Gross anatomy of the hindlimb skeletal system of theGalago senegalensis

The saltatory type of movement of the Prosimian,Galago senegalensis or Lesser Bushbaby is consistent with the gross anatomy of the skeletal system of the hindlimbs. The ilium and foot are elongated as in the tarsius, another prosimian having a similar mode of locomotion. The length of the foot is due largely to an elongated navicular and calcaneus. Other details of the complete bony structure of the hindlimbs are described and compared with animals of the suborders Prosimii and Anthropoidea when such is appropriate.Support for this study was supplied by the Easter Seal Research Foundation of the National Society for Crippled Children and Adults, Inc. and USPHS grant NS08590.     

Comparative anatomy and evolution of the odontocete forelimb

Previous studies of the odontocete forelimb have not considered flipper anatomy in an evolutionary context. This study of 39 cetacean species (1 extinct archaeocete, 31 extant and 3 extinct odontocetes, and 4 mysticetes), provides a detailed comparative analysis of the major bones and muscles of the odontocete flipper. Differences across families in the anatomy of the deltoid, supraspinatus, coracobrachialis, and subscapularis muscles correspond directly to size and shape of forelimb elements. Specialization of the different shoulder girdle muscles allows for more maneuverability of the flipper by independent control of muscular columns. Maximum likelihood analyses helped determine the correlation of characters studied by ancestral state reconstruction, and revealed independent evolution of osteological and external characters among various lineages. Comparative Analyses by Independent Contrast (CAIC), found several contrasts between flipper area and body length for several extant odontocetes and a linear relationship was inferred. Degree of hyperphalangy and the soft tissue encasing the flipper helped determine three flipper morphologies based on aspect ratio (AR) and qualitative data. These results suggest that differences in flipper shape have an evolutionary component and are likely largely in response to ecological requirements.     

Functional analysis of the shoulder girdle of cats during locomotion

The movements of the shoulder girdle of eight adult cats during overground stepping were studied, using standard slow motion cinematographic techniques. The patterns of activity of shoulder muscles were examined, using simultaneous intramuscular electromyography. Walking, trotting and galloping steps were analyzed from digitized single motion picture frame images. Angular movements of the shoulder girdle consist of biphasic flexion and extension of the shoulder joint and a monophasic flexion-extension alternation of the scapula on the thorax during each step cycle. In addition, the center of the scapula moves craniad during the swing phase and caudad during the stance phase with respect to a fixed reference point on the animal. Similar vertical movements of the center of the scapula also occur in each step cycle. Results of EMG studies of the 17 muscles capable of acting on the shoulder girdle indicate that three overall patterns of activity are found: (1) a pattern typical of extensor muscles, active during all the extension epochs; (2) a pattern typical of flexor muscles, active during the flexion epoch; and (3) a biphasic pattern of activity, active twice in each step. There data are used, along with a re-examination of previous models of the mechanics of the shoulder girdle of carnivores to examine the function and mechanics of shoulder motion. It is concluded that the rotary and translatory movements of the shoulder girdle during stepping combine to enhance step length.     

Integration of the mammalian shoulder girdle within populations and over evolutionary time

Morphological integration has the potential to link morphological variation within populations with morphological evolution among species. This study begins to investigate this link by comparing integration among shoulder girdle elements (e.g. scapular blade, glenoid, coracoid, etc.) during the origin and evolution of therian mammals, and within modern bat, opossum and mouse populations. In this study, correlations among skeletal elements and patterns of allometry are used as proxies for integration. Results suggest that shoulder girdle elements tended to vary and evolve independently during the origin of mammals and subsequent radiation of placentals, consistent with the elements’ distinct developmental and evolutionary origins. This finding suggests that skeletal element correlations, and therefore integration, can be conserved over large taxonomic and temporal scales. However, marsupials display a different pattern in which shoulder girdle elements tend to be more integrated, with the exception of the coracoid. This finding is consistent with a shift in the pattern of skeletal element integration coincident with the appearance of the marsupial mode of reproduction. This finding provides further evidence that development can play a significant role in the establishment of patterns of skeletal element correlation and that patterns of skeletal element correlation can themselves evolve when faced with sufficient selective pressures.     

Lessons learnt from the painful shoulder; a case series of malignant shoulder girdle tumours misdiagnosed as frozen shoulder

Adhesive capsulitis or frozen shoulder is a common condition characterized by shoulder pain and stiffness. In patients in whom conservative measures have failed, more invasive interventions such as arthrographic or arthroscopic distension can be very effective in relieving symptoms and improving range of movement. However, absolute contraindications to these procedures include the presence of neoplasia around the shoulder girdle. We present five cases referred to our institution where the diagnosis of shoulder joint malignancy was delayed, following prolonged, ineffective treatment for frozen shoulder. These cases highlight the importance of careful review of the radiology and the need for reconsideration of the diagnosis in refractory "frozen shoulder".     

