Before our rapidly increasing knowledge of gene interactions can be connected with the morphologic defects in mutant zebrafish, the normal course of skeletal development must be understood. Here, we describe the developmental morphology of the axial skeleton of zebrafish and review it in relation to the morphology of related species. The relative sequence of ossification in the skeleton is described. Two separate centers of development were found in the axial skeleton (Weberian apparatus and caudal fin) in contrast to tetrapods, which have a single anterior center. Slight variation was found in the overall relative timing of development. The extensive ichthyological literature on teleost anatomy and recent genetic data form the basis for our review and interpretation of homologies of various elements of the axial skeleton. Because homology forms the basis for all evolutionary comparisons, these data are critical for integration in evo-devo studies. Developmental Dynamics 228:337-357, 2003.

Detailed examples of how hierarchical assemblages of modules change over time are few. We found broadly conserved phylogenetic patterns in the directions of development within the median fins of fishes. From these, we identify four modules involved in their positioning and patterning. The evolutionary sequence of their hierarchical assembly and secondary dissociation is described. The changes in these modules during the evolution of fishes appear to be produced through dissociation, duplication and divergence, and co-option. Although the relationship between identified median fin modules and underlying mechanisms is unclear, Hox addresses may be correlated. Comparing homologous gene expression and function in various fishes may test these predictions.The earliest actinopterygians likely had dorsal and anal fins that were symmetrically positioned via a positioning module. The common patterning (differentiation) of skeletal elements within the dorsal and anal fins may have been set into motion by linkage to this positioning module. Frequent evolutionary changes in dorsal and anal fin position indicate a high level of dissociability of the positioning module from the patterning module. In contrast, the patterning of the dorsal and anal fins remains linked: In nearly all fishes, the endo- and exoskeletal elements of the two fins co-differentiate. In all fishes, the exoskeletal fin rays differentiate in the same directions as the endoskeletal supports, indicating complete developmental integration. In acanthopterygians, a new first dorsal fin module evolved via duplication and divergence. The median fins provide an example of how basic modularity is maintained over 400 million years of evolution.

BackgroundThe canals of the mechanosensory lateral line system are components of the dermatocranium, and demonstrate phenotypic variation in bony fishes. Widened lateral line canals evolved convergently in a limited number of families of teleost fishes and it had been hypothesized that they evolved from narrow canals via heterochrony and explore modularity in the lateral line system. Two species of cichlids with different canal phenotypes were used to test a hypothesis of heterochrony. Histological material prepared from ontogenetic series of Aulonocara stuartgranti (widened canals) and Tramitichromis sp. (narrow canals) was analyzed using ANCOVA to determine rates of increase in canal diameter and neuromast size (length, width) and to compare the timing of onset of critical stages in canal morphogenesis (enclosure, ossification).ResultsA faster rate of increase in canal diameter and neuromast width (but not length), and a delay in onset of canal morphogenesis were found in Aulonocara relative to Tramitichromis. However, rates of increase in canal diameter and neuromast size among canals, among canal portions and among canals segments reveal similar trends within both species.ConclusionThe evolution of widened lateral line canals is the result of dissociated heterochrony - acceleration in the rate of increase of both canal diameter and neuromast size, and delay in the onset of canal morphogenesis, in Aulonocara (widened canals) relative to Tramitichromis (narrow canals). Common rates of increase in canal diameter and neuromast size among canal portions in different dermatocranial bones and among canal segments reflect the absence of local heterochronies, and suggest modular integration among canals in each species. Thus, canal and neuromast morphology are more strongly influenced by their identities as features of the lateral line system than by the attributes of the dermatocranial bones in which the canals are found. Rate heterochrony manifested during the larval stage ensures that the widened canal phenotype, known to be associated with benthic prey detection in adult Aulonocara, is already present before feeding commences. Heterochrony can likely explain the convergent evolution of widened lateral line canals among diverse taxa. The lateral line system provides a valuable context for novel analyses of the relationship between developmental processes and the evolution of behaviorally and ecologically relevant phenotypes in fishes.

