Selectively reshaping a muscle phenotype: functional overload of cat plantaris.

INTRODUCTION: Functional overload (FO) of the fast plantaris muscle was studied in treadmill-exercised (FO-Ex) or sedentary (FO-Sed) adult cats. METHODS: Mechanical, phenotype, and kinematics analyses were performed. RESULTS: Plantigrade vs. normal digitigrade posture was observed early post-FO. Relative plantaris mass was greater in FO-Sed (10%) and FO-Ex (60%) cats than in controls 12 weeks post-FO. Specific tension was similar across groups, indicating functional hypertrophy. Fiber size was greater, percent slow fibers higher, percent IIa myosin heavy chain (MHC) higher, and IIx MHC lower in FO-Ex than controls. Twitch and half-relaxation times were longer, and the frequency-tension curve shifted toward that observed in slow muscles. Electromyography (EMG) and tendon force amplitudes during stepping were larger, and the yield (lengthening) phase occurred at a longer muscle length before compared with after FO. DISCUSSION: Reshaping the plantaris phenotype was highly dependent on the overload stimulus, indicating that electrical stimulation paradigms used during rehabilitation should be performed with the muscles under "loaded" conditions.

Does daily activity level determine muscle phenotype?

The activation level of a muscle is presumed to be a major determinant of many mechanical and phenotypic properties of its muscle fibers. However, the relationship between the daily activation levels of a muscle and these properties has not been well defined, largely because of the lack of accurate and sustained assessments of the spontaneous activity levels of the muscle. Therefore, we determined the daily activity levels of selected rat hindlimb muscles using intramuscular EMG recordings. To allow comparisons across muscles having varying activity levels and/or muscle fiber type compositions, we recorded EMG activity in a predominantly slow plantarflexor (soleus), a predominantly fast plantarflexor (medial gastrocnemius, MG), a predominantly fast ankle dorsiflexor (tibialis anterior, TA) and a predominantly fast knee extensor (vastus lateralis, VL) in six unanesthetized rats for periods of 24 h. EMG activity levels were correlated with the light:dark cycle, with peak activity levels occurring during the dark period. The soleus was the most active and the TA the least active muscle in all rats. Daily EMG durations were highest for soleus (11-15 h), intermediate for MG (5-9 h) and VL (3-14 h) and lowest for TA (2-3 h). Daily mean EMG amplitudes and integrated EMG levels in the soleus were two- to threefold higher than in the MG and VL and seven- to eightfold higher than in the TA. Despite the three- to fourfold difference in activation levels of the MG and VL vs the TA, all three predominantly fast muscles have been reported to have a similar, very low percentage of slow fibers. Comparing these relative EMG levels to the published fiber type profiles of these muscles yields a very poor relationship between daily activity level and fiber type composition in the same muscles across several species. Although it is clear that changing the levels of activity can modulate the expression of the myosin phenotype, these results indicate that factors other than activation must play critical roles in determining and maintaining normal phenotypic properties of skeletal muscle fibers.

Mechanical properties of rat soleus aponeurosis and tendon during variable recruitment in situ.

The in vitro mechanical properties of tendons are well described, whereas little data exist for conditions mimicking those found in vivo. Descriptions of the in situ mechanical properties of aponeuroses are more common, but the results are variable. Our goal was to examine the mechanical properties of these tissues under conditions mimicking the in vivo state. Tissue strains were measured in the rat (Rattus norvegicus) soleus muscle directly from the spacing of metal markers implanted within the tissues of interest using an X-ray video microscope. Strains were measured for the tendon and three regions (proximal, middle and distal) of the aponeurosis. Muscle stimulation was accomplished through isolated ventral rootlets, allowing force to be graded in seven repeatable increments independent of muscle-tendon unit length. Peak strains (during maximal tetanic contraction at optimum length; P(o)) were approximately 5% in tendon and approximately 12% in all regions of the aponeurosis. At forces above 50% of P(o), tissue stiffness was nearly constant in all regions, and a pronounced toe region was observed only at forces below approximately 25% of P(o). Stiffness increased in all regions as the muscle-tendon unit was lengthened. These results suggest that using mechanical properties measured ex vivo or during single contractile events in situ to estimate the in vivo behavior of tendon and aponeurosis may lead to errors in estimating the distribution of strain among the contractile and series elastic elements of the muscle.

Adaptations in skeletal muscle disuse or decreased-use atrophy.

Those factors that seem to play some role in inducing adaptations of skeletal muscle in vivo are discussed. The role of myogenesis in maintaining and repairing muscle during atrophic and hypertrophic states is discussed, including pointing out that the modulation of myonuclear number is one means of adapting to varying chronic levels of neuromuscular activity. Finally, we point out the potential consequences of muscle atrophy on the control of movement and the susceptibility to fatigue.

Mechanical properties of the electrically silent adult rat soleus muscle.

The isometric and isotonic in situ mechanical properties of the soleus muscle of adult female rats were determined after 60 days of inactivity induced by spinal cord isolation (SI). Compared to control, the absolute muscle mass, physiological cross-sectional area, and maximum tetanic tension of the soleus in SI rats were reduced by 69%, 66%, and 77%, respectively. Isometric twitch time-to-peak-tension and half-relaxation times were 41% and 60% shorter in SI than control rats. The maximum velocity of shortening (mm/s), as determined using the afterloaded technique, was 66% faster in SI than control rats, whereas unloaded shortening velocity was similar in the two groups (9% faster in SI rats). Peak power was 48% lower in SI than control rats. The SI soleus was 39% more fatigable than control. Thus, the soleus became a smaller, faster, and more fatigable muscle following 60 days of inactivity. In general, the results indicate that the adaptations are of a lesser magnitude than those reported previously following denervation for the same duration. These data provide a baseline for future efforts to experimentally define the mechanisms of neurally mediated, but activity-independent, regulation of the mechanical properties of the rat soleus muscle.