Bibliothek

Notizen: Summary To provide information on the potential stresses imposed on muscle receptors by natural movements, a cinematographic analysis was undertaken of the unrestrained walking, trotting, galloping, jumping and landing movements of 11 adult cats. Appropriate lever arm measurements were also made on 50 freshly displatched cats such that instantaneous representative lengths of soleus, medial gastrocnemius and tibialis anterior could be calculated. Use was made of a previous electromyographic analysis of hind limb muscles during unrestrained locomotion (Engberg, 1964; Engberg and Lundberg, 1969) and consideration given to recordings made by Severin, Shik and Orlovskii of the discharge of dorsal and ventral root filaments supplying ankle extensors during controlled walking of the high decerebrate cat (1967a and b). This approach revealed that locomotor movements involve 3 basic patterns of muscle displacement for the ankle extensors: a large near-static stretch that is alpha-gamma passive (as occurs in crouching); a large dynamic stretch that is also alpha-gamma passive (as occurs in the F phase of the step and after launch-off in the jump); and another dynamic stretch which involves co-activation of alpha and gamma systems and becomes extensive with an increase in forward speed (as occurs in the E2 phase of the step and during landing). There is discussion of the simulation of locomotor type stretch for ankle extensors and the more complex behaviour of tibialis anterior.

Notizen: A cinematographic analysis of the unrestrained walking, trotting, galloping, jumping and landing movements of 11 adult cats was undertaken to provide previously unavailable information concerning the demands imposed on the nervous system for the control of low and high speed movements and the demands imposed by such natural movements on muscle performance and proprioceptive response.With due regard for the swing (F and E1) and stance (E2 and E3) phases of the step cycle of an individual limb, single frame analysis of the film permitted measurement of instantaneous angles of the lower spine, hip, knee, ankle and metatarsophalangeal joints. Appropriate lever arm measurements were also made on 50 freshly dispatched cats and 25 cadavers such that the Law of Cosines could be used to calculate instantaneous lengths of select hind limb muscles that would apply to the natural movements of adult cats of small (1.5-2.5 Kg), intermediate (2.6-3.5 Kg) and large (3.6-4.5 Kg) size. Muscle displacements were analyzed relative to maximum and minimus in situ lengths and the lengths associated with quiet standing. Use was also made of a previous electromyographic analysis of hind limb muscles during unrestrained locomotion (Engberg and Lundberg, '69).The sequential relations between the four phases of the step cycle are maintained as forward speed increases from walking ( 〈 2 mph) to high speed galloping ( 〉 16 mph). There are significant differences in the time consumed by each phase, however, with a greater reduction in the E3 phase, little reduction in the E2 and E1 phases and virtually no reduction in the F phase. When each phase is expressed as a relative percentage of the duration of the total step cycle, the greatest reduction is again in E3 with little change in the E2 phase. In contrast F and E1 phases increase in the percent of time they occur in each cycle, with the greatest increase in the F phase. For all speeds, analysis of the phase relations between movements of various sections of the hind limb revealed a remarkable unity of knee and ankle joint movement. The hip joint is largely out of phase with the knee and ankle during E1 and E2, all three joints being in phase in F and E3. The digits are essentially out of phase with the other joints except in the stance phase of the gallop.Rates and extents of muscle displacement during natural movements are greater than might be anticipated when expressed in absolute mm's and mm/sec but not when considered in relation to maximum and minimum in situ length and the length associated with quiet standing (Ls). During stepping a progressive increase in forward speed results in: (a) a greater usage of muscles at lengths between Ls and maximum in situ length; (b) for knee and ankle extensors, pronounced increase in the lengthening contraction associated with the E2 (yield) phase of step; and, (c) for both flexor and extensor muscles, an increased active phase of lengthening or near isometric contraction immediately prior to periods of active shortening. In contrast to these changes in active muscle status, the change from walking to galloping has little effect on the extent and rate of passive muscle displacements, particularly the F phase stretch of extensors.For the soleus muscle, calculations were made of the relation between changes in overall muscle length during natural movements and the length of the average muscle fiber and the tendon of insertion. These measurements revealed that the increases in fiber length when passive and decreases in length during active shortening are less than would be anticipated from the extensive liteature on extirpated fibers. In contrast, the increase in fiber length when active is greater than would be expected from the admittedly sparse literature on this subject.The results of this study are discussed largely in relation to two points of neurophysiological interest: the physiological range of muscle stretch as it pertains to the responsiveness of muscle spindles and tendon organs; and those mechanical aspects of lengthening contractions that give insight into the neural control of stepping. For exciting both spindles and tendon organs passive muscle stretch and shortening contractions are shown to be relatively ineffective and lengthening and isometric contractions particularly effective movements. It is suggested that, just as recent literature has emphasized the co-activation of efferent alpha and gamma motoneurons as a muscle becomes active, so too is there a synchronous activation of afferents, particularly the Ia and group II endings of muscle spindles and Ib endings of tendon organs. Finally the thesis is advanced that, while it has been convenient to separate E2 from E3 in the description of the stance phase of the step cycle, extensor muscles are actually undergoing a single mechanical event: an active stretch-shorten cycle for knee and ankle extensors and an active isometric-shorten cycle for hip extensors. This hypothesis has significant implications for the neural control program that regulates the stepping sequence in that it emphasizes the extent to which appropriate changes must be preprogrammed in the mechanical properties of muscles for the smooth execution of stepping.