SummaryThe dermomyotome is a pool of progenitor cells on the surface of the myotome. In zebrafish, dermomyotome precursors (anterior border cells, ABCs) can be first identified in the anterior portion of recently formed somites. They must be prevented from undergoing terminal differentiation during segmentation, even while mesodermal cells around them respond to signaling cues and differentiate.T-box containing transcription factors regulate many aspects of mesoderm fate including segmentation and somite patterning. The fused somites (fss) gene is the zebrafish ortholog of tbx6. We demonstrate that in addition to its requirement for segmentation, fss/tbx6 is also required for the specification of ABCs and subsequently the central dermomyotome. The absence of Tbx6-dependent central dermomyotome cells in fss/tbx6 mutants is spatially coincident with a patterning defect in the myotome.Using transgenic fish with a heat-shock inducible tbx6 gene in the fss/tbx6 mutant background, we further demonstrate that ubiquitous fss/tbx6 expression has spatially distinct effects on recovery of the dermomyotome and segment boundaries, suggesting that the mechanism of Fss/Tbx6 action is distinct with respect to dermomyotome development and segmentation.We propose that Fss/Tbx6 is required for preventing myogenic differentiation of central dermomyotome precursors before and after segmentation and that central dermomyotome cells represent a genetically and functionally distinct subpopulation within the zebrafish dermomyotome.

In amniotes, BMP signaling from lateral plate and dorsal neural tube inhibits differentiation of muscle precursors in the dermomyotome. Here, we show that BMPs are expressed adjacent to the dermomyotome during and after segmentation in zebrafish. In addition, downstream BMP pathway members are expressed within the somite during dermomyotome development. We also show that zebrafish dermomyotome is responsive to BMP throughout its development. Ectopic overexpression of Bmp2b increases expression of the muscle precursor marker pax3, and changes the time course of myoD expression. At later stages, overexpression increases the number of Pax71 myogenic precursors, and delays muscle differentiation, as indicated by decreased numbers of MEF21 nuclei, decreased number of multi-nucleated muscle fibers, and an increased myotome angle. In addition, we show that while BMP overexpression is sufficient to delay myogenic differentiation, inhibition of BMP does not detectably affect this process, suggesting that other factors redundantly inhibit myogenic differentiation. Developmental Dynamics 239:806-817,

Cypriniformes (which includes the minnows, carps, loaches, algae-eaters, stone loaches, and suckers) is a morphologically diverse and incredibly speciose order of teleosts. It has been suggested that a number of evolutionary innovations, key to improved hearing and feeding, have played an important role in cypriniform fishes' success. One such innovation, the Weberian apparatus, is a novel assemblage of vertebral elements and modified ribs that relay and amplify sound pressure changes from the gas bladder to the inner ear. The Weberian apparatus unites Cypriniformes with other major orders into an extremely species-rich group of fishes, the Otophysi. Together, otophysan fishes comprise one of the largest groups of fishes in the world, as well as the majority of freshwater fishes. Here we present a detailed comparison of the Weberian apparatus in a number of cypriniform families using cleared and stained specimens. We present data regarding inter- and intrafamilial morphological variation within Cypriniformes. With few, but evolutionarily important, exceptions we find that diagnostic features of the Weberian apparatus characterize each family. Interspecific variation within each of the families Balitoridae, Gyrinocheilidae, and Catostomidae is only slight, whereas variation among subfamilies within Cyprinidae and Cobitidae is far more significant. This comparative study identifies a number of distinct morphologies, some of which appear highly correlated with ecological niche. For example, inhabiting swift-moving waters appears to be a key factor in the encapsulation of the anterior gas bladder in some cobitids, balitorids, and gobionin cyprinids.

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The relatively simple structural organization of the cranial lateral line system of bony fishes provides a valuable context in which to explore the ways in which variation in post-embryonic development results in functionally distinct phenotypes, thus providing a link between development, evolution, and behavior. Vital fluorescent staining, histology, and scanning electron microscopy were used to describe the distribution, morphology, and ontogeny of the canal and superficial neuromasts on the head of two Lake Malawi cichlids with contrasting lateral line canal phenotypes (Tramitichromis sp. [narrow-simple, well-ossified canals with small pores] and Aulonocara stuartgranti [widened, more weakly ossified canals with large pores]). This work showed that: 1) the patterning (number, distribution) of canal neuromasts, and the process of canal morphogenesis typical of bony fishes was the same in the two species, 2) two sub-populations of neuromasts (presumptive canal neuromasts and superficial neuromasts) are already distinguishable in small larvae and demonstrate distinctive ontogenetic trajectories in both species, 3) canal neuromasts differ with respect to ontogenetic trends in size and proportions between canals and between species, 4) the size, shape, configuration, physiological orientation, and overall rate of proliferation varies among the nine series of superficial neuromasts, which are found in both species, and 5) in Aulonocara, in particular, a consistent number of canal neuromasts accompanied by variability in the formation of canal pores during canal morphogenesis demonstrates independence of early and late phases of lateral line development. This work provides a new perspective on the contributions of post-embryonic phases of lateral line development and to the generation of distinct phenotypes in the lateral line system of bony fishes. J. Morphol. 277:1273-1291, 2016. © 2016 Wiley Periodicals, Inc.