Metastatic transitional cell carcinoma of the urinary bladder to the shoulder girdle

Transitional cell carcinoma (TCC) of the bladder typically metastasizes to the pelvic lymph nodes and to visceral sites including the lungs, liver, and bones. Other sites include the brain, especially after systemic chemotherapy. To our knowledge, we report the first case of TCC metastatic to the soft tissue of the shoulder girdle and discuss common and unusual sites of metastasis in TCC.     

Genetics of shoulder girdle formation: roles of Tbx15 and aristaless-like genes

The diverse cellular contributions to the skeletal elements of the vertebrate shoulder and pelvic girdles during embryonic development complicate the study of their patterning. Research in avian embryos has recently clarified part of the embryological basis of shoulder formation. Although dermomyotomal cells provide the progenitors of the scapular blade, local signals appear to have an essential guiding role in this process. These signals differ from those that are known to pattern the more distal appendicular skeleton. We have studied the impact of Tbx15, Gli3, Alx4 and related genes on formation of the skeletal elements of the mouse shoulder and pelvic girdles. We observed severe reduction of the scapula in double and triple mutants of these genes. Analyses of a range of complex genotypes revealed aspects of their genetic relationship, as well as functions that had been previously masked due to functional redundancy. Tbx15 and Gli3 appear to have synergistic functions in formation of the scapular blade. Scapular truncation in triple mutants of Tbx15, Alx4 and Cart1 indicates essential functions for Alx4 and Cart1 in the anterior part of the scapula, as opposed to Gli3 function being linked to the posterior part. Especially in Alx4/Cart1 mutants, the expression of markers such as Pax1, Pax3 and Scleraxis is altered prior to stages when anatomical aberrations are visible in the shoulder region. This suggests a disorganization of the proximal limb bud and adjacent flank mesoderm, and is likely to reflect the disruption of a mechanism providing positional cues to guide progenitor cells to their destination in the pectoral girdle.     

The anatomy of the forelimb in the anteater (Tamandua) and its functional implications

The forelimbs of anteaters play a major role in obtainment of food, defense, and locomotion. The greatly enlarged claws on the manus are used for ripping open insect nests and insect-infested wood; the claws also serve as the animals' only defensive weapons, since they lack teeth. Specialization of the claws for these functions has also had a substantial effect on the ways in which the forelimb is used for posture and locomotion. Modifications of the forelimb in the anteater Tamandua include the following. Attachments of the medial head of triceps are rearranged so as to greatly increase capability for powerful flexion of the claws. Ability to flex the elbow and to retract the humerus is also augmented; these movements would assist digital flexion in applying traction with the claws to material being torn away during food procurement. This traction can be supplemented by a variety of powerful side-to-side and/or twisting movements of the hand, brought about primarily by axial rotation of the upper arm and forearm. The digital joints are reinforced to resist the deviational and torsional loading to which the digits would be subjected during such movements. The morphological modifications of the forelimb in Tamandua are discussed in terms of how they affect the mechanical capabilities of the limb, what functions the limb is best designed to perform, how they may relate to what little is known about the specialized behavior of this animal, and what behavioral predictions may be made based on mechanical design.     

Similarity,parsimony and conjectures of homology: The chelonian shoulder girdle revisited

Much has been written about the definition and recognition of biological homology. Homology is usually defined as similarity inherited from a common ancestor (e.g., papers in Hall, 1994). It is recognised through cladistic analysis: Patterson (1982) and de Pinna (1991) have cogently argued that homology can be equated with synapomorphy (a shared evolutionary novelty uniting a monophyletic group). Such identification involves two stages: first, a possible homology is proposed on the basis of morphological similarity. This similarity might be structural, topological, developmental, or any combination thereof. Next, a cladistic analysis is performed, involving the trait in question and all other informative traits identified. If the trait is congruent with the resultant phylogeny, it is accepted as homologous in all taxa which possess it. If the trait is incongruent with the phylogeny, it is interpreted as homoplasious in certain taxa. This has been termed the test of congruence (Patterson, 1982; de Pinna, 1991). Rieppel (1996) has recently suggested that the test of congruence might be circular, and that as a result certain inferences about the evolution of the chelonian shoulder girdle (Lee, 1996) are poorly substantiated. Here I argue that the test of congruence is not circular, and that the disputed conclusions about the evolution of chelonian shoulder girdle can be defended on the basis of parsimony. More generally, I suggest how considerations of parsimony can and should be used to arbitrate between conflicting conjectures of homology that are both congruent with an accepted phylogeny.