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The acquisition of a movable jaw and a jaw joint are key events in gnathostome evolution. Jaws are derived from the neural crest derived pharyngeal skeleton and the transition from jawless to jawed vertebrates consists of major morphological changes, which must have a genetic foundation. Recent studies on the effects of bapx1 knockdown in fish and chicken indicate that bapx1 has acquired such a role in primary jaw joint development during vertebrate evolution, but evidence from amphibians is missing so far. In the present study, we use Ambystoma mexicanum, Bombina orientalis, and Xenopus laevis to investigate the effects of bapx1 knockdown on the development of these three different amphibians. Using morpholinos we downregulated the expression of bapx1 and obtain morphants with altered mandibular arch morphology. In the absence of bapx1 Meckeĺs cartilage and the palatoquadrate jaw joint initially develop separately but during further development the joint cavity between both fills with chondrocytes. This results in the fusion of both cartilages and the loss of the jaw joint. Despite this the jaw itself remains usable for feeding and breathing. We show that bapx1 plays a role in jaw joint maintenance during development and that the morphants morphology possibly mirrors the morphology of the jawless ancestors of the gnathostomes.

The vertebrate head/trunk interface is the region of the body where the different developmental programs of the head and trunk come in contact. Many anatomical structures that develop in this transition zone differ from similar structures in the head or the trunk. This is best exemplified by the cucullaris/trapezius muscle, spanning the head/trunk interface by connecting the head to the pectoral girdle. The source of this muscle has been claimed to be either the unsegmented head mesoderm or the somites of the trunk. However most recent data on the development of the cucullaris muscle are derived from tetrapods and information from actinopterygian taxa is scarce. We used classical histology in combination with fluorescent whole-mount antibody staining and micro-computed tomography to investigate the developmental pattern of the cucullaris and the branchial muscles in a basal actinopterygian, the Longnose gar (Lepisosteus osseus). Our results show (1) that the cucullaris has been misidentified in earlier studies on its development in Lepisosteus. (2) Cucullaris development is delayed compared to other head and trunk muscles. (3) This developmental pattern of the cucullaris is similar to that reported from some tetrapod taxa. (4) That the retractor dorsalis muscle of L. osseus shows a delayed developmental pattern similar to the cucullaris. Our data are in agreement with an explanatory scenario for the cucullaris development in tetrapods, suggesting that these mechanisms are conserved throughout the Osteichthyes. Furthermore the developmental pattern of the retractor dorsalis, also spanning the head/trunk interface, seems to be controlled by similar mechanisms.

The pectoral girdle is a unique skeletal element that underwent drastic morphological changes during its evolution, especially in association with the fin-to-limb transition. Comparative studies of its development are needed to gain a deeper understanding of its evolution. Transplantation experiments using the quail-chick chimeric system have revealed that not only lateral plate mesoderm but also somites contribute to the pectoral girdle in birds. Studies in mice and turtles also document somitic contributions to the pectoral girdle, but extirpation experiments in a salamander did not affect shoulder girdle development. Somitic contributions to the pectoral girdle therefore have been interpreted as a feature unique to amniotes. Here, we present a long-term fate map of single somites in the Mexican axolotl, based on transplantations of somites two to six from GFP-transgenic donors into wild-type hosts, as well as injections of fluorescein dextran into single somites. The results show a somitic derivation of the dorsal region of the suprascapula, demonstrating that somitic contributions to the pectoral girdle are not restricted to amniotes. Comparison with the few other species studied so far leads us to suggest a position-dependent origin of the pectoral girdle. We propose that embryonic origin is determined by the proximity of the developing pectoral girdle to the somites or to the lateral plate mesoderm, respectively. This position-dependent origin and the diversity of the anatomy of the pectoral girdle among vertebrates implies that the embryonic origin of the pectoral girdle is too variable to be useful for defining homologies or for phylogenetic analysis.

The fate of single somites has not been analyzed from a comparative perspective with modern cell-marking methods. Most of what we know is based on work using quail-chick chimeras. Consequently, to what degree cell fate has been conserved despite the anatomical differences among vertebrates is unknown. We have analyzed the cell fate of the cranialmost somites, with the focus on somite two, in the Mexican axolotl (Ambystoma mexicanum). Somite cells were marked by injection of dextran-fluorescein and detected using immunofluorescence after 2 months of development in paraffin sections. Our data confirm and extend earlier studies based on classical histology in salamanders. We show that somite two contributes to different muscles, skeletal elements, and connective tissues of the head and cranial trunk region. Cells from somites two and three migrate latero-ventrally and contribute to the hypobranchial muscles mm. geniohyoideus and rectus cervicis. We provide evidence that the specific formation of the hypobranchial musculature from ventral processes of the somites might be variable in different classes of vertebrates. We further demonstrate that mm. cucullaris and dilatator laryngis, which were earlier thought to have a branchial origin, arise from somitic material in a manner very similar to the findings in quail-chick chimeras. Our findings indicate that the pattern of somitic derivatives is highly conserved within tetrapods.