There is increasing evidence that the architectural design and arrangement of the fibers within a motor unit have important physiological and developmental ramifications. Limited data, however, are available to directly address this issue. In the present study the physiological properties of one motor unit in each of seven cat tibialis anterior (TA) muscles were determined. Each of these units then was repetitively stimulated to deplete the glycogen in all muscle fibers within the unit. Subsequently, the length, type of ending, and spatial distribution of fibers sampled from these physiologically and histochemically typed motor units were determined. Four fast fatigable (FF), one fast, fatigue resistant (FR), and two slow (S) motor units (MU) were studied. The samples consisted of all those glycogen-depleted fibers (9-27) contained within a single fascicle or a circumscribed area of each of the motor unit territories. The mean fiber lengths for the two slow motor units were 35.9 and 45.5 mm. The mean fiber lengths for the fast motor unit samples ranged from 8.8 to 48.5 mm. Some fibers of both the fast and slow units reached lengths of 58 mm. Most of the fibers in the slow units extended the entire distance between the proximal and distal musculotendinous planes, had relatively constant cross-sectional areas, and terminated at the tendon as blunt endings. In contrast, the majority of the fibers in the fast units terminated intrafascicularly at one end, and the cross-sectional area decreased progressively along their lengths, that is, showed a tapering pattern for a significant proportion of their lengths. Therefore, the force generated by units that end midfascicularly would appear to be transmitted to connective tissue elements and/or adjacent fibers. All fibers of a fast unit within a fascicle were located at approximately the same proximo-distal location. Thus, developmentally the selection of muscle fibers by a motoneuron would seem to be influenced by their spatial distribution. The architectural complexities of motor units also have clear implications for the mechanical interactions of active and inactive motor units. For example, the tension capabilities of a motor unit may be influenced not only by the spatial arrangement of its own fibers, but also by the level of activation of neighboring motor units.
The motor unit is the basic unit for force production in a muscle. However, the position and shape of the territory of a motor unit within the muscle have not been defined precisely. The territories of five motor units in the cat tibialis anterior muscle were reconstructed three-dimensionally (3-D) from tracings of the glycogen-depleted fibers belonging to each unit. The motor unit territories did not span the entire length of the muscle and their cross-sectional areas tapered along the proximodistal axis producing a conical shape. In addition, the position of the territory of each unit shifted in an anterior-posterior plane along the longitudinal axis of the muscle, presumably as a consequence of the pinnation of the fibers. The area of the motor unit territory at any given level along the proximodistal axis was highly correlated with the number of fibers within the territory at that level. Connective tissue boundaries (outlining fascicles) appeared to have a strong influence on the shape of the territory, territories showed abrupt changes at connective tissue boundaries as groups of motor unit fibers within a fascicle often terminated together while motor unit fibers in neighboring fascicles did not terminate. It is likely that the mechanical impact of the recruitment of a motor unit is affected by the location and shape of motor units within the same muscle area.(ABSTRACT TRUNCATED AT 250 WORDS)
The geometric shape of the filamentous, intrafascicular type of muscle fiber ending was reconstructed as a basis for understanding the pattern in relay of the fiber's force to the muscle tendon. Single motor units (MUs) identified physiologically as being fast and slow, respectively, were isolated in cat tibialis muscles and glycogen-depleted for recognition in cross sections of the muscle frozen at its Lo. Serial measurements of cross-sectional area (CSA) using an image processing system were made along 14 intrafascicular endings of MU fibers and an additional seven, non-depleted fibers identified histochemically as slow. Comparison of coefficients of variation for the linear relation of the CSAs and of the equivalent diameters with length along the taper indicated that in both fast and slow fibers the areas bore a closer relationship, that is, the taper had the equivalent of a parabolic, rather than a conical outline. The implications of these two conformations to relay of the fiber's contractile force to surrounding structures are displayed graphically.
The physiological cross-sectional area (CSA) of a motor unit (MU), taken as the sum of fiber areas measured on a single section through the approximate midlength of the MU, has been compared with the physiological CSA more strictly defined as the sum of the maximal areas to be found anywhere along the length of each of the MU fibers. The CSA at intervals along the fiber length was measured in fibers selected from four glycogen-depleted, isolated MUs in the cat tibialis anterior (TA), and profiles of the summed areas made. In one MU, measurements were also taken on all the MU's fibers at less frequent intervals. The profiles demonstrate that the summed CSA based on each fiber's maximum CSA may exceed that derived from observation on any single section by as much as 20%. As a consequence, values that have been reported for specific tension (force per unit area) of MUs in the TA and probably other muscles may have been overestimated, especially for those MUs of fast type. Estimates were also made of the share of the MU's total force transmitted directly to the tendons of origin and insertion via endings of the blunt musculotendinous type as distinct from tapering intrafascicular endings acting through in-series connective tissue and non-MU fibers. In two MUs of slow type in which most fibers ran from tendon to tendon, "partial tapering" extending over 1 cm of the fiber length accounted for a third of the total physiological CSA, and indicated yet another mode for relay of the MU's force to the tendon.
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