The zebrafish unplugged gene controls motor axon pathway selection.
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En route to their targets, motor axons encounter choice points at which they select their future path. Experimental studies predict that at each choice point specialized cells provide local guidance to pathfinding motor axons, however, the identity of these cells and their signals is unknown. Here, we identify the zebrafish unplugged gene as a key component for choice point navigation of pioneering motor axons. We show that in unplugged mutant embryos, motor neuron growth cones reach the choice point but make inappropriate pathway decisions. Analysis of chimeric embryos demonstrates that unplugged activity is produced by a selective group of mesodermal cells located adjacent to the choice point. As the first motor growth cones approach the choice point, these mesodermal cells migrate away, suggesting that unplugged activity influences growth cones by a contact-independent mechanism. These data suggest that unplugged defines a somite-derived signal that elicits differential guidance decisions in motor growth cones. (+info)
Fate and function of the ventral ectodermal ridge during mouse tail development.
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In the mouse embryo, the body axis continues to develop after gastrulation as a tail forms at the posterior end of the embryo. Little is known about what controls outgrowth and patterning of the tail, but it has been speculated that the ventral ectodermal ridge (VER), a morphologically distinct ectoderm on the ventral surface near the tip of the tail, is a source of signals that regulate tail development (Gruneberg, H. (1956). Nature 177, 787-788). We tested this hypothesis by ablating all or part of the VER and assessing the effects of such ablations on the development of tail explants cultured in vitro. The data showed that the VER produces signals necessary for somitogenesis in the tail and that the cells that produce these signals are localized in the middle and posterior region of the VER. Dye labeling experiments revealed that cells from these regions move anteriorly within the VER and eventually exit it, thereby colonizing the ventral surface ectoderm anterior to the VER. In situ hybridization analysis showed that the genes encoding the signaling molecules FGF17 and BMP2 are specifically expressed in the VER. Assays for gene expression in VER-ablated and control tails were performed to identify targets of VER signaling. The data showed that the VER is required for expression of the gene encoding the BMP antagonist noggin in the tail ventral mesoderm, leading us to speculate that one of the major functions of the VER in tail development is to regulate BMP activity. (+info)
The expression of the homeobox gene Msx1 reveals two populations of dermal progenitor cells originating from the somites.
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Experimental manipulation in birds has shown that trunk dermis has a double origin: dorsally, it derives from the somite dermomyotome, while ventrally, it is formed by the somatopleure. Taking advantage of an nlacZ reporter gene integrated into the mouse Msx1 locus (Msx1(nlacZ) allele), we detected segmental expression of the Msx1 gene in cells of the dorsal mesenchyme of the trunk between embryonic days 11 and 14. Replacing somites from a chick host embryo by murine Msx1(nlacZ )somites allowed us to demonstrate that these Msx1-(beta)-galactosidase positive cells are of somitic origin. We propose that these cells are dermal progenitor cells that migrate from the somites and subsequently contribute to the dorsalmost dermis. By analysing Msx1(nlacZ) expression in a Splotch mutant, we observed that migration of these cells does not depend on Pax3, in contrast to other migratory populations such as limb muscle progenitor cells and neural crest cells. Msx1 expression was never detected in cells overlying the dermomyotome, although these cells are also of somitic origin. Therefore, we propose that two somite-derived populations of dermis progenitor cells can be distinguished. Cells expressing the Msx1 gene would migrate from the somite and contribute to the dermis of the dorsalmost trunk region. A second population of cells would disaggregate from the somite and contribute to the dermis overlying the dermomyotome. This population never expresses Msx1. Msx1 expression was investigated in the context of the onset of dermis formation monitored by the Dermo1 gene expression. The gene is downregulated prior to the onset of dermis differentiation, suggesting a role for Msx1 in the control of this process. (+info)
Dorsoventral axis determination in the somite: a re-examination.
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We have repeated classic dorsoventral somite rotation experiments (Aoyama and Asamoto, 1988, Development 104, 15-28) and included dorsal and ventral gene expression markers for the somitogenic tissue types, myotome and sclerotome, respectively. While the histological results are consistent with those previously published, gene expression analysis indicates that cells previously thought to be 'sclerotome' no longer express Pax1 mRNA, a sclerotome marker. These results, together with recent quail-chick transplantation experiments indicating that even very late sclerotome tissue fragments are multipotential (Dockter and Ordahl, 1998, Development 125, 2113-2124), lead to the conclusion that sclerotome tissue remains phenotypically and morphogenetically plastic during early embryonic somitogenesis. Myotome precursor cells, by contrast, appear to be determined within hours after somite epithelization; a finding consistent with recent reports (Williams and Ordahl, 1997, Development 124, 4983-4997). Therefore, while these findings support a central conclusion of Aoyama and Asamoto, that axis determination begins to occur within hours after somite epithelialization, the identity of the responding tissues, myotome versus sclerotome, differs. A simple model proposed to reconcile these observations supports the general hypothesis that determinative aspects of early paraxial mesoderm growth and morphogenesis occur in early and late phases that are governed principally by the myotome and sclerotome, respectively. (+info)
Crustacean (malacostracan) Hox genes and the evolution of the arthropod trunk.
