Posterior tail development in the salamander Eurycea cirrigera: exploring cellular dynamics across life stages.

During embryogenesis, the body axis elongates and specializes. In vertebrate groups such as salamanders and lizards, elongation of the posterior body axis (tail) continues throughout life. This phenomenon of post-embryonic tail elongation via addition of vertebrae has remained largely unexplored, and little is known about the underlying developmental mechanisms that promote vertebral addition. Our research investigated tail elongation across life stages in a non-model salamander species, Eurycea cirrigera (Plethodontidae). Post-embryonic addition of segments suggests that the tail tip retains some aspects of embryonic cell/tissue organization and gene expression throughout the life cycle. We describe cell and tissue differentiation and segmentation of the posterior tail using serial histology and expression of the axial tissue markers, MF-20 and Pax6. Embryonic expression patterns of HoxA13 and C13 are shown with in situ hybridization. Tissue sections reveal that the posterior spinal cord forms via cavitation and precedes development of the underlying cartilaginous rod after embryogenesis. Post-embryonic tail elongation occurs in the absence of somites and mesenchymal cells lateral to the midline express MF-20. Pax6 expression was observed only in the spinal cord and some mesenchymal cells of adult Eurycea tails. Distinct temporal and spatial patterns of posterior Hox13 gene expression were observed throughout embryogenesis. Overall, important insights to cell organization, differentiation, and posterior Hox gene expression may be gained from this work. We suggest that further work on gene expression in the elongating adult tail could shed light on mechanisms that link continual axial elongation with regeneration.

Evolving possibilities: Post-embryonic axial elongation in salamanders with biphasic (Eurcyea cirrigera, E. longicauda, E. quadridigitata) and paedomorphic life cycles (Eurycea nana and Ambystoma mexicanum). Submitted Acta Zoologica.

Typically the number of vertebrae an organism will have post-embryonically is determined during embryogenesis via the development of paired somites. Our research investigates the phenomenon of post-embryonic vertebral addition in salamander tails. We describe body and tail growth, and patterns of postsacral vertebral addition and elongation in context with caudal morphology for four plethodontids (Eurycea), and one ambystomatid. Eurycea nana and A. mexicanum have paedomorphic life cycles; E. cirrigera, E. longicauda and E. quadridigitata are biphasic. Specimens were collected, borrowed and/or purchased, and cleared and stained for bone and cartilage. Data collected include snout-vent length (SVL), tail length (TL), vertebral counts and centrum lengths. Eurycea species with biphasic life cycles had TLs that surpassed SVL following metamorphosis. Tails in paedomorphic species elongated but rarely exceeded body length. Larger TLs were associated with more vertebrae and longer vertebrae in all species. We observed that rates of postsacral vertebral addition varied little amongst species. Regional variation along the tail becomes prominent following metamorphosis in biphasic developers. In all species vertebrae in the posterior one-half of the tail taper towards the tip. We suggest a developmental link might exist between the ability to continually add vertebrae and regeneration in salamanders.

Early differentiation and migration of cranial neural crest in the opossum, Monodelphis domestica.

Marsupial mammals are born at a highly altricial state. Nonetheless, the neonate must be capable of considerable functional independence. Comparative studies have shown that in marsupials the morphogenesis of many structures critical to independent function are advanced relative to overall development. Many skeletal and muscular elements in the facial region show particular heterochrony. Because neural crest cells are crucial to forming and patterning much of the face, this study investigates whether the timing of cranial neural crest differentiation is also advanced. Histology and scanning electron microscopy of Monodelphis domestica embryos show that many aspects of cranial neural crest differentiation and migration are conserved in marsupials. For example, as in other vertebrates, cranial neural crest differentiates at the neural ectoderm/epidermal boundary and migrates as three major streams. However, when compared with other vertebrates, a number of timing differences exist. The onset of cranial neural crest migration is early relative to both neural tube development and somite formation in Monodelphis. First arch neural crest cell migration is particularly advanced and begins before any somites appear or regional differentiation exists in the neural tube. Our study provides the first published description of cranial neural crest differentiation and migration in marsupials and offers insight into how shifts in early developmental processes can lead to morphological change.

Tail development and regeneration throughout the life cycle of the four-toed salamander Hemidactylium scutatum.

The salamander tail displays different functions and morphologies in the aquatic and terrestrial stages of species with complex life cycles. During metamorphosis the function of the tail changes; the larval tail functions in aquatic locomotion while the juvenile and adult tail exhibits tail autotomy and fat storage functions. Because tail injury is common in the aquatic environment, we hypothesized that mechanisms have evolved to prevent altered larval tail morphology from affecting normal juvenile tail morphology. The hypothesis that injury to the larval tail would not affect juvenile tail morphology was investigated by comparing tail development and regeneration in Hemidactylium scutatum (Caudata: Plethodontidae). The experimental design included larvae with uninjured tails and with cut tails to simulate natural predation. The morphological variables analyzed to compare normally developing and regenerating tails were 1) tail length, 2) number of caudal vertebrae, and 3) vertebral centrum length. Control and experimental groups do not differ in time to metamorphosis or snout-vent length. Tails of experimental individuals are shorter than controls, yet they display a significantly higher rate of tail growth and less resorption of tail tissue. Anterior to the site of tail injury, caudal vertebrae in juveniles display greater average centrum lengths. Results suggest that regenerative mechanisms are functioning not only to produce structures, but also to influence growth of existing structures. Further investigation of juvenile and adult stages as well as comparative analyses of tail morphology in salamanders with complex life cycles will enhance our understanding of amphibian development and of the evolution of amphibian life cycles. J Morphol 233:15-29, 1997. © 1997 Wiley-Liss, Inc.