Many climbing mammals are able to reverse normal hind foot posture to effect the grip necessary to descend headfirst or to hang upside down. Such hind foot reversal is known in sciurids, procyonids, felids, viverrids, tupaiids, prosimians, and marsupials. The joint movements involved, however, have never been documented unequivocally although various interpretations (some contradictory) have been made. We report here radiographic data from species of the genera Didelphis, Felis, Nasua, Nycticebus, Potos, Sciurus, and Tupaia. In the six eutherians studied, three joints are involved, and there is a common pattern in the mechanism: crurotalar plantarflexion, subtalar inversion, and transverse tarsal supination. Hind foot reversal represents the development of an unusual degree of excursion at these joints, rather than the appearance of any new type of movement. In Didelphis the mechanism is quite different: a bicondylar, spiral tibiotalar joint is the principal site of inversion/abduction movements. This specialization is characteristic of didelphids and phalangerids, and occurs in the extinct multituberculates as well; it is not found in macropodids (which are like eutherians in crurotalar joint structure) or other marsupial families. This diversity in pedal structure and function is evidently the result of parallel evolution from the type of tibiotalar joint of cynodonts and early mammals. In Morganucodon the bulbous, hemispheroidal proximal surface of the talus bears two tibial facets. These facets are represented in didelphids and multituberculates as sulci, whereas in macropodids and eutherians they developed as the proximal and medial surfaces of the talar trochlea. Among living mammals, the primitive hemispheroidal joint is retained among monotremes as a ball and socket joint.
Muscle architecture, moment arms, and locomotor movements in the distal limb segments of the procyonids Nasua (coati) and Procyon (raccoon) are analyzed with reference to patterns of muscle fiber length. This study addresses the hypothesis that relative fiber lengths among muscles in a muscle group can be predicted on the basis of correlates of muscle tension. The results include the following: consistent patterns of fiber length of muscles in a muscle group exist within and between the two genera. Differences in fiber length between muscles can be accounted for by two principal correlates of muscle excursion--length of a muscle's moment arm about a joint and joint-angle excursion. Muscle fiber pinnation permits increased tendon excursion, but this effect is relatively small in comparison to the effects of moment-arm length and joint-angle excursion. Corollary action between two or more joints (or lack thereof) is an important factor in determination of fiber lengths.
Understanding the mechanisms of muscle pattern formation requires that the complete sequence of ontogenetic events be defined, particularly in the emergence of architectural complexity and in the spatial relations between muscles and skeletal elements. This analysis of visceral arch myogenesis in quail (Coturnix coturnix japonica) embryos identifies the location of premuscle condensations and subsequent segregation of individual muscles, documents the initial orientation of myofibers and changes in alignment associated with maturation, and describes the spatial and temporal relations between muscle development and the formation of connective tissues. Premuscle condensations form within the visceral arches on embryonic days 2-4, before skeletal elements make their appearance. Discrete muscles may form from the subdivision of a muscle mass after fiber orientations have been established (e.g., jaw adductor and hyobranchial muscles) or by the segregation of a mesenchymal cluster from the condensation prior to the appearance of oriented fibers (e.g., protractor, muscle of the columella). The rate and pattern of subsequent muscle maturation are closely associated with the development of the hard tissues. Myogenesis in 4-9-day embryos centers around the quadrate cartilage, the retroarticular process of the mandibular (Meckel's) cartilage, and the epibranchial cartilage. Muscles form attachments on these elements and remain without additional attachments until the appropriate elements (e.g., otic capsule, pterygoid bone) develop. No single description of myogenic events applies to all visceral arch muscles, nor is there an arch-specific pattern of ontogeny. Rather, each muscle has distinctive characteristics based on its spatial relations within the developing head.
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