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Representatives of the Insecta and the Malacostraca (higher crustaceans) have highly derived body plans subdivided into several tagma, groups of segments united by a common function and/or morphology. The tagmatization of segments in the trunk, the part of the body between head and telson, in both lineages is thought to have evolved independently from ancestors with a distinct head but a homonomous, undifferentiated trunk. In the branchiopod crustacean, Artemia franciscana, the trunk Hox genes are expressed in broad overlapping domains suggesting a conserved ancestral state (Averof, M. and Akam, M. (1995) Nature 376, 420-423). In comparison, in insects, the Antennapedia-class genes of the homeotic clusters are more regionally deployed into distinct domains where they serve to control the morphology of the different trunk segments. Thus an originally Artemia-like pattern of homeotic gene expression has apparently been modified in the insect lineage associated with and perhaps facilitating the observed pattern of tagmatization. Since insects are the only arthropods with a derived trunk tagmosis tested to date, we examined the expression patterns of the Hox genes Antp, Ubx and abd-A in the malacostracan crustacean Porcellio scaber (Oniscidae, Isopoda). We found that, unlike the pattern seen in Artemia, these genes are expressed in well-defined discrete domains coinciding with tagmatic boundaries which are distinct from those of the insects. Our observations suggest that, during the independent tagmatization in insects and malacostracan crustaceans, the homologous 'trunk' genes evolved to perform different developmental functions. We also propose that, in each lineage, the changes in Hox gene expression pattern may have been important in trunk tagmatization. (+info)
Uncx4.1 is required for the formation of the pedicles and proximal ribs and acts upstream of Pax9.
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The expression of the homeobox gene Uncx4.1 in the somite is restricted to the caudal half of the newly formed somite and sclerotome. Here we show that mice with a targeted mutation of the Uncx4.1 gene exhibit defects in the axial skeleton and ribs. In the absence of Uncx4.1, pedicles of the neural arches and proximal ribs are not formed. In addition, dorsal root ganglia are disorganized. Histological and marker analysis revealed that Uncx4.1 is not necessary for somite segmentation. It is required to maintain the condensation of the caudal half-sclerotome, from which the missing skeletal elements are derived. The loss of proximal ribs in Pax1/Pax9 double mutants and the data presented here argue for a role of Uncx4.1 upstream of Pax9 in the caudolateral sclerotome. Our results further indicate that Uncx4.1 may be involved in the differential cell adhesion properties of the somite. (+info)
The paired homeobox gene Uncx4.1 specifies pedicles, transverse processes and proximal ribs of the vertebral column.
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The axial skeleton develops from the sclerotome, a mesenchymal cell mass derived from the ventral halves of the somites, segmentally repeated units located on either side of the neural tube. Cells from the medial part of the sclerotome form the axial perichondral tube, which gives rise to vertebral bodies and intervertebral discs; the lateral regions of the sclerotome will form the vertebral arches and ribs. Mesenchymal sclerotome cells condense and differentiate into chondrocytes to form a cartilaginous pre-skeleton that is later replaced by bone tissue. Uncx4.1 is a paired type homeodomain transcription factor expressed in a dynamic pattern in the somite and sclerotome. Here we show that mice homozygous for a targeted mutation of the Uncx4.1 gene die perinatally and exhibit severe malformations of the axial skeleton. Pedicles, transverse processes and proximal ribs, elements derived from the lateral sclerotome, are lacking along the entire length of the vertebral column. The mesenchymal anlagen for these elements are formed initially, but condensation and chondrogenesis do not occur. Hence, Uncx4.1 is required for the maintenance and differentiation of particular elements of the axial skeleton. (+info)
DNA dendrimers localize MyoD mRNA in presomitic tissues of the chick embryo.
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MyoD expression is thought to be induced in somites in response to factors released by surrounding tissues; however, reverse transcription-PCR and cell culture analyses indicate that myogenic cells are present in the embryo before somite formation. Fluorescently labeled DNA dendrimers were used to identify MyoD expressing cells in presomitic tissues in vivo. Subpopulations of MyoD positive cells were found in the segmental plate, epiblast, mesoderm, and hypoblast. Directly after laying, the epiblast of the two layered embryo contained approximately 20 MyoD positive cells. These results demonstrate that dendrimers are precise and sensitive reagents for localizing low levels of mRNA in tissue sections and whole embryos, and that cells with myogenic potential are present in the embryo before the initiation of gastrulation. (+info)