The contractile properties of fast-twitch and slow-twitch skeletal muscles are primarily determined by the myosin isoform content and... Show moreThe contractile properties of fast-twitch and slow-twitch skeletal muscles are primarily determined by the myosin isoform content and modulated by a variety of sarcomere proteins. X-ray diffraction studies of regulatory mechanisms in muscle contraction have focused predominately on fast- or mixed-fiber muscle with slow muscle being much less studied. Here, we used time-resolved x-ray diffraction to investigate the dynamic behavior of the myofilament proteins in relatively pure slow fiber rat soleus (SOL) and pure fast fiber rat extensor digitorum longus (EDL) muscle during twitch and tetanic contractions at optimal lengths (Lo), 95% Lo, and 90% Lo. Before the delivery of stimulation, reduction in muscle length led to decrease in passive tension. The x-ray reflections upon reduction in length showed no transition in the myosin heads from ordered OFF state, where heads are held close to the thick filament backbone, to disordered ON states, where heads are free to bind to thin filament, in both muscles. When stimulation was delivered to both muscles for twitch contractions at Lo, x-ray signatures indicating the transition of myosin heads to ON states were observed in EDL but not in soleus muscle. During tetanic contractions, changes in the disposition of myosin heads as active tension develops is a cooperative process in EDL muscle whereas in soleus muscle this relationship is less cooperative. Moreover, this high cooperativity was maintained in EDL at all lengths tested here, but cooperativity decreased upon reduction in lengths in soleus. The observed reduced extensibility of the thick filaments in soleus muscles as compared to EDL muscles indicate a molecular basis for this behavior. These data indicate that for the EDL thick filament activation is a cooperative strain-induced mechano-sensing mechanism, whereas for the soleus thick filament xiii activation has a more graded response. Lastly, x-ray data collected at different lengths demonstrated that the effect of length on soleus is more pronounce compared to EDL, particularly noticeable in the thick filament during relaxation phase after stimulation ceased. These observations indicate that soleus is more length-dependent than EDL. These different approaches to thick filament regulation in fast- and slow-twitch muscles may be designed to allow for short duration, strong contractions versus sustained finely controlled contractions, respectively. Show less