Notizen: The effects of changing muscle length on the mechanical properties of 89 motor units from adult cat medial gastrocnemius have been studied in eight experiments.Few differences were found between the effects of length on tetanic tension, twitch tension, twitch-tetanus ratio, twitch contraction time, twitch half relaxation time, rate of force development and electrical activity for fast contracting (twitch contraction time ≤ 45 msec) and slowly contracting ( 〉 45 msec) units. Those differences that did appear did not persist when these two groups were matched by tetanic tension. It is concluded that the biophysical mechanisms responsible for the changes in mechanical and electrical properties with length must be similar for fast and slow twitch units and not related to potential differences in their muscle fiber type.The effects of changing muscle length on the mechanical properties of the eight whole muscles suggest that changes in force output with length are of minor importance during normal movements as the muscle is found to be electrically active over a relatively narrow range of lengths close to the optimum length for tetanus of the whole muscle. The very shortest muscle lengths at which there is only minimal force development are not used in natural movements, while the declining limb of the length tension curve is at muscle lengths beyond the maximum in situ length.

Notizen: A broad survey of muscle unit properties in 14 muscles of the cat hind limb is presented which emphasizes some general features of unit properties in mammalian muscles. A more detailed analysis of muscle unit properties in three muscles of the posterior compartment of the lower leg is then presented using Burke's tetrapartite (FF, FI or F (Int.), FR, and S) unit classification scheme. Our data on the properties of motor units in cat tibialis posterior (TP) have been compared to those generated by Burke and colleagues on units in flexor digitorum longus (FDL) and medial gastrocnemius (MG). In all three muscles, twitch contraction time was distinctly slower for type S units and specific tension outputs were substantially greater for type FF units than for type S units. The innervation ratios of type FR units were slightly lower than for type S units but the specific tension of the FR units was closer to FF units than to type S units. The FF units controlled 70-74% of the cumulative force output of each muscles, indicating a substantial capacity for powerful rapid contractions of all three of these muscles despite their differences in “size,” action, and force generation. Distinctive features of the three muscles included differences in the unit types' force producing capabilities and in the relative representation of “nonfatigable” type FR and S units in each muscle. In particular, TP is endowed with some unusually powerful type FF units and a high percentage (42%) of type S units. In contrast, FDL has units that develop relatively little force and an unusually high representation (56%) of type FR units. The possible relationships between these muscle features and their presumed role in posture and locomotion is discussed.