. Parameter values were based on morphological characteristics, which made it possible to model individual units. However, for a number of parameters the values reported in the literature vary considerably. It was investigated how sensitive model results were for variation of these parameters. Slack length of the series elastic element, mean moment arm, maximum force, and length of the contractile element appeared to be the most important determinants of the behavior. For mean moment arm and contractile element length, morphology-based methods of estimation could be recommended. Slack length and maximum force were obtained through optimization. It was concluded that the model does not contain parameters on which its output depends strongly and which are difficult to estimate as well, with two exceptions: slack length of the series elastic element and maximum force.
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S., Kwa, Weijs W. e T. van Eijden. "Effects of activation rate on contractile properties of rabbit masseter motor units". Experimental Brain Research 142, n. 2 (1 gennaio 2002): 221–26. http://dx.doi.org/10.1007/s00221-001-0931-6.
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Kwa, S. H. S., W. A. Weijs e T. M. G. J. van Eijden. "Electromyographic and contractile properties of rabbit masseter motor units during fatiguing stimulation". Experimental Brain Research 149, n. 1 (19 dicembre 2002): 96–106. http://dx.doi.org/10.1007/s00221-002-1338-8.
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Elek, J. M., K. Wohlfarth, M. Schubert e R. Dengler. "PS-25-5 Contractile properties of single motor units in Parkinson's disease". Electroencephalography and Clinical Neurophysiology/Electromyography and Motor Control 97, n. 4 (settembre 1995): S146. http://dx.doi.org/10.1016/0924-980x(95)92961-k.
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Abd-El-Basset, Ebtesam M., e Sergey Fedoroff. "Contractile units in stress fibers of fetal human astroglia in tissue culture". Journal of Chemical Neuroanatomy 7, n. 1-2 (luglio 1994): 113–22. http://dx.doi.org/10.1016/0891-0618(94)90012-4.
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