Myotube heterogeneity in developing chick craniofacial skeletal muscles.

Avian skeletal muscles consist of myotubes that can be categorized according to contraction and fatigue properties, which are based largely on the types of myosins and metabolic enzymes present in the cells. Most mature muscles in the head are mixed, but they display a variety of ratios and distributions of fast and slow muscle cells. We examine the development of all head muscles in chick and quail embryos, using immunohistochemical assays that distinguish between fast and slow myosin heavy chain (MyHC) isoforms. Some muscles exhibit the mature spatial organization from the onset of primary myotube differentiation (e.g., jaw adductor complex). Many other muscles undergo substantial transformation during the transition from primary to secondary myogenesis, becoming mixed after having started as exclusively slow (e.g., oculorotatory, neck muscles) or fast (e.g., mandibular depressor) myotube populations. A few muscles are comprised exclusively of fast myotubes throughout their development and in the adult (e.g., the quail quadratus and pyramidalis muscles, chick stylohyoideus muscles). Most developing quail and chick head muscles exhibit identical fiber type composition; exceptions include the genioglossal (chick: initially slow, quail: mixed), quadratus and pyramidalis (chick: mixed, quail: fast), and stylohyoid (chick: fast, quail: mixed). The great diversity of spatial and temporal scenarios during myogenesis of head muscles exceeds that observed in the limbs and trunk, and these observations, coupled with the results of precursor mapping studies, make it unlikely that a lineage based model, in which individual myoblasts are restricted to fast or slow fates, is in operation. More likely, spatiotemporal patterning of muscle fiber types is coupled with the interactions that direct the movements of muscle precursors and subsequent segregation of individual muscles from common myogenic condensations. In the head, most of these events are facilitated by connective tissue precursors derived from the neural crest. Whether these influences act upon uncommitted, or biased but not restricted, myogenic mesenchymal cells remains to be tested.