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.